w \W. A SURVEY or THE coancomus PYRENOC'ARPCBUS ucams or THE GREAT LAKES REGION Thesis for the Degree of M. S. MICHIGAN STATE UNIVERSITY Richard Clinton Harris I966 LIB RA R 1’ Michigan Stab: University 1 Has)! ABSTRACT A SURVEY OF THE CORTICOLOUS PYRENOCARPOUS LICHENS OF THE GREAT LAKES REGION by Richard Clinton Harris This survey of the corticolous pyrenocarpous lichens in the Great Lakes region treats nine genera and 23 species. The study is based on field work and examination of material from the Farlow Herbarium, Michigan State University, Smithsonian Institution, Univer- sity of Michigan, University of Wisconsin and the personal herbarium of J. W. Thomson. The morphology and ecology of these lichens is discussed briefly. Spores are illustrated for all species treated. Molestia and Plagio- 22523 are described as genera new to science. Arthopyrenia gyisguiliae, A. thomsonii, A. willgyana, Pyrenula agawae and P. neglecta are described as species new to science. Arthopyrenia myricae (Nyl.) Zahlbr., Leptorhaphis atomaria (Ach.) Szatala, L. parameca (Massal.) Koerb., and Polyblastiopsis meridionalis Zahlbr. are reported as new to North America. Microthelia wallrothii (Hepp) Rehm is reported as new to the Great Lakes Region. Pyrenula nitida (Weig.) Ach. and Arthopyrenia alba (Schrad.) Zahlbr. are excluded from the flora. A SURVEY OF THE CORTICOLOUS PYRENOCARPOUS LICHENS OF THE GREAT LAKES REGION BY Richard Clinton Harris A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE Department of Botany and Plant Pathology 1966 CONTENTS List of appendices ---------------------------------------- ~v List of plates ------------------ - ......................... vi Introduction-- ------ ~ ---------- - —————————— - ............... -1 Morphology Methods ------------------------------------------------ 3 Thallu8---- --------------------- Dun-cucu- ............... ‘A Aacocarp---- ----- --- ------- - ------ ----¢- ————— ... —————————— 7 Paraphyses and paraphysoids-------~ ------------------ -12 Asci---—---------—----------- .......... - ..... - ...... -_13 Spores ------ ------ -------------- - ............. - ...... -15 Pycnidia-----~--- ----------------------------------- --16 ECOlOgyd-o-------------o ---------- C--- CCCCCC - aaaaa u- ........ 17 Taxonomy Family classification—------------~ ---------------- ---23 Key to species---------—-----——------—- ............... 25 Pleosporaceae—-~-----~----~-—---------—--- ...... ------28 ArthOpyrenia---------------- .................. ---28 A. finkiie-~——---- ------- - ——————— --—------.-29 A. myricae----~- ----- -----—-----e-----------31 A. punctiformis-------—--—--------- ......... 32 A. quinqueseptata----------------—----------34 A. quisquiliae-----------o---—-~ ............ 36 A. sphaeroides------o--- .................... 37 A. thomsonii~------—-------------— ...... ----39 A. willgyana-o------ ------- - ................ 39 ii Dermatina ........................................ 42 Do Pjrenocarpa ------------ qa-u..- --------- ---_I+2 Leptorhaphis-o- ---------------------------- - ..... hh LO atomaria------- ........ -D-O-Db-O-o uuuuuuu #5 Le contorta--- --------- -—------ ............. #6 Lo ePidermidiB‘------ ------ -u-------- ....... 1+? La jarameca------~----------------o-o---~--—50 Microthelia-------o-----------—--------- ..... ----51 Me micula~- -------- - ----- ------- ..... -OOQ—a-SZ MO nicula Var. maladr11oculata“--------—-O--—54 MC 'allrOthii’“-"--"'“C".------------~------—55 M0198tia‘--"“’--------‘-"--C-HI'C'IIN-ODCOO-CDCO-nncoa56 M. leucgplaca-----—--—---—--- ------- - ..... --57 Melanomma up. confused with Molestia--------6O Po laCtCa------’ --------------- can-Io--- ..... 65 P. meridionalis-- ------ --—- ----- - ----------- 66 Pernulaccae----------——--——--------- ............... --67 Plagiocagpa-------- ....... ¢--- ---------- ---------67 P. hyalospora -------------------- -—- ..... ---69 gzgenula----------- ----------------- ---—---------72 Po ngglect8--“--------*-------~-~--~-~~-----72 PO agwae‘—--"----‘- ----------------- d-Q—O-u7s Tryphetheliaceae-----~-- ------ - ------- - ............... 76 Tryphethelium----------------------— ............ _76 T. virens-------—-------- ------ -------- ..... 76 iii Summary ---------------------------------------------------- 78 Literature cited ........................................... 79 iv List of Appendices Appendix I. Basic ecological data ------------------------- 82 Appendix II. Pyrenolichen substrates in the Great Lakes region ---------- - --------- - ............ 92 List of plates Plate 1. Map of Great Lakes region showing sites of ecological sample stands—---------------- --------- --93 2. Spores of Arthopyrenia-------------------- ---------- 95 3. Spores of Leptorhaphis, Microthelia and Polyblastiopsis ------- --------------- ----------- ---_97 h. Spores of Dermatina, Melanomma, Molestia, Plagiocarpa, Pyrenula and Tryphethelium-- ------- ----99 vi INTRODUCTION This study was designed primarily as a taxonomic survey of the corticolous pyrenocarpous lichens in the Great Lakes region. Very little work has been done in this group in North America. There have been no previous systematic collections made of these lichens made in the Great Lakes region which centers on Michigan including the area adjacent to the Great Lakes as shown in Pl. 1. Bruce Fink collected on the western edge of the area in Iowa and Minnesota, but he apparently had reached no final treatment of this group at the time of his death. Specimens were tentatively named pending a final decision and the same species was often found under more than one name. Thus in order to clarify Fink's species concepts and because of their historical importance many of his collections have been included in this study although they were not collected in the Great Lakes region proper. Several new species have been included in order to place them on record since they probably will be collected in this region eventually. Specimens seen from outside of the Great Lakes region are cited only for taxa of special interest. The pyrenocarpous lichens seem to be on the boundaries of lichen- ization and are worthy of study for this reason alone. In view of the difficulty of determining lichenization it was decided to collect all pyrenocarpous fungi in the field and then after preliminary examina— tion in the laboratory to treat further only such material as could be determined to belong to a genus or species which traditionally was considered to be a lichen whether or not an obvious algae containing thallus was present. A secondary purpose of this study was to become acquainted with the problems of morph010gy and morphoIOgical terminology in the pyrenocarpous lichens preparatory to monographic work in the group. For this reason the discussion of morphology is somewhat more exten- sive than it otherwise might be. The lack of knowledge concerning the pyrenocarpous lichens is especially great in regard to their ecology. On account of this the field work also included a brief ecological study designed to provide some basic ecological data about their abundance and relations with the‘substrate. I would like to thank the curators of the following herbaria for generous loans of material for this study: Farlow Barbarian, Harvard Univ. (FE), Michigan State Univ. (MSG), Smithsonian Institution (Us), Univ. of Michigan (MICE), Univ. of Wisconsin (WIS), and the personal herbarium of J. W. Thomson (Thomson). I would especially like to thank H. A. Imshaug for his advice and guidance during the course of the study. MORPHOLOGY Methods External morphology was studied under a dissecting microscope. Measurements were made with an eyepiece micrometer. For microscOpic studies the material was moistened with water and freehand sections were cut with a sharp razorblade. Freezing microtome sections were made for some collections during the earlier stages of the study. Although microtome sections are generally superior, especially for details of ascocarp structure, the large amount of time and difficul- ty involved precluded their routine use. Thallus sections were mounted in 10-20% potassium hydroxide (KOH) and examined for the presence of algae. If algae were not immediate- ly obvious, additional sections were mounted in lactOphenol-cotton blue, warmed briefly and examined for algae. The cotton blue stains the cytoplasm of the algae but not the bark cells of the substrate making the presence and abundance of the algae easily observable. Ascocarp sections were routinely mounted in Lugol's iodine (IKI, iodine lgm, potassium iodide lgm, water lOOgm). Occasionally there are color reactions with the gelatinous material in the ascocarp nucleus. In general, however, the contents of the asci are colored bright orange-red, making them easier to locate. The cytOplasm of the paraphyses or paraphysoids is often colored pale yellow. After it was observed that dilute KGB changed the appearance of the asci, 20% KOB was run in under the edge of the coverslip of the IKI mount and its effects on the ascus were observed. Several biological stains were tried in an attempt to investi- gate ascus structure. None were particularly successful. Congo red, 3 recommended by Richardson & Morgan~Jones (196%) is a good cytOplasmic stain but does not seem sufficiently superior to the more easily handled IKI to warrant its routine use. Janus green shows promise for differential staining of parts of the ascus wall but I have not been able as yet to devise a consistently satisfactory procedure for its use. R. A. Shoemaker (1964) has suggested that blue writing ink is very good for staining certain ascus structures. I have not as yet tried it. Paraphyses and paraphysoids were examined in 10-20% KOH. This dissolves most of the gelatinous material around then and allows their septation and branching to be studied. Potassium hydroxide often causes considerable swelling and if measurements were made in KOH,this fact should be indicated. Ascus and spore measurements were made in both IKI and KOH. There is little, if any, difference in overall ascus or spore size in the two reagents. Critical morphological features were recorded for the majority of the specimens by drawing them with the aid of a Wild drawing tube. Thallus The thallus in all of the pyrenocarpous lichens examined so far is of the endophloeodal type. The fungus hyphse and algae, if pres- ent, occur beneath the upper layers of the bark, penetrating into the bark to varying depths. The alga, in all cases where there is a clear algal layer, is Trentepohlia. The amount of algae and hyphae in the thallus varies considerably from species to species. In many cases what appears to be a thallus is in reality an area of roughened, loosened and modified bark cells with relatively few hyphae and often 1, no algae. it the Opposite extreme the external layer of the thallus is an almost cortex-like layer of bark cells consolidated with very abundant hyphae. The bark and the hyphae may almost lose their iden- tity. In this type a well-develOped algal layer is always present. There seems to be at least a partial series connecting these two end- points. The loose bark, no algal layer type is found in Polyblasti- cpsis, Eeptorhaphis and some Species of Arthopyrenia. Dermatina pyrenocarpa has a thallus in which there are scattered clumps of algae in the loose bark-hyphal layer. This species seems to be the only one intermediate in amount of algae. In all other cases there is either a clearly defined algal layer, at least when young, or no algae at all. Many ArthOpyrenia species, Plagiocarpa and Molestia have a good algal layer underneath a roughened bark exterior. The most ad- vanced type, in which the thallus appears shiny and smooth due to the cortex-like upper layer, occurs in gyrenula neglects and Tryphe- thelium sirens. Both of which, it may be noted, are ascohymenial. Almost half of the species in the Great Lakes region lack an obvious algal layer. Algae are apparently entirely lacking or only a few scattered cells are present on the surface of the bark. These scattered cells never seem to be Trentepohlia. I do not know whether these algae are bark epiphytes or are associated with the fungus. If these species are indeed lichens, it is clear that they are on the borderline of lichenisation. Lichenologists and mycologists, on the whole, have paid very little attention to genera which have been considered traditionally to belong in the others domain. Genera and families have been circumscribed to include only fungi or only lichens on the basis that they are two fundamentally different types of organisms. Lichens are now generally considered to be fungi which 5 which differ from other fungi only in the source of their nutrition. Gradually less and less emphasis is being placed on the source of nutrition, i.e., saprOphytic or parasitic and if parasitic whether on higher plants or algae. Distinctions on this basis are being replaced by ones based on ascocarp, ascus and spore morphology. Lichenologists, in general I think, have realised the relative unimportance of nutri- tion in taxonomy earlier than nycologists. Bruce Fink (1913, pp. 106- 107) over fifty years ago stated what he considered the ideal solu- tion to this problem. "If we are ever to know the relationships of algicolous to non-algicolous Ascomycetes, lines of cleavage in study must cease to be determined by the food habits of these plants. Forms that appear to be closely related must be studied together, whether all are algicolous, all non- algicolous, or part of them one and part the other." However, it is only quite recently that mycologists have begun to re- consider some of the genera traditionally passed over as lichens. Butler (1940) suggested that the genus Buellia should include lichen- ised fungi, lichen parasites and saprOphytes. The first comprehen- sive work to take up this suggestion is that of Mueller and von irx (1962). H. Riedl (1963, p. 26k), a mycologist experimenting with the application of modern mycologioal taxonomic principles to a group of pyrenocarpous lichens, says in a discussion of Arthopyrenia, “Vom Standpunkt der Flechtenorganisation aus, ist Artho- enia besonders primitiv nu werten, es finden sich bei ihr saemtliche Uebergaenge von nicht lichenisierten ueber seitweilig lichenisierte su obligat lichenisierten irten. Aber such in letzterem Fall handelt es sich wohl nur um eine Form von obligates Parasitismus." Riedl also points out that if parasitism on algae is a generic or even a specific criterion, then facultative lichen fungi would have to recieve different names depending on whether algae were present or absent when collected. This is, of course, nomenclaturally absurd. 6 Even if some of the pyrenocarpous genera can be shown to be complete- ly non-lichenized, they probably will continue to be treated prima- rily by lichcnologists due to the inertia of tradition. These are the reasons why many species which are described as not having asso- ciated algae are treated here in a work on lichens. Ascocarp Within the class Ascomycetes two fundamentally different types of ascocarp develOpment have been distinguished. Nannfeldt (1932) introduced the terms Ascoloculares and Ascohymeniales for the two types and although they have never formally been given taxonomic rank, they make euphonious descriptive adjectives. The two basic types will be referred to as ascolocular and ascohymenial. The ascocarp in the ascohymenial type possesses a wall derived from the basal cells of the archicarp, which is the name given to the ascogonium and the cells immediately surrounding it resulting from the stimulus of sexual fusion. A pyrenomycete ascocarp with such a wall around it is preperly called a perithecium. The peri- thecial wall is probably best called the exciple or excipulum. Exciple has been used for both apothecial and perithecial walls by lichenologists, although it is often restricted to apothecia by mycologists. In view of the presumed phylogenetic relationship of ascohymenial discomycetes and pyrenomycetes and in view of the iden- tical origin and function of this tissue in both, exciple seems to be the best term for the perithecial wall. The sterile elements of the hymsniun arise from the basal ascogenous layer before the asci and grow upwards. These paraphyses are usually unbranched, not anasta- nosed and free at their upper ends. The peritheciun Opens by an 7 ostiole which is schizogenous in origin, resulting from differential growth of the exciple. The terms perithecium, paraphyses and ostiole are usually restricted to ascohymenial ascomycetes by contemporary mycologists. The ascolocular ascocarp begins as a small patch of vegetative cells which multiply and become carbonized to form an area of stro- matic tissue. There is apparently no sexual fusion stimulus invol- ved in the formation of this tissue. Sexual fusion, if it occurs, takes place within this tissue after it has develoPed. In the sim- plest types the asci which develop within the stroma are randomly scattered and each dissolves its own chamber in the stroma as it expands. The probable next stage has the asci aggregated into a group or several groups in the stroma. The asci may again dissolve chambers or they may be surrounded by the compressed remains of the original stroma tissue. This tissue has been called either para- physoid threads (Singer 1963) or interthecial tissue (Luttrell 1965). The most advanced type has a distinct paraphysis—like tissue which grows downward from the tap of the stroma before the asci develop. This type of tissue has been called paraphyseid hyphae (Singer) or pseudoparaphyses (Luttrell). They are attached at the top, branched and anastamosed. The ascolocular type of ascocarp with a single chamber is a pseudothecium. Its Opening is simply called a pore. Singer (1963) suggests that if the origin of the sterile elements is not definitely known the general term paraphysoid should be used. Since I have made no develoPmental studies, I will use the term para- physoid. The major feature of the ascolocular type is that the ascocarp is a modified stroma in which the asci develop without any seperating 8 wall derived from the archicarp. If a perithecium is embedded in a stroma, the fertile tissue is separated from the stromatic tissue by a distinct wall derived from the archicarp. Stromata occur in ascohymenial pyrenolichens as well as forming the basis of the ascocarp in the ascolocular pyrenolichens. A stroma may be defined generally as a cushion—like mass of fungus cells and host tissue, in or on which fructifications develOp. Some workers, including Johnson (l9h0) prefer to call a stromatic tissue containing numerous cells from the substrate a pseudostroma. In the asco- hymenial species the stress is an accessory structure surrounding a peritheciua or group of perithecia. It tends to contain bark cells and is therefore in most cases a pseudostroma in the sense of Johnson. In Tryphethelium virens the raised area around groups of perithecia is almost all bark mixed with relatively few hyphae. It may or may not be carbonized. In gyrenula and Plagiocarpa_the stroma is reduced to a fairly thin carbonaceous wall-like layer surrounding a single perithecium with various degrees of completeness or forming a shield at the tip of the perithecium. A general term for this stromatic layer is involucrellua (Swinscow 1962). The exciple often separates from the involucrellum. The involucrellum may contain crystals of unknown composition. The involucrellum in some cases is restricted to a shield around the ostiole and should be referred to by the nycological term, clypeus. If the involucrellum is lacking at the base of the peritheciua, it may be called a dimidiate involucrellum. The majority of the ascolocular species have only a single chamber in the stroma which does not usually contain bark cells. Such pseudothecia are found in érthOpyrenia, Leptorhaphis, golyblastiOpsis, Molestia and Micrgthelia. The ascolocular stroma in Dermatina and 9 related genera usually contain c. 2-10 distinct chambers separated from each other by thin layers of stromatic tissue. Each chamber has its own exit pore. Since the pseudothecium is a modified stroma, the nomenclature used for the wall and accessory structures of an ascohymenial asco- carp cannot be used. It also seems inadvisable to create terms for ascolocular structures by adding -oid to ascohymenial terms. The layer of carbonized strosatic tissue surrounding the fertile tissue is not a wall in the sense that the perithecial wall is, but since there is no other satisfactory term, I will use pseudothecial wall. The term excipuloid layer has been used, but it seems to me that when, as is often the case, the outer part of the stroma is darker than the inner, there is too much temptation to use the term involucrelloid layer also. This is contrary to the fact that there is, in reality, only a single tissue involved. Also the stromatic origin of the involucrellua as opposed to the archicarpal origin of the exciple might tend to be forgoten. My own studies of the ascocarp, especially in ascolocular species, have been somewhat superficial. In monographic work more use would have to be made of pseudothecial wall characters. Mycologists have based species and even genera on its cellular characteristics (Scheinpflug 1958). Virtually the only character which has been used by lichenologists is whether the "wall" is entire or lacking below (dimidiate). In both ascolocular and ascohymenial types this can only apply to stromatic tissues. This character must be used with great care. There seems to be variation between dimidiate and entire forms within a single species. Therefore the character is probably best treated as a quantitative rather than a qualitative character. 10 The pseudothecia of several genera may be surrounded by a dark circular or elliptical stain. Keissler (1938) calls this the "hof". It is composed of dark brown hyphae extending outward from the pseudo- thecial wall and may possibly serve to anchor the pseudothecium to the substrate. It is most highly develOped on Betula whose bark flakes off very readily. This structure is not found in any species possessing a well-developed algal layer. The fertile tissue of the ascocarp taken as a unit is often termed the nucleus, especially when being considered on a more-or- less lacroscOpic level. The term is very useful in discussing color reactions since the reacting substance has not been definitely local- ised, although it seems to me to be in the gelatinous material euro rounding the paraphyses or paraphysoids. It may be a polysaccharide in view of the blue-green or red-orange reaction with iodine. Iodine reactions are found in both ascolocular and ascohymenial species. These reactions are often used to distinguish species or occasionally varieties. Iodine reactions must be interpreted with great care since the concentration of iodine in the IKI solution used to test the ascocarp section affects the color produced. Also the IKI solution tends to deteriorate fairly rapidly with age and should be freshly made for any critical work. For example, in Plagiocarpa hyalospora too low an iodine concentration results in no reaction. A moderate concentration ” gives a strong blue-green color in the nucleus which may be changed to red-orange by increasing the con- centration of iodine. By raising and lowering the concentration the cycle of colors can be run thru several times before it remains per- manently redoorange. It is apparently a general rule that a high enough concentration of iodine will finally shift a blue-green color 11 to red-orange, indicating perhaps that there is only a single sub- stance reacting. In Leptorhaphis atomaria the orange color appears at such low iodine concentrations that the blue-green color is not seen or is visible only fleetingly. If the IKI solution is not al- lowed to become too old these reactions seen quite reliable and are usually correlated with morphological characters. The nucleus may contain oily substances in large or small amounts and appear as draplets in a K08 mount. The presence or absence of oil in the nucleus seems to be a consistent character at the species level. Paraphyses and Paraphysoids is mentioned in the discussion of the ascocarp the sterile ele- sents among the asci have different origins and characteristics. The ascolocular type, paraphysoids, are attached at tap and bottom without any free tips. They are usually clearly branched and much anastamosed. They are always septate. The amount of septation and branching varies greatly. it one extreme, found in heptorhaphis and Polyblastiopsis especially, the paraphysoids are more septate with short cells so that they appear almost parenchynatous. The other extreme is repre- sented by some species of £3£EQBIE°nia where septation and branch- ing are almost non-existent, so that these paraphysoids are diffi- cult to distinguish from true paraphyses. In such cases the ascus type may become the sole criterion of the ascolocular nature of these species. In our species of the ascohymenial genus gzgenula the paraphyses are unbranched, apparently non-septate and free at the tips. Plagio- carpa has slightly branched and septate paraphyses. In Tryphethelius 12 the paraphyses pose some difficulty since they are anastamosed into a broad meshed net. However, they are not much if at all scptate. This broad paraphysis reticulum has not been found in any genera known to have ascolocular develOpment or bitunicate asci. This paraphysis type seems to be an independent development in ascohymenial pyreno- mycete evolution. I have also observed anastamosed paraphyses in discomycete lichens, such as Pertusaria, but no one has ever suggest- ed that their develOpment is ascolocular. isci As far as I am aware no studies have been made to definitely establish ascolocular or ascohymenial develOpment in any of the pyre— nocarpous lichens. This information is merely inferred from the presence of characters which have been shown in other well studied cases to be associated with one type or the other. One is the pre- sence of paraphyses as apposed to paraphysoids. The other major cha- racteristic is that the structure of the ascus can be divided into two types which generally correlate with the type of ascocarp deve- lopment. The walls of the asci in both types consist of two layers, excascus and endoascus. In the ascohymenial type the two layers are permanently fused together. The spores are liberated in most cases, by means of a complicated pore mechanism at the ascus tip. This type is called the unitunicate ascus. In the ascolocular type the two. wall layers are not fused. The exoascus is thin and relatively in- elastic. The endoascus is usually thick and elastic. When the spores are to be liberated the exoascus splits or an apical exoascus cap is pushed off and then the exoascus expands becoming greatly elongated. The spores are then expelled from the stretched and extended endoascus. 13 The pore mechanism at the tip of the ascus is less complex. This type is called the bitunicate ascus. The bitunicate ascus has been demonstrated and illustrated for at least one species in most of the genera treated in this paper as ascolocular (Richardson & Morgan-Jones 1964). I have also observed bitunicate asci in many of the Great Lakes species, especially éfiiflfl‘ pyrenia and Polyblastiopsis. A second phenomenon has been noted which is apparently mainly a feature of bitunicate asci. Unfortunately, it was noticed only re- cently so that consistent detailed studies have not been made for very many species. When an ascus is treated with dilute KOH the wall at the tip increases considerably in thickness. Sometimes an ascus tip may become 4-5 times thicker. This thickening has not been ob- served is all species studied but it may occur so slowly that I simply have not watched long enough. .This phenomenon seems associated with another common characteristic of bitunicate asci, the dimple of Richardson and Morgan-Jones (l96h) or the chambre sous-apicale of Magne.(l9#6). This structure is a narrowing of the ascus cavity just below the tip to form what looks like an indentation in the ascus tip. Th0 dimple may be almost unnoticable, broad and shallow, or small and thisble-shaped, or larger and subglobose. The shape and size of the dimple as seen in asci mounted in IKI or Congo red is a useful cha- racter. The shape in KOH is also useful but needs more care in interpretation, since the shape is often changed in KOH. Also in K03 dimples become evident which were not evident or only indistinct- ly so in IKI. The dimple in XOR is usually a short thimble-shaped tube projecting somewhat into the ascus cavity. It should be noted that the exoascus is apparently unchanged by KOH. The changes in IR the ascus tip are due to an eXpansion of the exoascus tip downward into the ascus cavity. This expansion, occurring a short distance back from the tip, could narrow the ascus cavity and emphasize or even create a dimple. Occasionally the endoascus swells for a con- siderable distance back from the tip and a long narrow canal is formed. It is possible that this expanding layer is responsible for the mechanical splitting of the exoascus or pushing off an exoascus cap in the process of spore liberation. Its reactions in KOH are almost certainly abnormal. Spores The classification of the pyrenocarpous lichens is based pri« marily on differences in spore septation and color. No non-septate spores are known in the temperate—boreal corticolous pyrenolichens. The ascolocular genera have a wide variety of shape and septation but all lach.any endospore thickening. The spore lumina are, there- fore, cylindrical. They are transversally septate or muriform, colorless or brown. A few species passes a colorless outer layer which expands in K03 and eventually dissolves. It may be gelatinous in nature. Its presence or absence seems to be a reliable character at the species level. The ascohymenial genera in the Great Lakes region possess a thickened endospore. It is not well-develOped in Plagiocarpa but it is highly developed in gzgenula and Tryphethelium. Thickening of the endospore results in the lumina becoming lenticular, ellipsoidal or globose instead of cylindrical. The position of the lumina in relation to the outer spore wall is important. All three of the genera mentioned have transversally septate spores. Pyrenula spores 15 are 3-septate and brown, Plagiocarpa 3-septate and colorless, Tryphe- thelium 7-9-septate and colorless. Pycnidia The structures which I am treating as the imperfect states of the pyrenolichens are considered by some to be spermatia. The fusion of the spores produced with a trichogyne has never been demonstrated and until such information is available it seems best to treat them as asexual reproductive imperfect states. The type of perfect state recognized for the pyrenolichens is the pycnidial type. Pycnidia are quite common scattered among the ascocarps. They are generally similar in appearance to the ascocarps but usually much smaller, c. 1mm in diameter. The pycnidia known for all the ascolocular species and for Pygenula are simple one-chambered pycnidia. In Tryphethelium virens the pycnidia are labyrinthiform. The conidia produced in these pycnidia are normally borne on relatively short unbranched conidiOphores. Conidia in the pyreno- lichens have been divided into two kinds on the basis of size: macro- conidia and sicroconidia. The macroconidia are often pigmented and septate while sicroconidia are usually hyaline and simple. The con- idia in ascolocular species tend to be short rods, while in the ascohymenial species they tend to be longer, often filiform and curved. More attention should be paid to conidia since they may reveal fundamental groupings in the pyrenolichens. l6 ECOLOGY In view of having only a short time available in the field and in view of the taxonomic difficulties in attempting any detailed ecological studies of corticolous lichens, it was decided to keep the ecological study as simple as possible, especially since not even the most basic data have been collected previously for any corticolous pyrenocarpous lichens. In this study ten different forest stands were chosen for their interest or convenience. A tree was picked in what seemed on inspection to be a fairly homogeneous area. The next hundred trees over 10 cm in diameter at breast height away from the starting point were sampled moving in a single direc- tion to avoid repetition. The species and diameter were recorded for each tree. All pyrenocarpous lichens were collected. The height above ground, the side of the tree and a rough approximation of the area covered were recorded for each collection. The data are in no way rigorous and provide only general indications of the abundance, relations to the substrate and distribution of this little known group of lichens. Summations of the basic data for each stand are to be found in Appendix I. The stands have been roughly character— ized according to the system of Curtis (1959). The location of the sample stands is shown in Pl. 1. The percentage of trees with pyrenolichens (Table I) normally seems to range from 4 to 23. The Newaygo stand (7) which was ex- tremely dry (Opuntia nearby) had no pyrenolichens at all. The Pine Grove stand (5) had a very high pyrenolichen cover but it was almost all a single species, Polyblastiopsis’fallaciosa, which coated all of 17 sp the young maples dominating the stand. The number of pyrenolichen species ranged from 0 to S; the number of tree species from 6 to 1}. Table l. Summation of the number of pyrenolichen and tree species and the percentage of the trees with pyrenolichens. Stand # lichen species # tree species % trees with lichens 1 2 7 6 2 2 6 1+ 3 5 10 17 lo 2 7 7 5 1+ 8 5) 6 5 13 15 7 o 8 o 8 4 10 9 9 h 11 2) 10 3 11 16 If the number of tree species is graphed against the number of lichen species for each stand, there seems to be a tendency for both to increase tagether. l3 6 12 f. ll 10 9 spiZizs 10 8 3 9 8 7 5 7 4.1 6 2 O l 2 3 h 5 # pyrenolichen species Likewise the percentage of trees with pyrenolichens increases with an increase in the number of pyrenolichen species. 18 50 5 45 40 35 % trees 30 with 25 9 pyreno- 20 10 5 lichens 15 6 10 8 “1 5 2 o 1 2 3 h 5*1 # pyrenolichen species If the stands are arranged in a north to south order and the percent occurrence of each species is listed for the stands,it seems evident that there are two groups; one occurring south of Clare, Michigan and the other north of Clare. Of the three species which occur on both sides of the line, it is interesting to note that hicrothelia micula represents a species group and I have sus- picions that there may be more than one species included in Poly- blastiopsis fallaciosa. Table 2. Species t South Stand North } 9 10 8 7 6 5 h 2 3 l ArthOpyrenia sphaeroides E l A. finkii ; l A. willeyana 17 ll 5 Tryphethelium virens 2 2 PolyblastiOpsis fallaciosa' 13 3 l 2 #9 l 2 Microthelia micula . 5 l l 35 h 5 Pyrenula neglects l Molestia leucoplaca l 1 6 “ Leptorhaphis epidermidis 6 L. atomaria 1 Pyrenula agawae J 1 There are faint indications that the occurrence of the pyreno- lichens is related to the age of the tree which it grows on. In general it can be seen that the percentage of trees with pyreno- lichens increases with the percentage of trees in the 20-39 cm. diameter class and decreases with the increase of trees under 20cm. or over kOcn. 55 75 50 70 Outside scale: “5 65 fi'trees with “0 60 lichens 35 55 30 50 Inside scale: 25 “5 % trees 20-39 20 #0 cm. in diam. 15 35 10 30 5 25 O 20 12345678910 Stand There are two exceptions to the tendency for correlation of tree diameter and occurrence of pyrenolichens as shown in the graph above. The very dry Newaygo stand (7) had no lichens at all. The Pine Grove stand (5) was dominated by PolyblastiOpsis fallaciosa which seems to have a preference for the conditions on young gear saccharum but older Quercus alba as shown in Table 3. Table 2. Diameter in cm. 10-19 20-29 30,39 40- %jhcer saccharum ”I available 45 32 10 11 %.Acer saccharum _ with Polyblastiopsis 66 52 2 O %~Quercus alba available 20 26 2“ 3O % Quercus alba with Polyblastiopsis O O A“ 66 20 The only other species with enough collections to show any trend in preferring substrate trees with a certain diameter is Arthopyrenia willeyana which apparently has a preference for older trees. Since there were not enough collections from any one tree species, all the species which A. willeyana occurred on have been combined in a single table (Table h). Table h Diameter in cm. 10-19 20-29 30-39 to- % trees available 2#.5 22 20 5 trees with 0 2k 22 i. willeyana The final ecolOgical factor which may be worthy of comment is the tendency for species with well-deveIOped algal layers to be able to live on a greater variety of sustrates (Table 5). The data are compiled from herbarium material and my own collections. Table i Species ArthOpyrenia willeyana Plagiocarps hyalospora Pyrenula neglecta Molestia leucOplaca Arthopyreaia sphaercides Dermatina pyrenocarpa ArthOpyrenia finkii i. quinqueseptata A. quisquiliae Tryphethelium virens Pyrenula agawae Average Polyblastiopsis fallaciosa ArthOpyrenia punctiformis Polyblastiopsis meridionalis Laptorhaphis parameca L. atomaria L. contorta L. epidermidis Algae good layer layer scattered good layer layer 01 good layer layer absent N n M M u I! 21 # substrate trees 9, on lignum also 5, on lignum also , on 5 outside G.L. HHHNNtm uwwwmwuwtr O k Polyblastiopsis lactea absent 1 Melanomma sp. " 1 Hicrothelia wallrothii " l Arthopyrenia nyricae " 1 Average 1.8 It would appear that on the average those pyrenolichens with a good algal layer occur on twice as many tree species as those without algae e A table of the species of tree which the various pyrenolichens occur on will be found in Appendix II. 22 TAXONOMY Family Classification The delimitation of families in the pyrenocarpous lichens is at present in a state of chaos due to the relatively recent upheaval caused by the discovery of the two fundamental types of ascocarp development. According to Zahlbruckner (1926) the genera occurring in the Great Lakes would be placed in three families. Tryphethelium is our only representative of the Tryphetheliaceae Zahlbr. It can be char- acterized as possessing perithecia embedded in a pseudostroma. Mycoporaceae Zahlbr.. having more than one chamber in the ascocarp, is also known from the Great Lakes by a single genus, Dermatina. All of the remaining genera were included in the Pyrenulaceae Zahlbr. Of these Arthopyrenia, Leptorhaphis, Polyblastiopsis and Microthelia have been shown to possess bitunicate asci (Richardson & Morgan- Jones 196%) and are, therefore, presumably ascolocular. gyrenula, however, is ascohynenial. Since it is the type genus of the Pyrenulaceae, the ascolocular genera must be excluded from the family. gygenula and Plagiocarpa are the two genera of the Pyrenulaceae occurring in the Great Lakes region. Riedl (1962) has united the Mycoporaceae, which is clearly ascolocular, with the other ascoa locular genera removed from the Pyrenulaceae. This seems to me to be taxonomically correct but there is a nomenclatural problem as to what the correct name for this family is. This group seems to be closely related to genera in the traditionally non—lichen family Plecsporaceae Lindau. Many of the species treated in this paper are at best facultative lichens. Thus it seems best not to make any 23 family distinction based on lichenization. The Pleosporaceae, dating from 1897, is clearly the oldest 'available name for the family. It should be noted here that I am not reccomending that all lichen fam- ilies should be united with non-lichen families. It is clearly not warranted in most cases, but in this case where the relationships are clear and where the boundaries of lichenization are tenuous, uniting the two groups seems justifiable. If one wishes to maintain a separate lichen family, the correct name would seem to be the lanthOpyreniaceae Zahlbr. (Engler & Prantl, Nat. Pflfam. ed. 2. 8: 91. 1926). Riedl (1962) uses as the oldest name Hycoporaceae Zahlbr. (1903) but the generic name fichporum Plotow in Koerb. (Grundriss der Cryptogaa. 199. 1848) is a later homonys of MchROrum Meyer (Nebenstudien 327. 1825) and is therefore illegitimate. The family name is also illegitimate unless conserved (Int. Bot. Code. l961ed. Art. 6“, Art. 17 note 1). Article 17 was modified in the 1961 edition of the Code so that Riedl's choice of names was possible at the time he made it. ganthopyrenia Bachmann is distinguished from Artho- pyrenia only in possessing Xanthccapsa rather than Trentepchlia as the phycobiont. The general trend as indicated in the discussion of thallus morphology is to regard the phycobiont as unimportant and not worthy of generic recognition (Riedl 1965, Morgan-Jones & Swinscow 1965). Thus if Myc0poraceae is unavailable and Xanthopyrenia is included in Arthopyrenia, the next available name is Xanthopyreniaceae. If one does not consider ganthcpyrenia to be closely related to ArthOpyrenia and maintains them in separate families the next avail- able name is Arthopyreniaceae Watson (New Phytol. 28: 107. 1929). The ArthoPyreniaceae, as circumscribed by Watson, is actually the best fit for the present grouping and would require very little 24 modification. However, since I do not believe lichenization to be imporatant in this group, I will include the ascolocular pyreno- carpous lichens in the Pleosporaceae Lindau. Key to the species of Great Lakes pyrenocarpous lichens 1. 1. 3. 3. 5. 5. 7’- Ascocarp containing more than one chamber, chambers separated by thin uncarbonized sterile layers--- ----------- -- ------------ -- 2 Ascocarp containing only a single chamber (if more than one are joined then each with an entire carbonaceous wall)-----—-------} 2. Spores muriform, brown--~~---~-------Dermatina pyrenocarpa 2. Spores transversally l-septateo------~-o----(Myc0porellum) Walls of spore thickened, lumina of spores therefore lenticular or .PhOI‘iCa-l, 3070]? mu1torn¢¢* -------------------- anno--o----“ Walls of spore not thickened, lumina of spores therefore cylin- drical, sometimes muriform—---~---—---------------o-----------6 k. Spores hyaline, 7-9—septate, perithecia in groups embedded in a pseudostroma------—--¢-----------Tryphethelium virens h. Spores brown, 3-septate, perithecia not in a pseudo- .trQua-DO----OCOOOOCOOO-GIOGO-e-n-oooucan---—----------------5 Thallus well-developed, hymeniua with abundant oil droplets, IKI+ blue-green, terminal lumina of spores directly against the outer spore wall------—-------------------§Igenula neglecta Thallus poorly develOped, hymenium without much oil, IKI-, terminal lamina of spores separated from the outer spore wall by ‘ 1‘yer of .ndogpar.-------—---o------co----Pz:enula asawae 6 . spor.‘ brO'n.--"--------¢--accuses-sac----ooh-nonoa—aouunooq 6. Spores colorless (occ. becoming tinted in old age but if so almost always shrunken and obviously defunct)---~----~---10 spor.‘ l-..pt‘t.OO-COQOOOO-CC----------------COO--OCO—O-------8 aSpores more than loseptate-------~-------------------c~---e~--9 8. Hyphae at base of pseudothecium, not nucleus, IKI+ violet, occurring on Betula papyrifera-----Microthelia wallrothii 25 8. Hyphae at base of pseudothecium lKI-, on a wide variety of trees other than Betula papyrifera ---Microthelia micula group 9. Spores constantly 3-septate with a thick gelatinous outer layer, always on Populus tremuloides, algae lacking---Melanomma sp. 9. Spores more than 3-septate, usually 5-6-septate, without gela- tinous outer layer, on a variety of trees, algal layer well- deve10ped---o-----~----—--¢-------—-------Molestia leucOplaca 10. Spore“ uniform-nes-c-n—uasoone-nuances-see-cow--Q—nonuuooooovfill 10. Spores transversally septate----------—--~------o--—---13 ll. Asci initially with four spores, spores 35-h5 x 12-13p-----~-- Polyblastistis lactea ll. Asci initially with eight spores, spores up to 25 x qu-----12 12. Pseudothecial wall green or greenish-brown-- . PolyblastiOpsis meridibnalis 12.,Pseudothecia1 wall brown------Polyblastiopsis fallaciosa 13. Spores cylindrical-fusiform to acicular, in one or two bundles in the ascus, which is little longer than the spores---—-o--lh 13. Spores oval to fusiform, strung out in the ascus, which is several to many times longer than the spores----~---~----~--l7 lh. Spores under 30m in length-------~----—-------------—--15 lb. Spores over BOP in length---~--~v--------------------~-16 15. Hucleus IKI-, spores often with pointed ends, on Betula papyrifera-------------------o------Leptorhaphis epidermidis 15. Nucleus IKI+ orange (occ. initially pale bluish), spores with rounded ends, on Populus----------------Leptorhaphis atomaria 16. Nucleus IKI-, on Populus tremuloidego-a--—-------—.——--- Beptorhaphis contorta l6. Nucleus IKI+ bluish, on Prunus and Betula papyrifera---- Lpptorhaphis parameca 1?. Spores in a single row in the ascus, ellipsoid with rounded ends and two equal cells-—-------—-------------—~--c-—-----18 17- spores in more than one row in the ascus, or if in a single row not ellipsoid and equal two-celled--------------~---—---19 26 18. Spores 35-60 2 15-239. pseudothecia 0.5-1.0mm in diameter ArthOpyrenia finkii 18. Spores 21-27 x lO-lZP, pseudothecia 0.5-0.8mm in diameter (ArthOpyrenia alba) l8. Spores 11-15 x 6*8P9 pseudothecia 0.3-0.5mm in diameter Arthopyrenia sphaeroides 19. Spores more than 1-septate--—--------~------------- ------ ---20 19. Spores l-septate (very rarely becoming 3-septate)---- ------- 22 20. Mature spores 5-7-septate, nucleus IKI-, --------- ----- Arthopyrenia quinqueseptata 20. Mature spores )- rarely 4-septate--~----- ----- ---------21 21. Nucleus IKI+ greenish-blue becoming orange, pseudothecial wall brown-----------¢ ----------- . ------- —-glagiocarpa hyalospora 21. nucleus IKI-, pseudothecial wall green-avArthOpyrenia myricae 22. Spores narrow, Z-hr, in a single row in the ascus-—----— (ArthOpyrenia quisquiliae) 22. Spores broader than by, in more than one row in the ascus -Q-O---‘-- ....... --------O------’----------------------23 23. Pseudothecium hemispherical and shiny, not much immersed, asci clavate or cylindrical-elevate, spores coming to a slightly rounded point at both ends---------~---Arthopyrenia willeyana 23. Pseudotheciua not hemispherical and shiny, often immersed, spores truncate at least at one end------e------ ----- --¢----2# 2h. Pseudothecium c. 0.1mm in diameter, flattened, often with a dark hyphal ring, asci obclavate, 2-4 times as long as broad¢-—------—-----------—-e--Arthgpyrenia punctiformis 2h. Pseudothecium 0.3-0.5mm in diameter, hemispherical, asci long cylindrical or cylindrical-clavate, (#)5-7 times as long as broad--~-~----~---~--------§rthcpyrenia thomsonii 27 PLEOSPORACEAE Lindau Engler & Prantl, Natuer. Pflansenfam. 1. Teil, Abt. 1. 428. 1897. ARTHOPYRENIA Hassal. Ricerch. Auton. Lich. 165. 1852. Verrucaria‘Leiophloea Ach. Method. Lich. suppl. 2%. 1803. Leiophloea (Ach.) 8. Gray, Natur. Arrang. Brit. Plants 1: 495. 1821. Eyrenillium Clem. Gen. Fungi #1. 1909. Ascocarp a pseudothecium, spores hyaline, transversally septate with cylindrical lumina. The oldest name for this genus is Leicphloea (Ach.) 3. Gray. Arthgpyrenia Massal. will have to be conserved unless one wishes to split the genus as Riedl (1962) has done and use LeiOphloea for those species with l—septate spores and Arthopyrenia for those with more than l-septate spores. Riedl fails to discuss Clements' (1909) treat- ment which also separated the l-septate spored species from the multiseptate spored species. Arthopyrenia_was retained by Clements as the multiseptate genus and typified it with A.gpyrenuloides (Fee) Huell. Arg. The typification is incorrect since this species was not included in the original publication of the genus. A genus, Pyrenillium, was erected for the l-septate species which is, there- fore, a synonym of LeiOphloea. From the few observations I have made in this genus it seems that the trend toward spores with more than a single septum occurs in several natural groups within the artmus which are not closely related. The division suggested by Clements and Riedl seems unnatural and can be justified only on grounds of convenience in pigeon-holing species. Until relation- Ships within the genus can be more carefully assessed it seems best 28 to maintain a single genus and to conserve Arthopyrenia Massal against LeiOphloea (Ach.) 8. Gray. Arthopyrenia is often cited as having been emended by Mueller irg. (Mem. Soc. Phys. Hist. Nat. Geneve, 16: 428. 1862.). Mueller actually emended Arthopyrenia Hassal. 1859(not 1852). Massalongo (185k) erected the genus Acrocordia for some species removed from ArthOpyrenia. Hueller's emendation consists of placing Acrocordia back in Arthopyrenia as a section and adding two other sections. One of these, Pseudosagedia, has had all of its species transferred to Porina. The other, Leptorhaphis, is generally recognized as a distinct genus. Massalongo's (1852) original genus included a species, A. ggmmata which he later included in Acrocordia so that Mueller's emendation was unecessary in this respect. Therefore, unless one wishes to include Porina or Leptorhaphis, Arthopyrenia cannot be cited as having been emended by Mueller Arg. Arthopyrenia finkii Zahlbr. Cat. Lich. Univ. 1: 306. 1921. gzgenula gemmata var. macrocarpa Willey, Enumer. Lich. New Redford 38. 1892. Eygenula megalospora Fink, Minn. Bot. Stud. 2: 329. 1899. Arthopyrenia macrospora Fink, Contr. 0.8. Hat. Herb. 1a: 237. 1910. (non J. Stein. 1909). Thallus endophloeodal, pale gray to whitish, smooth, with a hyaline cortex-like layer 10-20(35)? thick. Trentepohlia abundant. Pseudothecia scattered, dark brown to blackish, matt, subspherical, (3«-£5-1.0mm in diameter, one third to almost completely immersed. POI-‘0 often at the end of a short neck, ocassionally displaced from 29 the vertical. Pseudothecial wall entire, brown above and below, or hyaline below appearing dimidiate, 30-70? thick, thinner below. Nucleus IKI-. Paraphysoids c. 1? thick, not obviously septate, bran- ched and anastamosed, persistent. Asci long cylindrical, usually with a distinct subglobose dimple, unchanged in XOR, 170-250 3 15-25r. Spores initially eight in the ascus but 1-9 spores often aborting, hyaline, ellipsoid with blunt or somewhat pointed ends, l-septate with equal cells, slightly constricted at the septum especially in age, wall uniformly c. 2? thick, 33-98 x 15-23}. Arthopyrenia finkii has the largest spores and pseudothecia of an apparent series within the genus Arthopyrenia, sometimes recognized as a separate genus Acrccordia Massa1., which have a similar sopre, ascus and parphysoid type differing primarily in size of spores and pseudothecia. ArthOpyrenia sphaeroides is the smallest member: A. alba (Schrad.) Zahlbr. is intermediate. Arthopyrenia alba appar- ently does not occur in North America. All specimens in herbaria so far seen under the name A. alba or A. gemmata have been incorrect- ly identified. This species occurs from southern Minnesota to western Penn- sylvania and on the East Coast in Massachusetts and Maryland. Specimens s8en. UNITED STATES IOWA: Jones County, Wapsipinicon, south of Anamosa, 1252235 3&2 (use). MARYLAND: Baltimore County, ravine at Catonsville, 2133: £92, lit Jan. 1911 (vs). Union Dan, M ggg, 12 .Sept. 1907 (vs). MASSACHUSETTS: Bristol County, New Bedford, Kills! (US). MICHIGAN: Kalamazoo County, Kellog Forest north of Augusta, 30 Harris lkl (MSG); Washtenaw County, Crooked Lake, Harris 870 (M80). MINNESOTA: Blue Earth County, Mankato, £135 222, 1 July 1899 (HIGH, TYPE); Yellow Medicine County, Granite Falls, {1&5 226, 11 July 1899 (MICE). PENNSYLVANIA: Huntingdon County, Penn State College Nature Camp, Thomson 1282 (Thomson). WISCONSIN: Green Lake County, T16N, RISE, sect. l8, E212.l§2l (WIS): Vernon County, T12N, 34w, sect. 26/35, gale 1222 (WIS), Wild Cat Mtn. State Park, Thomson 12818 (WIS). Arthopyrenia myzicae (Nyl.) Zahlbr. Annal. Naturh. Hofmus. Wien. 18: 365. 1903. Verrucaria myricae Nyl. Flora 52: 297. 1869. Verrucaria epidermidis var. aeruginella Nyl. Notiser ur Shllsk. Faun. Fl. Fenn. Forhandl. 8: 173. 1866. Eerrucaria aeruginella Nyl. Flora 55: 365. 1872. In 1872 Nylander combined V. myricae with V. epidermidis var. geruginella, raising the variety to species level as the name for the combined taxon, treating V. myricae as a synonym. However, according to the International Code of Botanical Nomenclature the correct epithet is ngicae since it is the first one used at the species level. Description modified from Keissler (1938). "Thallus" effuse, .indicated by a paler grayish or brownish area on the bark. No algae seen. Pseudothecia scattered, black to greenish-black, elliptical, 0-2-0.1.“ long by 0.1—0.2mm wide, flattened. Pseudothecial wall lack- iilla; below, including bark cell, green to greenish-brown or greenish- black, changing to reddish-purple in nitric acid, 25-35)! thick. 31 Nucleus IKI-. Paraphysoids c. 1r thick, septate, much branched and anastamosed. Asci cylindrical, tip 2—5P thick, without dimple; ex- panding in KOH to 6-10? thick, inner ascus tip protruding 2-3P into the ascus cavity, without any visible canal in the tip; 50-70 x lO-l5p. Spores eight in the ascus, roughly biseriate, fusiform with slightly pointed ends, 3-(#)—septate, cells approximately equal, constricted at the septa; 18-22 x 6'8P‘ This EuroPean species has not been reported from North America before. It is easily recognized by the greenish pseudothecial wall and the three—septate spores. Only a single collection from Isle Royale on 52323 can be definitely referred to this species. Several collections from Iowa and Illinois on guercus also have a greenish pseudothecial wall and three—septate spores but the asci are longer, up to 120u, and the spores are somewhat larger and not at all pointed at the ends. Exaiccati examined. Krypt. Vind. 861 (FH) and 3477 (Us); Magn. 127 (PB). Specimens seen. UNITED STATES MICHIGAN: Keweenaw County, Isle Royals National Park, Rock Barber, Lowe 262 (MICE). Arthopyrenia punctiformis Hassal. Ricerch. Auton. Lich. 168. 1852. Massalongo in his publication of this species clearly is not transferring a species as conceived by any earlier author and does 11:31: cite any basionym except Verrucaria punctiformis auct. Thus the Priority of this name must date from 1852 and not from Persoon 17911 52 or Schrank 1789, whose species Massalongo is often cited as having transferred into ArthOpyrenia. Vainio (1921) credits Arthgpyrenia punctiformis to Arnold (1871), specifically excluding Massalongo's species. He does not give any reasons for this treatment and if the Massalongo type should prove to be a different species, a new epithet will be needed since A. punctiformis Arnold is a later homo- nyne gescription. Thallus end0phloeoda1 or apparently absent, indicat- ed only by a lighter area on the bark. No algae seen. Pseudothecia scattered or rarely clustered and fused, dark brown or black, cir- cular to elliptical in outline, hemispherical to flattened, c. 0.1mm in diameter; occasionally surrounded by a ring of dark hyphae; initially covered by a thin layer of bark but soon entirely uncover- ed. Pseudothecial wall lacking below, 20-30? thick. Nucleus IKI-. Paraphysoids thick, 2-5}, septate and much branched with an almost parenchymatous appearance, often appearing moniliform in KOH. Asci obclavate, clearly broadest below the middle, wall thickest at the tip, expanding strongly in EUR; (30)-40-60 x lZ-ZOP. Spores eight in the ascus, irregularly arranged, hyaline, l-septate, cells some- what unequal with the lower one longer and narrower than the upper, usually not constricted at the septum; occasionally with a gelat- inous outer layer; 16-22 x 5-6P' There is variation in the collections which I have included in this species in regard to the size of the spores, reactions of the asci in K03 and the nature of the paraphysoids. However, they all liave short squat asci enlarged in the lower part and much thickened alt the tip in KOB. Until these variations can be interpreted thru Study of more material than is now available, they are best left together under A. punctiformis. ‘2" This species occurs on smooth, young bark of Acer, Quercus, Populus and Betula throughout the Northeastern United States. A single specimen has been seen from Arizona. Specimens seen. UNITED STATES MICHIGAN: Alger County, Kingston Lake, Harris 661 (MSG), Laughing . Fish Point, Deerton, pm 12;}; (MICE); Charlevoix County, Fox Lake, Beaver Island, Imshaug 22382,(HSC); Keweenaw County, Isle Royale Nat. Park, Angleworm Lake, McFarlin a m 580a (MICH, us). MINNESOTA: County unknown, Minneapolis, £33k 32, 1896 (MICE); Ottertail County, Battle Lake, £355 18, 18 June 1900 (MICH). OHIO: Franklin County, Georgesville, Segue, 1895 (MICH, US). Arthopyrenia quinqueseptata (Nyl.) Muell. Arg. Flora 68: 326. 1885. Verrucaria quinqueseptata Nyl. Mem Soc. Acad. Maine-et-Lcire 4: 58. 1858. (Expos. Syn. Pyrenocarp.). Thallus endophloeodal, light gray to whitish, smooth. Trentepohlia abundant. Pseudothecia scattered to clustered and partially fused, black, occasionally slightly shiny, hemispherical, mostly only about one third immersed but rarely almost entirely immersed. Pseudothecial wall dimidiate or entire but very thin below, brown above, hyaline below if entire and appearing dimidiate, 24-30? thick. Nucleus IKI-. Paraphysoids c. If thick, septate but not much branched or anasta- mosed. Asci cylindrical-elevate, somewhat thickened at the tip, with m.small apical dimple, expanding very slightly in KOH; 60-65 x 10-15». Spores eight in the ascus, biseriate or triseriate, hyaline, fusiform, Often slightly narrower at one end, 5-(7)-septate transversally, ot‘ten with 1-2 cells longitudinally septate, not or only slightly 34 constricted at the septa; 19-25 x 5-7P. Pycnidia often present, black, c. 0.1mm in diameter. Macro- conidia hyaline, cylindrical fusiform with rounded ends, mostly 7-septate, 22 x 5P' This is a very interesting species in several respects. The tendency for the spores to become longitudinally septate would lead one to think that it might be intermediate between Arthopyrenia and Polyblastigpsis. However, the extremely paraphysis-like paraphysoids make this unlikely since the paraphysoids in PolyblastiOpsis are usually almost parenchymatous. The species is also difficult to separate from Porina since the paraphysoids are so little branched and anastamosed. A very closely related species, Arthopyrenia faginea (Schaer.) Swinsc., has only very recently been transferred from Porina (Swinscow 1965). The American species differs only slightly from that species. The spores in A. faginea are larger and consistently have seven transverse septa with no tendency to become submuriform. Both species, as does the also related Porina affinis (Massa1.) Zahlbr., possess a unique type of fusiform multiseptate macroconidia. Porina affinis has not yet been transferred to Arthg- pygenia but Swinscow (1965) has suggested that it should be. I have not collected this species in the Great Lakes region. Specimens have been seen from Iowa, Minnesota and Pennsylvania so that it seems that the species should eventually be found in our area. Exsiccati examined. Cummings I. 299 (HIGH, MSC); Cummings II. 250 (MSG). Specimens seen. UNITED STATES ILLINOIS: Menard County, Athens, Hall, 1878 (HIGH). IOWA: Fayette County, Fayette, Fink, June 1893 (WIS), Pink, 35 June 1899 (HIGH), £12k, 1895 (US), 3335, 1896 (US). MINNESOTA: Beltrami County, Red Lake, Eigk 882, 26 July 1900 (HIGH); Blue Earth County, Mankato, {325 _2_9_§_, A July 1899 (MICH). PENNSYLVANIA: Chester County, locality unkown, collector un- known, Hb. E. Michner, 1852 (HIGH), West Chester, E33213, July 1898 Arthopyrenia quisquiliae sp. nov. Thallus endOphloeodal, pale gray, smooth, Trentepohlia present, well-developed but not abundant. Pseudothecia scattered or clustered and occasionally fused, black, mostly shiny, subglobose to slightly conical, about one third immersed, 0.3-O.#mm in diameter. Pseudo- thecial wall entire or dimidiate, brown above, 30-60u thick, thinner below and colorless, or lacking below. Nucleus IKI-. Paraphysoids c. 1? thick, septate, slightly branched and anastamosed. Asci cylin- drical, not thickened at the tip, without an obvious dimple; tip ex- panding in KOH forming a small dimple. Spores eight in the ascus, obliquely uniseriate, fusiform, l-septate with equal or slightly unequal cells, slightly constridid at the septum, occasionally some- what bent, 19-20 x 2-RP. No species has been described in the temperate or boreal floras with such narrow spores uniseriate in the ascus. Unless there is a trepical species name available, this is clearly a new species. This species has not yet been found in the Great Lakes region but it should be expected since it occurs in Iowa and Minnesota. The only collection for which the substrate is identified is on Ulmus. 36 Specimens seen. UNITED STATES ILLINOIS: Fulton County, Canton, ESlE (US). IOWA: Boone County, Fink, 27-8 July 1903 (MICE); Fayette County, Fayette, {125, 1896 (MICE, MSC, US); Jones County, Napsipinicon, south of Anamosa, Imshaug 28122 (MSG). MARYLAND: Baltimore County, Seaman's Run, 2113;, 1 Dec. 1911 (US). MINNESOTA: Beltrami County, Red Lake, Fink (MSG). ArthOpyrenia sphaeroidee (Nallr.) Zahlbr. Engler & Prantl, Natuerl. Pflanzenfam. I Teil, Abt. I‘: 65. 1903. Verrucaria sphaeroides Wallr. Flora Crypt. Germ. 3: 300. 1831. Thallus endOphloeodal, pale gray to whitish, slightly roughened to smooth. Trentepghlia abundant, well-developed, forming short chains. Pseudothecia scattered, dark brown to black, occasionally somewhat shiny, hemispherical, one third to two thirds immersed, 0.3-0.5mm in diameter. Pseudothecial wall entire, brown above but mostly hyaline below and appearing dimidiate, 60-80u thick above, thinner below. Nucleus IKI-. Paraphysoids c. 1? thick, not obvious- ly septate, branched and anastamosed. Asci cylindrical, with an obvious dimple in the tip, unchanged in KOH, 80-90 x 9-10P (80-120 x 8-18», Keissler 1938). Spores eight in the ascus, uniseriate, hyaline, oval with blunt ends, l-septate with equal cells, slightly constricted at the septum, spore wall minutely roughened; 12-15 x 7-8u (11-21 x 8-12P, Keissler 1938). ArthOpyrenia sphaeroides has usually not been recognized in North America by American lichenologists. It is commonly found in herbaria under A. alba (A._gemmata) or A. biformis. All of the 37 European specimens of A. sphaeroides seen have the spore wall mi- nutely roughened. However, in the Northeastern United States there are a large number of collections in which the spores are smooth- walled and slightly smaller. The spores are quite close in appear- ance to those of A. biformis (Borr.) Massal. but A. biformis has much smaller, completely superficial pseudothecia and also differs in ascus structure and spore arrangement. These collections prob- ably represent an undescribed species. However, it is part of a very complex species-group centered around A. biformis, A._sphaeroides, and A. alba. I do not feel well-enough acquainted with this group to describe a new species in it at this time. All of the smooth spored collections come from Maine, Massachusetts and Maryland with one exception, a single collection from Michigan (Crooked Lake, Washtenaw County, Harris 869 MSG). ArthOpyrenia sphaeroides proper‘ with the exception of collections from the Gaspe Peninsula)has been found only in Iowa and western Michigan. It occurs mainly on Quercus alba'but also once on Age: and 22215: In EurOpe it has been reported on Fagus,garpinus, Fraxinus and Pepulus. Exsiccati examined. Krypt. Vind. 2152 (FE, US); Malbr. 398 (MSG). Specimens seen. CANADA QUEBEC: Gaspe Peninsula, Eckfeldt, 1880-1883 (MICH), Macoun 12 Aug. 1882 (US). UNITED STATES IOWA: Boone County, The Ledges, 2335, 27-8 July 1903 (MICH); Fayette County, Fayette, Fink, July 1899 (HIGH, WIS). MICHIGAN: Keweenaw County, Isle Royals Nat. Park, Siskowit Lake, Lowe 863 (MICH); Newaygo County, near Hesperia, Imshaug 21196 (MSG). 38 ArthOpyrenia thomsonii sp. nov. Thallus endOphloeodal, gray, smooth to roughened. Trentepohlia present but very scarce. Pseudothecia mostly clustered and occasion- ally fused, rarely scattered, dark brown, rough, hemispherical, com- pletely immersed when young, becoming up to three-fourths exposed in age. Exit pore mostly depressed, pale, large, up to 60p in diameter. Pseudothecial wall dimidiate, h0-75P thick, the outer layers consis- ting mostly of bark cells, only the inner 25u or so consisting en- tirely of fungal tissue. Nucleus white or yellowish-white, opaque not translucent as is usual in Arthopyrenia. Paraphysoids c. 1-2P thick, septate, branched and anastamosed. Asci cylindrical to cylin- dric-clavate, without a dimple or with a very small one, tip not ex- panding in KOH, 80-90 x 12-15p. Spores eight in the ascus, obliquely uniseriate to biseriate, hyaline or very pale brown in old age, fusi- form to elliptic-fusiform, l-septate becoming 3-septate in old age, cells equal or slightly unequal, 22-25 x 7-8P' The only material seen of this species is an exsiccat, Thomson 96 (sub A. leucochlora). on Gleditsia triacanthos. sPecimens seen. UNITED STATES ‘WISCONSIN: Fond du Lac County, 2 miles north of Rosendale, Thomson, 16 May 1997 (HIGH, MSG, WIS). Arthopyrenia willeyana sp. nov. Thallus endOphloeodal, light gray to whitish, well-deveIOped, often somewhat powdery appearing but mostly smooth. Trentepohlia very abundant and well-develOped. Pseudothecia mostly scattered, black, shiny, hemispherical to subglobose, almost completely immersed 39 when young but very soon becoming almost entirely superficial, 0.15-0.25mm in diameter. Pseudothecial wall entire, brown or occa- sionally somewhat greenish in KOH, hyaline below and appearing di- midiate, 20-30p thick. Nucleus IKI-. Paraphysoids c. 1r thick, septate, much branched and anastamosed, persistent, when dry forming a layer just within the pseudothecial wall leaving a hollow in the center of the ascocarp. Asci cylindric or clylindric-clavate, not much thickened at the tip, small dimple evident or not; expanding in KOH to 5-7P thick at tip, the dimple becoming more pronounced; #0-90 x lO-ldr. Spores eight in the ascus, biseriate, hyaline, l-septate, often some spores becoming 2-3-septate, cells unequal, the lower usually narrower and shorter than the upper. spore wall not constric- ted at the septum, this, without gelatinous outer layer; 11-18 x 3-5r. Material of this species has been found in herbaria 35553227 pyrenia leucochlora Muell. Arg. to which it is quite similar. Artho- pyrenia leucochlora was described from material sent by Willey. Material under this name from Willey's herbarium in the Smithsonian Institutuion is mixed. Two-of his collections which have a rather peculiar thallus seem to fit Mueller's description best. Unfortun- ately, they are not conspecific with the Great Lakes collections. How- ever, the rest of Willey's material is. I have not seen Mueller's type but feel sure from the original description of the species that it is the species which does not occur in the Great Lakes. On this, basis I have described the Great Lakes material as a new species. Thomson's Lich. Wise..Exsic. 96,issued as A. leucochlora, is neither of the species involved here but anonher, apparently new species, described in this paper as A. thomsonii. 40 Most of the collections of A. willeyana come from either guercus 2183 or Ulmus americana but it also occurs on guercus borealis, Carya, Fraxinus, Platanus, Acer saccharum, Pepulus tremuloides, Betula lutea and lignum. This species is the most common and abundant pyrenolichen in southern Michigan. I have also seen specimens from Illinois, Ohio, Massachusetts and Long Island. Specimens seen. UNITED STATES IOWA: Dickson County, Lake Okoboji, Thomson 19289 (WIS); Jones County, Wapsipinicon, south of Anamosa, Imshaug 28093, 28105 (MSG). MICHIGAN: Alger County, Kingston Lake, Harris 821 (MSG); Barry County, Shaw Lake, Harris 888 (MSG); Berrien County, Warren Woods, Harris 21 . g1§§,_g12 (MSG); Clare County, Thuja swamp west of Farwell, Harris 221. 228 (use); Clinton County, near Rose Lake, Harris 22g (use); Eaton County, near Vermontville, Imshaug £2121, 1232§_(MSC); Emmet County, Harbor Springs, 831gg,112, 8 Sept. 1931 (MICE); Ingham County, Sanford Woodlot, East Lansing, Harris 821, 58, 32, 228, 82, 82, 82. 292. 228, 298. 212 (MSG), Baker Woodlot, East Lansing, §3££1£ 22, 21 (MSG), Holt Bog southwest of Holt, Harris 22; (MSG); Iosco County, Silver Valley State Forest campground north of Tawas, Harris §1§ (MSG); Kalamazoo County, oak woods west of Richland, Harris 118, 118, 118, 11;, 121_(MSC), Kellog Forest north of Augusta, Harris 278, 311, 312, 26, #1 , E18, 212,{21&_(MSC); Montcalm County, Coley Lake north of Sheridan, Harris 1981 (MSG); Washtenaw County, Crooked Lake, Harris 1 , $21.§20 12, 18, 8 , 822, 822, 881, 885, 866, 862, 868 (MSG) WISCONSIN: Dane County, Univ. of Wise. Arboretum, gagggg, 23 Oct. 1965 (WIS); Rock County, Thibault Point, Lake Koshkonong, Thomson 1222 (Thomson). #1 DERMATINA Almq. K. Svensk. Vetensk.-Akad. Handl. l7: 8. 1880. Mycoporum Flot. in Koerb. Grundriss der Cryptogam. 199. 1848. (non Meyer 1825). Lscocarp with more than one distinct multiascal chamber, each with its own exit pore; spores brown and murifora. Dermatina pyrenocarpa (Nyl.) Zahlbr. Cat. Lich. Univ. 1: 550. 1922. Mycoporun pyrenocarpum Nyl. Flora #1: 381. 1858. Thallus endophloeodal, indicated by a lighter area on the bark. Trentepohlig_present in scattered clumps, not abundant. Ascocarps scattered, dark brown to black, often slightly shiny, cushion-shaped, elliptical in outline, 0.2-1.0mm in length, usually 0.1-0.2mm less in width; containing l-lO separate chambers, the upper part of the chambers often raised giving the ascocarp a bumpy appearance. Pores of chambers large, usually easily visible due to their lighter color. Upper layer of stroma dark, carbonaceous, c. 20-40P thick, not pene- trating very far downward into the sterile layers separating the chambers which are mostly hyaline and c. 10-20? thick; stromatic layer at the base hyaline or somewhat carbonaceous, especially be. tween the chambers. Nuclei IKIa. Paraphysoids not distinct, short septate, such branched and anastamosed, appearing almost parenchy- matous. Asci obclavate-saccate, much thickened at the tip which ex- pands greatly in KOH, 70-85 x 35-50p. Spores eight in the ascus, irregularly arranged, initially hyaline but soon brown, oblong- ellipsoid, murifors with 7-9-(11) transverse septa and 1-3 longi- tudinal septa, constricted only at the median septum dividing the #2 spore into two, usually unequal, halves; 30-50 x 12-15? (33-46 x 13-19} in Fink 1935). This species is the only Great Lakes representative of a group of three genera which have more than one chamber in the ascocarp. Care must be taken not to confuse these genera with members of the Arthoniaceae where single asci are randomly scattered throughout an irregular or disk-like stroma. Perhaps the best distinguishing character is that under a dissecting microscope Dermatina and its relatives have obvious exit pores for each chamber, while such Open- ings are never present in the Lrthoniaceae. Also the sultichambered nature of Dermatina is usually obvious and the fruit resembles a bunch of small black beads glued together. The other two genera, EZEE‘ porellum and MycoporOpsis have not yet been found in the Great Lakes region. I have, however, seen material from Ohio (without locality) and from Iowa which seems to be an undescribed species of MZCOECr- ElEEE‘ The two Ohio collections from the Pink Herbarius are appar- ently type material of Dermatina ohiense (Fink) Zahlbr. The name was originally published by Nylander (1857) as a variety of D.+pyrenocarp§without any description. Fink (1935) validated the name at the species level. According to his description, the spores are murifors but I have found only l-septate spores in Fink's mater- ial. Since Fink's specimens are at variance with his description and since they are without locality, Dermatina chiense is best treated as a doubtful species until further studies can be made. Dermatina pyrenocarpa has been found on the bark of QEEE rubrum, Carpinus caroliniana, and [2525 grandifolia. Specimens verified come from the southeastern Great Lakes and the East Coast from Massa- chusetts to Tennessee. The species was originally described from 43 Mexico but I have not seen the type. Fink (1935) reports it from California but the only specimen in his herbarium under this name from California is an Arthothelium. sticcati examined. Cummings I. 286 (MSG); Cummings II. 218 (HIGH, MSG); Lojka 150 (MICE); Macoun II. 199 (MICE). Specimens seen. CANADA ONTARIO: Hastings County, Central Ontario Junction, Macoun, 22 Oct. 1893 (MICE). UNITED STATES DELAWARE: New Castle County, Newark, Commons, 6 July 1891 (MICH). DIST. OF COLUMBIA: Takoma Park, Williams & 8238;, 1898 (MICK). MASSACHUSETTS: Bristol County, New Bedford, Willey, 1885 (MICK). MICHIGAN: Cheboygan County, gorge near Douglas Lake, Nichols, June-Aug. 1925 (MICE). OHIO: Fairfield County, Sugar Grove, 8853: 838, 28 Jan. 1893 (MICE). TENNESSEE: Hamilton County, Chatanooga, Calkins 2h? (MICE). LEPTORHAPHIS Koerb. Syst. Lich. Germ. 371. 1855. conserved over Endophis Norm. Nyt. Hag. Naturv. 7: Zko. 1853. Ascocarp a pseudothecium, spores long narrow fusifors-cylindric- al to acicular, in one or two bundles in the ascus which is little longer than the spores. a». Leptorhaphis atomaria (Ach.) Szatala Magy. Bot. Lap. 26: 31. 1928. Lichen atomarius Ach. Lich. Suec. Prodron. 16. 1798. Leptorhaphis tremulae Koerb. Syst. Lich. Germ. 372. 1855. Vainio (1921) examined the Acharian material of Lichen atomarius and determined that it was a Leptorhaphis and not an Arthopyrenia as has often been assumed. The transfer of Lichen atomarius to £2232? rhaphis has been attributed to Magnusson (1936). "Thallus" not visible or indicated by a lighter area on the bark. No algae seen. Pseudothecia scattered, dark brown to black, subglobose to hemispherical, initially immersed but very soon almost completely exposed, 0.1-0.2nm in diameter. Pseudothecial wall dark brown, almost entire but usually lacking at the very base, 15-3OP thick. Nucleus IKI+ orange or occasionally pale bluish initially. Paraphysoids not distinct, c. In thick, short celled, much branched and anastamosed. Asci short clavate, not much thickened at the tip, mostly without a dimple; tip thickening slightly in KOH and forming a dimple; #0-50 x 8-15P. Spores eight in the ascus, in one or two bundles, cylindric-fusiform, usually with rounded ends, often curved, l-3-septate, 20-28 x Z—hn. This is a European species which has not been previously re- ported from North America. It occurs on Pepulus on both continents. North American material is IKI+ orange which has occasionally been taken as the basis for a distinct variety in Europe. I have seen no collections of this species from outside the Great Lakes region. Exsiccati examined. Magn. 126 (FH); Rabenh. 187 (MICE). “5 Specimens seen. CANADA ONTARIO: Algoma District, Lake Superior Prov. Park, lookout h miles north of campground, Harris 282.(MSC); Thunder Bay District, Sibley Prov. Park, one mile east of Lake Marie Louise, Erbisch 8&8 (MSG). UNITED STATES fl IOWA: Allamakee County, Effigy Mounds Nat. Monument, bluffs along Mississippi River, Inshaug 22906 (MSG); Delaware County, Maquoketa River north of Dundee, Imshaug £8911 (MSG). MICHIGAN: Emmet County, Wilderness State Park, Harris 889 (MSG); Iosco County, Silver Valley State Forest campground north of Tawas, Harris 832 (MSG); Iron County, junction USP hwy. 139 and Co. 657 south of Kenton, Harris 2§;.(MSG); Keweenaw County, Isle Royale Nat. Park, Hay Bay, 9313,22; (MICE), Forbes Lake, Wetmore 888g (MSG); Ontonagon County, Porcupine ths.‘% mile east of parking lot, Harris §§2 (use). Leptorhaphis gontorta Degel. Ark. Bot. 30A: 13. 19HO. "Thallus" not visible or indicated only by a lighter area on the bark. No algae seen. Pseudothecia scattered, black, occasion~ ally quite shiny, subglobose to hemispherical, 0.1~O.2mm in diameter. Pseudothecial wall dimidiate, lacking below, dark brown, 15-25P thick, Nucleus IKI-. Paraphysoids c. 1? thick, septate with short cells, branched and anastamosed. Asci cylindrical, not much thick- ened at the tip; tip thickening in KOH and occasionally forming a dimple; 50-65 x IO'IZP (45-5“ x 10°5'13P in Degelius 1940). Spores eight in the ascus, in a single bundle, hyaline, acicular, l-(3)- septate; 35-50 x 2-3P. #6 Degelius (1940) described this species from Maine on guercus borealis and also found it on Prunus in North Carolina (Degelius 19kl). The Great Lakes material is all from Populus and differs from the type in having larger, more rounded and emergent pseudothecia. It differs from L. paramega in lacking any IKI reaction and may not be distinct. Specimens seen. CANADA ONTARIO: Algoma District, Lake Superior Prov. Park, north of Sand River, Imshaug £812; (MSG), lookout 9 miles north of campground, Harris 288 (MSG). UNITED STATES MAINE: Sagadahoc County, Prince's Point near Brunswick, Degelius, 3 Sept. 1939 (isotype US). MICHIGAN: Cheboygan County, Pine Grove State Forest campground east of Wolverine, Harris 282 (MSG); Iron County, junction of USF hwy. 139 and Co. 657 south of Kenton, Harris 288, 282 (MSG); Keweenaw County, Isle Royals Nat. Park, McGargo Cove, Eggg‘flfi;_(MIGH); Menominee County, Big Cedar River State Forest campground north of Cedar River, Harris Zég_(MSC); Otsego County, Grass Lake, Harris, 10 Oct. 1962 (MSG). WISCONSIN: Vilas County, Trout Lake, Cglbersonl58h (WIS). Leptorhaphis epidermidis (Ach.) Th. Fries Lich. Arctoi 273. 1860. Lichen epidermidig Ach. Lich. Suec. Prodr. 16. 1798. Lichen epidermidis has been transferred to both ArthOpyrenia and Leptorhaphis. The first to realise that Acharius and others had #7 included several taxa in this species was Nylander (1852). He used the name Verrucaria epidermidis for what is now reCOgnized as an Arthopyrenia and gave a new name to the Leptorhaphis, Verrucaria oxzspora. This treatment was followed in essence, notably by Koerber and Massalongo, until Th. Fries Lichenes Arctoi in 1860. Fries re- versed the previous treatment and used gpidermidis in Leptorhaphis and placed the Arthopyrenia in synonymy. From that time until the early 1900's oxzspora and epidermidis were used about equally for the Leptorhaphis. Nylander's interpretation is supported by the use of Arthopyrenia epidermidis by the describer of that genus, Massalongo, and by the fact that the type of Koerber's genus, Leptorhaphis, is L. oxygpora (Nyl.) Koerb. Th. Fries seemingly arbitrary reversal has been supported by the two modern treatments of European pyrenolichens, Vainio (1921) and Keissler (1938). If the type contained spores, the problem would be easily resolvable. Vainio (1921) has examined what he says is the Acharian type, although he does not give his reasons for his choice of a specimen. It is, according to Vainio, a mixed collection, neither part containing spores. Thus the identity of neither half can be definitely determined but Vainio suggests that one is Leptorhaphis and the other an Artthyrenia. It is hard to believe that Acharius' material was limited to a single collection. Further research should be done in the Acharian herbarium to verify Uainio's conclusion. If there is no satisfactory way of typifying the species, the name should be dropped in both Leptorhaphis and Arthopyrenia. Description. "Thallus" not detectable. Algae absent or per- haps a few protococcoid cells at the base of the pseudothecia. Pseudo- thecia scattered, black, dull to slightly shiny, hemispherical or #8 slightly flattened, c. 0.2mm in diameter, surrounded by a dark elliptical hyphal ring, c. 0.1mm wide. Pseudothecial wall dimidiate, lacking below, dark brown, 20-30p thick. Nucleus IKI-. Paraphysoids c. 1‘2} thick, septate with short cells, branched and anastamosed. Asci short clavate, often somewhat truncate at the tip, not much thickened at the tip, sometimes with a shallow dimple; expanding in KOH to c. 5? thick with a pronounced cylindrical dimple; 32-k5 x 10-15P. Spores eight in the ascus, mostly in a single bundle, hya- line, cylindrical-fusiform, often with one or both ends sharp- pointed, usually curved or twisted, l-(3)-septate, not constricted at the septum, 20-30 x z-fin. 'Thie is a widespread circumboreal species rare south of the Hemlock-White Pine-Northern Hardwoods region. It occurs on Betula papyrifera and 318p0pulifolia. Exsiccati examined. Malbr. 100 (MSG); Norrlin 389 (MSG); Thomson #1 (HIGH, MSG, Thomson, WIS). Specimens seen. CANADA ONTARIO: Algoma District, Lake Superior Prov. Park, just north of Sand River, Imshaug gégifi (MSG); Simcoe County, Blue Mtns. between Singhampton and Duntroon on hwy. 2h, Imshaug géggz (MSG). UNITED STATES MICHIGAN: Alger County, Kingston Lake, Harris 888, 882 (MSG); Charlevoix County, Fox Lake, Beaver Island, Imshaug £1822 (MSG); Clare County, Thuja swamp north of Harrison on old US 27, Haggis figfi_(MSG); Keweenaw County, Isle Royals Nat. Park, Rock Harbor, £212 222 (HIGH); Montcalm County, Coley Lake north of Sheridan, 835583 1022 (MSG); Ontonagon County, Porcupine Mtns.‘% mile east of the “9 parking lot, EEEEEE 673, 678, 686,_888, 68 , 82; (MSG). MINNESOTA: Beltrami County, Bemidji, §$£§.§Z£* 12 July 1900 (MICH); Blue Earth County, Mankato, §i£5.1299 27 June 1899 (MIGH); Lake County, Beaver Bay, £325.8221 13 July 1897 (MICE); Otter Tail County, Henning, ZlEE.l§2v 25 June 1900 (MIGH); St. Louis County, Duluth, Kimball, 1900 (US). ' WISCONSIN: Douglas County, Brule River Valley, Thomson, 9 Aug. 1996 (MICE, MSG, Thomson, WIS). Leptcrhaphis parameca (Massal.) Koerb. Parerg. Lich. 386. 1856. Sagedia parameca Massal. Symmict. Lich. 97. 1855. Description modified from Keissler (1938). "Thallus" not vis- ible. No algae seen or a few scattered protococcoid cells on the bark surface. Pseudothecia scattered or clustered and occasionally fused, dark brown, flattened, circular or elliptical in outline, 0.15-0.2mm in diameter, with a dark hyphal ring (poorly developed in Great Lakes material). Pseudothecial wall dimidiate, lacking below, dark brown, 0. 25-30u thick. Nucleus IKI+ blue-green. Para- physoids septate with short cells, branched and anastamosed. Asci clavate, slightly thickened at the tip, not much changed in KOH, 30-50 8 7-10P. Spores eight in the ascus, in a single bundle, hya- line, acicular, pointed at both ends, with several indistinct septa, 30-50 x 2.5-3P. This European species has not previously been reported from North America, but American material seems to agree closely with EurOpean material. Known only from three Michigan collections, one on Prunus, the others on Betula papyrifera. 50 Specimens seen. UNITED STATES MICHIGAN: Alger County, Kingston Lake, Harris 882, 888 (MSG); Clare County, Thuja swamp north of Harrison on old US 27, Harris 288’(MSC); Roscommon County, Thuja swamp, Backus Lake Flooding, Imshagg 25991 (MSG). MIGROTHELIA Koerb. Syst. Lich. Germ. 372. 1855. The status, both nomenclatural and biological, of the genus Microthelia has been and still is in a state of confusion. Mueller and von Arx (1962) have discussed these difficulties in detail. I shall comment briefly on them. Microthelia was erected by Koerber as a lichen genus. Later workers considered it to be synonymous with Didymosphaerig Fuck. (1869), a non-lichen genus. Although Microthelia is the older name, they refused to use it for a non-lichen genus since it was a lichen name. This cannot be done any longer since for the purposes of nomenclature the Int. Code of Botanical Nomenclature does not recognise lichens as distinct from fungi. Currently both Microthelia and Didymosphaeria are regarded as valid, distinct non- lichenized genera. Vainio (1921), however, divided Microthelia Ioerb. into two genera, Microthelia, saxicolous and lichenized, and Didymosphaeria, corticolous and non-lichenized. The genus Micro- thelia Koerb. has been typified, both thru usage and lectotypifi- cation, by M. micula Koerb. (Mueller A von Arx 1962), a corticolous species. Therefore, Vainio's Didymosphaeria must bear the name Microthelia Hoerb. This leaves the saxicolous lichen genus without a name. It seems to be distinct from Microthelia on other grounds 51 in addition to lichenization. It is probably worthy of generic recognition. This would result in three distinct and nomenclaturally correct genera. gicrothelia Koerb. and Didymosphaeria,Fuck., non- lichenized, and a third unamed genus which is lichenized. In this paper the only species considered are referable to the genus £1352- thelia as traditionally treated by lichenologists as well as by modern mycologists. There is a great need for close study of this group to determine more clearly both generic and specific limits. I have no "feel" for this group and as a result have not tried to resolve the material centered around M. micula into definite taxo- nomic entities as indicated under the species. Microthelia micula Koerb. Syst. Lich. Germ. 373. 1855. Mueller and von Art (1962) use the name M. biformis (Leight.) Hassal. This name is based on a nisidentification by Leighton of gzgenula biformis Borr. Leighton clearly attributes his name to Borrer. Borrer's species is an Arthopyrenia so that Leighton'e combination cannot be used as a basionya in Microthelia. ”Thallus" indistinct or indicated by a gray blotch on the bark. No algae seen. Pseudothecia scattered to clustered, black, dull, hemispherical, 0.2-0.3mm in diameter. Pseudothecial wall dimidiate, occasionally almost entire, brownish-black, 50-60? thick. Pore large. Nucleus IKI+ bluish or IKI-. Paraphysoids septate, branched and anastamosed. Asci cylindrical to slightly obclavate, tip thicker than the walls, with an indistinct dimple; expanding greatly in KOH to form a long narrow dimple. Spores eight in the ascus, biseriate or irregular, brown, ovoid, l-septate, constricted at the septum, cells usually unequal, 12-16 x S-8p. “I According to Scheinpflug (1958) the spores in this species are equal-celled and unequal in M. applanata (Fr.) Mueller, an otherwise very similar species. All of the collections which I have examined vary in this respect within a single pseudotheciun. My main diffi- culty here seems to be that the characters which I have tried to use to separate collectionsseem to be varying entirely independently of one another. There seems to be a group with smaller spores, mostly under 22p long and 10? wide, but an overlap occurs with a group which has spores mostly 22~3OP.long by lO-er wide. The majority of the collections have a minutely roughened spore wall but a few have smooth walls. Hepp's Fleet. Europ. 108, cited by Mueller, has smooth spore walls. Some spores have a gelatinous outer layer, others do not. According to Vainio (1921) the nucleus is IKI-, while Keissler (1938) finds that it is IKI+ bluish. Degelius (1941) comments that he has found H. micula to be both IKI+ and IKI-. There is also a great deal of variation in the completeness of the pseudothecial wall. Specimens seen. CANADA ONTARIO: Algoma District, Lake Superior Prov. Park, % mile north of Doc Grieg Lake, Harris 226, iél’{2§§’ 262, 26E, 262, 266, :82 (1450). UNITED STATES MICHIGAN: Alger County, Kingston Lake, Harris 612, 616, 612, 650, ézl, 622, ééé (MSG); Charlevoix County, old orchard, Beaver Island, Imshaug £1222 (MSG); Cheboygan County, Pine Grove campground east of Wolverine, Harris 781, 802 (MSG); Clare County, oak woods north of Harrison, Harris 231 (M50); Emmet County, Wilderness State Park, Harris 802 (MSG); Ingham County, Baker Woodlot, East Lansing, Harris 53 22, 26, 6 , er(MSC); Iosco County, Silver Valley campground north of Tawas, Harris 812, 820 (MSC); Iron County, Teepee Lake south of Kenton, Harris 695 (MSG); Isabella County, woods north of Mt. Pleas- ant, Imshaug glgég (MSG); Kalamazoo County, Kellog Forest north of Augusta, Harris 322, 223, 2;, 23. 122. :32, :12 (use); Keweenaw County, Isle Royale Nat. Park, outlet of Siskowit Lake, L312 6% , £82.(MICH); Mackinac County, vicinity of Sugar Loaf Rock, Mackinac Island, Imshaug 222200 (NSC); Menominee County, Big Cedar River camp- ground north of Cedar River, Harris 222 (MSG); Ontonagon County, Porcupine Mtns.,'% mile east of parking lot, Harris ézg,‘§§21 ééfl, gig (MSC); Washtenaw County, Crooked Lake, Harris 3 (M50). MINNESOTA: Blue Earth County, Mankato, Eing’éé, 24 June 1899 (MICE), [igg’lg2, 26 June 1899 (MICE); Pennington County, Thief River Falls, {$35,228, 18 July 1900 (MICE); Iellow Medicine County, Granite Falls, Fink 212, 1“ July 1899 (MICE). Microthelia micula var. quadriloculata (Fink) Zahlbr. Cat. Lich. Univ. I: 263. 1921. gyrenula cinerella var guadriloculata Pink, Contr. U.S. Nat. Herb. l#: 239. 1910. The type of this variety consists only of pycnidia with brown macroconidia, ellipsoid and )~(#)-septate with an occasional longi- tudinal septum. These pycnidia have not been found to my knowledge to be associated with any perfect state so that the material is in essence undeterminable. However, the name should be removed from Hicrothelia. Specimens seen. UNITED STATES MAINE: Knox County, Rockland, Merrill, 25 May 1910 (MICE). 54 MINNESOTA: Beltrami County, Bemidji, Fink 622, 11 July 1900 (MICH); Blue Earth County, Mankato (Rapidan), Fink 162a, 28 Jan. 1899 (MICE); Cook County, Grand Portage Island, Fink 82, 24 June 1897 (type, MICE, MSC). Microthelia wallrothii (Hepp) Rehm Bedwigia 18: 163. 1879. Expenula wallrothii Hepp. Flecht. Eur0p. 709. 1860. Description modified from Keissler (1938). "Thallus" indistinct. No algae seen. Pseudothecia scattered, black, often slightly shiny, hemispherical, often depressed in the center, 0.18-0.3mm in diameter; surrounded by a well-deveIOped dark elliptical hyphal ring which may be up to 0.15mm wide. Pseudothecial wall entire, brownish-black, thinner below, rarely lacking below in the center, often extending outward from the edges a little. Nucleus IKI-. Paraphysoids 1.5-2? thick, septate, branched and anastamosed. Asci clavate, tip some- what thickened, without a dimple, hO-SO x lk-18P. Spores eight in the ascus, biseriate, brown, ellipsoid, 1-septate, not constricted at the septum, 10-17 x 6-7P. A rather Peculiar type of sacroconidia has been attributed to this species, but I have found them associated with two other pyreno- lichens on Betula, both in this country and in Europe. Thus, spec- imens without ascocarps should not be assigned to this species on the basis of conidia alone. The pynidia are black, c. 0.05-0.1mm in diameter, with macroconidia brown, oval, l-septate, appearing polarilocular, 10~12 x 5-6u. Microthelia wallrothii_is unusual in that some of the hyphae at the base of the ascocarp and, perhaps, of the hyphal ring turn pale 55 violet in IKI. This reaction occurs sporadically in some collec- tions of Leptorhaphis and PolyblastiOpsis but appears to be constant in M. wallrothii. Both American and European material are variable in regard to the thickness of the pseudothecial wall below the nucleus and it is even occasionally lacking below. Microthelia oblongata Muell. Arg., described from material collected in Massachusetts by Willey, is very similar but lacks the IKI+ violet reaction of the pseudothecial hyphae and the ascus tip reacts differently in KOH. Microthelia wallrothii occurs only onoBetula papyrifera in the United States. The majority of collections come from Iowa and the Black Hills. It reaches the Great Lakes only in northern Minnesota. Specimens seen. UNITED STATES IOWA: Fayette County, Fayette, 2335, June 1894 (MICH, Thomson, WIS), £335, 19 Aug. 1897 (MICH). MINNESOTA: Cook County, Grand Portage, 2355;22, 21 June 1897 (MICE). MOLESTIL gen. nov. Ascocarp a pseudothecium, spores brown, with more than one transverse septum, lumina cylindrical. Type species: Verrucaria leucoplaca Wallr. The only species known is M. leucoplaca. It has previously been included in Eigenula. However, on the basis of the cylindrical spore lumina and the presumably ascolocular nature of the ascocarp it must be excluded from Pygenula, which has lenticular or spherical lumina and is ascohymenial. Vainio (1921) placed the species in the 56 non-lichenized genus Leptosphaeria but stated in his discussion, "Etiam sporarum indole haec species a Eigenulis differt, et inter lichenes, si gonidiis instructs esset, genus autonomum constitueret.". Vainio either did n08 find algae in his specimens or if he did, he attributed them to another lichen mixed in. However, algae are ab- undant in both American and EurOpean collections. M. leucOplaca could be included in Microthelia, which has been treated by lichen- ologists to allow inclusion of species which occasionally have more than one septum in the spores. These species are saxicolous and are never more than 3-septate. Microthelia has been taken over by mycoIOgists and redefined on the basis of the cellular characteristics of the pseudothecial wall. I have not tried to make use of this character as yet. The distinction of Molestia from Microthelia is admittedly at present based primarily on the multiseptate spores and general "feel". Molestia leucOplaca (Wallr.) comb. nov. :grrucaria leucoplaca Wallr. Flora Crypt. Germ. 3: 299. 1831. Eigenula leucOplaca (Wallr.) Koerh. Syst. Lich. Germ. 365. 1855. Lgptosphaeria leucoplaca (Wallr.) Vainio, icta Soc. Fauna Flora Penn. #9: 143. 1921. Eigenula leucoplaca var. pluriloculata Fink, Minn. Bot. Stud. 2: 709. 1902. The basionym of this species is often given as Verrucaria farrea Ach. (Math. Lich. 115. 1803.) but Vainio (1921) has examined the type of this species and states that it has nothing to do with M. leuco- 23.29.;- Thallus endophloeodal, white to pale gray, smooth. Trentepohlia abundant, very well—deveIOped, cells 9-12P in diameter, forming short 57 chains. Pseudothecia scattered, black to dark brown, shiny or appear- ing pruinose due to the remains of the covering bark layer, hemi- spherical to subglobose, c. 0.3mm in diameter, usually depressed in the center; pore in the depression, large, up to 50p across; initially just under bark surface but finally one third to one half immersed. Pseudothecial wall entire, brown above, 60-90P thick, colorless below, thinner below, appearing dimidiate. Nucleus IKI+ dark blue-green 'becoming orange. Paraphysoids c. 1? thick, only slightly branched and anastamosed. Asci long cylindric, without a dimple or with a very large one formed by a slight thickening of the wall at the tip; no reaction with KOH; 95-100 1 12?. Spores eight in the ascus, ob— 1iquely uniseriate, brown, with the terminal cells somewhat lighter in color, fusiform, 5-6-septate when mature, often slightly constrict- ed at the septa, the central cells largest decreasing in size toward the ends, lumina cylindrical and the walls of uniform thickness; no gelatinous outer layer present; 18-24 x 5-9P. The white thallus, shiny black pseudothecia depressed around the pore and brown spores with more than three septa are the diag- nostic characters of this species. The recognition of this species in North America has been com- plicated by the presence of a non-lichenized fungus found only on the living bark of Pogulus tremuloides. This fungus apparently does not occur in Europe. American collections under the name Ezrenula' leucoplaca in herbaria seem to be split about evenly between M. leuco- placa and the fungus. The fungus is easily distinguished by its smaller ascocarps, lack of IKI reaction and constantly B-septate spores with a thick gelatinous outer layer. These 3-septate spores caused Fink (1902), thinking that the EurOpean species was constantly 58 5-septate, to describe a variety pluriloculata for the 5-6-septate lichen. However, EurOpean material is also more than 3-septate and identical to ours, although EurOpean.authors tended to gloss over the extra septa in order to include it in Eigenula.‘ In the Great Lakes region M. leucoplaca has been collected on Ulmus americana, Acer saccharum, Tilig, POEulus, Quercus and Fraxinus. The only collection seen from North America outside of the Great Lakes is from New Hampshire. The range of the fungus, while somewhat more western and northern, overlaps M. leucoplaca, especially in north- ern Michigan and Minnesota. Exsiccati examined. Cummings I. 194 (MSG, WIS); Cummings II. 126 (HIGH); Krypt. Vind. 215} (FH); Rel. Far. #53 (MICH, WIS); Stenh. 179 (MICH). Specimens seen. CANADA ONTARIO: Carleton County, Ottawa, Macoun, 18 Apr. 1896 (US). QUEBEC: county unknown, Montmorency, Macoun, 7 Jan. 1905 (MIGH). UNITED STATES IOWA: Fayette County, Fayette, {325, June 189% (MICH, Thomson, WIS), {£35, July 189% (MICH, Thomson, WIS), £23k, 17 Aug. 1895 (MICH). 2225' h Dec. 1895 (MICH, MSG, WIS). MICHIGAN: Cheboygan County, Pine Grove campground east of Wolverine, Harris Z§Q_(MSC); Iosco County, Iargo Springs on the Au Sable River north of Tawas, Harris 827 (MSG); Menominee County, Big Cedar River campground north of Cedar River, Harris 222, Z22, Z22, 128,4222, 2&2 (MSG); Ontonagon County, Porcupine Mtns.‘fi mile east of parking lot, Harris 6 4, 617, 682, 690 (MSG). MINNESOTA: county unknown, Auburndale, Fink, 6 Sept. 1902 (MICH), 59 Minneapolis, §$E§.l2221 11 July 1896 (MICH), Pork Bay, Eink, 25 Aug. 1902 (MIGH); Beltrami County, Bemidji, Eggk_222, 9 July 1900 (MICH), Red Lake, £23§p£912, 3 Aug. 1900 (MICH); Koochiching County, Koochi- ching, {132.223, 26 July 1901 (MICE); Otter Tail County, Housing, 2125'221, 29 June 1900 (MIGH); Pennington County, Thief River Falls, {$35l282, 18 July 1900, §g;, 19 July 1900 (MICE); Yellow Medicine County, Granite Falls, 3335, 15 July 1899 (HIGH). NEW HAMPSHIRE: Carroll County, Chocorua, Farlow, May 1906 (MICH, WIS). For the sake of completeness and clarity I will include here a description of the fungus which has been confused with M. leuCOplaca. R. A. Shoemaker (in litt.) has suggested that the fungus is an un- described species of Melanomma. "Thallus" white, powdery, apparently consisting entirely of dead bark cells; occuring exclusively on the living bark of Pepulus tremuloides. Pseudothecia scattered, black, hemispherical, c. 0.1mm in diameter, mostly completely immersed in the loosened bark cells. Pseudothecial walls entire, brown, 25-h5P thick. Nucleus IKIe. Paraphysoids c. 1P thick, much branched and anastamosed. Asci cylindrical to elevate-cylindrical, tip not much thickened, dimple not evident, tip expanding in KOH and a small dimple becoming clearly evident; 60-80 x 12-15P. Spores eight in the ascus, biseriate, brown, fusiform, constantly 3-septate, usually stromdy constricted only at the median septum, spore halves equal or more commonly unequal, with the upper half broader; heavy gelatinous layer evident in KOH; 15-2# x 6-8p. Apparently common on Populus tremuloides in the northwestern Great Lakes region. Also common in the Black Hills. 60 Specimens seen. UNITED STATES MICHIGAN: Alger County, Rock River, Lowe 2202 2229 (MICE); Cheboygan County, Pine Grove campground east of Wolverine, Harris 222 (MSG); Iron County, junction of USF hwy. 139 and Co. 657 south of Kenton, Harris 222 (MSG); Keweenaw County, Isle Royale Nat. Park, Hay Bay, 2222122; (MICH), McCargo Cove, 22:2 222,(MICH), Rock Harbor, 2222 22; (MICE); Menominee County, Big Cedar River campground north of Cedar River, Harris 751 (MSG). MINNESOTA: Cook County, Grand Marais, £222, 31 July 1902 (MICE), Grand Portage Island, [225, 21 June 1897, Rose Lake, £225,222, 27 June 1897 (MICH, MSG, Thomson); Otter Tail County, Leaf Hills, 2225 222, 27 June 1900 (MICH); Pennington County, Thief River Falls, 2222 221, 17 July 1900 (MICE); St. Louis County, Harding, {$22 22;, 19 Aug. 1901 (MICH). POLIBLASTIOPSIS Zahlbr. Engler & Prantl, Natuerl. Pflanzenfam. I. Teil, Abt. I‘: 67. 1903. Ascocarp a pseudothecium, spores hyaline, submuriform to muriform. PolyblastIOpsis traditionally has been distinguished from MicrOglaena Koerb. on the basis of simple paraphyses and proto- coccoid algae in Microglaena and branched and anastamosed para- physoids and Trentepohlia in PolyblastiOpsis. Morgan-Jones and Swinscow (1965) have shown that Microglaena actually has branched and anastamosed paraphysoids and bitunicate asci as in Polyblasti- 22222. They have transferred one species of PolyblastiOpsis, P. sericea Massa1., to Microglaena but have not yet suggested that 61 the two genera should be combined. Since P. sericea seems to be quite typical of the genus, this combination would seem to be indi- cated. The combined genus would have to bear the name MicrOglaena as it is by far the oldest. Keeping the genera separate on the basis of the phycobiont does not seem valid if they are similar in all other respects. I have made no studies of Microglaena and prefer, there- fore, to keep the genera separate since Morgan-Jones and Swinscow have not seen fit to combine the genera. PolyblastiOpsis fallaciosa (Arn.) Zahlbr. Engler A Prantl, Natuerl. Pflanzenfam. I. Teil, Abt. I‘: 65. 1903. Polyblastia fallaciosa (Stizenb. ex Arn.) Arn. Flora #6: 604. 1865. This species has been something of a problem nomenclaturally. Arnold and some other European workers used the name Arthqpyrenia punctiformis fallax for collections of this species before the spores were examined. When Arnold (1863) discovered the spores were muri- fora and not l-septate as in Arthgpyrenia, he corrected his misiden- tification and described a new muriform-spored species, Polyblastia fallaciosa. American workers in the late 1800's and early 1900's were apparently unaware of Arnold's correction and continued to perpetuate the original error. As a result American herbarium mater- ial under the illegitimate combination PolyblastiOpsis ggiigx.or the legitimate one, Arthopyrenia fallax tends to be a mixture of both genera. Fink (1935) recognized P. fallax as distinct from P. falla- 22222 but they are nomenclaturally and taxonomically the same. Description. "Thallus" light gray to white or not at all evident when on Betula. Algae absent or a few scattered protococcoid cells around the pseudothecia. Pseudothecia scattered, black, sometimes 62 shiny, hemispherical especially when young but in age flat-t0pped or depressed in the center, 0.2-0.3mm in diameter; when on Betula with a dark elliptical hyphal ring. Pseudothecial wall entire, brown above, 20-30p thick, brown to hyaline below, often appearing dimi- diate, thinner below. Nucleus IKI-. Paraphysoids thick, 1.5-2u, septate with short cells, much branched and anastamosed, occasion- ally almost parenchymatous in appearance. Asci clavate to cylind- rical, without a dimple and only slightly thickened at the tip or with a shallow to medium flat-tepped dimple and the tip thicker; tip expanding in KOH to form a small thimble-shaped dimple in the much expanded tip; 50-102 x 15'25P° Spores initially eight in the ascus, leZ-(k) spores commonly not deveIOping but usually visible in the ascus, irregularly arranged, hyaline, oblongoelliptical, submuriform with 5-7 transverse septa and 1-2 longitudinal septa, usually con- stricted at the transverse septa, more so at the median septum, one half sometimes broader; usually with a thick gelatinous outer layer; 17-25 I 7-10Po Pycnidia usually present, black, 0.05-0.1mm in dia- meter. Microconidia c. 10 x 1P’ hyaline, simple, rod-shaped. Macro- conidia have also been attributed to this species which are brown, Boseptate, 9-12 x B-kp. Polyblastiopsis fallaciosa occurs commonly on Quercus 2222, Betula pgpyrifera and young geer saccharum and rarely on 2252222 cinerea and Fraxinus. Verified specimens range from Tennessee north to Massachusetts and around the Great Lakes west to Minnesota and Iowa. Specimens seen. CANADA ONTARIO: Algoma District, tundra strip along Lake Superior north of Batchawana, Imshaug 22922 (MSG), along Sand River Rapids, 63 Lake Superior Prov. Park, Imshaug 26269 (MSG), Kinny Lake, Lake Superior Prov. Park, gaggig‘égg (MSG); Simcoe County, Blue Mtns. between Singhampton and Duntroon on hwy. 24, Imshaug gégflé, EQQZI, g§221,(nsc). UNITED STATES ILLINOIS: Cook County, Calkins 22g7(wxs). IOWA: Fayette County, Fayette, §1g§,.kpr. 1898 (MICE, MSG). MICHIGAH: Allegan County, State game area southeast of Perch Lake, Barrie #00, #01 (MSG); Alger County, Kingston Lake, Harris §g§‘(MSC); Berrien County, Warren Woods, Harris Eli, glé (MSG); Charlevoix County, Fox Lake, Beaver Island, Imshaug EZEZé (MSG), old apple orchard, Beaver Island, Imshaug gzgég (MSG); Cheboygan County, Pine Grove campground east of Wolverine, Harris 775, ZZé, ZZZ, zzg, Zég (MSG); Emmet County, Straits west of Mackinaw City, Imshaug §2§é2,(MSC); Ingham County, Sanford Woodlot, East Lansing, Harris fill, 2;§_(MSC); Iosco County, Iargo Springs on in Sable River north of Tawae, Harris 830, QZQ, Q22’(MSC); Iron County, junction of USF 139 and Co. 657 south of Kenton, Harris le (MSG); Kalamazoo County, KeIIOg Forest north of Augusta, Harris lfig, l6, #1 , 2&2 (MSG); Keweenaw County, Fort Wilkins, Thomson 2998, 3006 (Thomson, WIS); Mackinac County, Mackinac Island, Sugar Loaf Rock, Imshaug 272708, 2221;, 22212 (MSG), Leslie Avenue, Imshaug 27289 (MSG); Menominee County, Big Cedar River campground north of Cedar River, Harris 239, Zfi§, 222, 22é_(MSC); Ontonagon County, Porcupine Mtns. ‘% mile east of parking lot, Harris 671, 672 (MSG); Washtenaw County, Crooked Lake, Harris I, Q, Q, §§Q, ggz, §§2 (MSG). MINNESOTA: Beltrami County, Bemidji, EEBE £29, 12 July 1900 (MICE); Cook County, Grand Marais, Pink, 12 Aug. 1902 (MICE).' 6h 28 July 1902 (US), Grand Portage, Fink 2Q, 21 June 1897 (MICE); Grand Portage Island, Fink fig, 23 June 1897 (MICE), Gunflint, Fink 276, 30 June 1897 (US), Misquah Hills, Fink 20 , 5 July 1897 (MICE). WISCONSIN: Oneida County, near Big Stone Lake, Culberson 1604 (WIS). Polyblastiopsis lactea (Massal.) Zahlbr. Engler & Prantl, Natuerl. Pflanzenfam. I. Teil, Abt. I‘: 65. 1903. Blastodesmia lactea Massal. Ricerch. Anton. Lich. 181. 1852. "Thallus" gray to whitish. Algae not found. Pseudothecia scattered, black, slightly shiny, hemispherical to conical, O.)- O.4mm in dimeter, initially immersed but finally appearing to be raised on a small mound in age, surrounded by a dark hyphal ring. Pseudothecial wall entire, brown, thicker above, c. 50p thick, thin- ner below, lS-ZOP. Nucleus IKI-. Paraphysoids c. l p thick, septate, branched and anastamosed. Asci cylindrical to clavate, with a broad rounded dimple in the tip which is little thicker than the walls; tip expanding in KOH to form a long narrow dimple in the tip which is now up to 10p thick. Spores tour in the ascus, occasionally fewer, irregularly arranged, hyaline, long elliptical, muriform, with 6-8 transverse septa and 2-3 longitudinal septa, slightly constricted at the septa, especially the median one; thick gelatinous outer layer evident in KOH; 35-45 x lZ-lzp. The only verified collection of P. lactea is on Fraxinue from Michigan. All of the other North American material of this EurOpean species under this name in herbaria has turned out to be P. fallaciosa. PolyhlaetioPsis lactea is apparently the only species in the genus with four spores to the ascus. All others are initially two or eight spores in the ascus. The initially eight-spored P. fallaciosa 65 has a strong tendency for spore abortion. It is commonly found with 6-7 spores and rarely with as few as four mature spores in the ascus. This, for the most part, accounts for the previous records of P. lactea in North America. The spores of P. fallaciosa are smaller and if asci with less than eight spores are carefully examined, the remains of the undeveloPed spores can be found. Massalongo (1852) allowed up to six spores per ascus in this species but only a single EurOpean collection has shown more than four spores. Exsiccati examined. Arn. 56% (FE); Koerb. 381 (US); Lojka 149 (FH); Hassal. 1&3 (FH); Rabenh. 201 (FR). Specimens seen. UNITED STATES MICHIGAN: Emmet County, Wilderness State Park, Harris 816 (MSG). Polyblastiopsis meridionalis Zahlbr. Annal. Naturh. Hofmus. Wien. 23: 223. 1909. "Thallus" pale gray. No algae found. Pseudothecia scattered, bluish- or greenish-black, circular to elliptical in outline, very much flattened and occasionally somewhat sunken in the center. Pseudothecial wall dimidiate, greenish—black or greenish-brown, 10-25? thick. Nucleus IKI-. Paraphysoids not abundant, c. In thick, septate, branched and anastamosed. Asci cylindrical, wall and tip uniform in thickness, e. 2P’ tip often appearing somewhat truncate; tip expanding slightly in KOH with the formation of a small dimple; 65-75 x 12'15P' Spores eight in the ascus, uniseriate. to biseriate, hyaline, submuriform, oblong-elliptical, with 3-(6) transverse septa and l-(Z) longitudinal septa, constricted at the septa especially the median one; gelatinous outer layer present; 21-25 x B-QP. 66 This species, which is distinctive in its flattened green pseudothecia, has not been recognized in North America previously although all of the specimens seen are from old herbarium material. The only collection from the Great Lakes region is from Northern Ohio. Usually found on smooth bark. Exsiccati examined. Krypt. Vind. l6h6 (Isotype?, FR, US); Merrill II. 93 (sub Eycoporum pyrenocarpum, MICE, MSG). Specimens seen. UNITED STATES IOWA: Black Hawk County, La Porte, £335, 21 Feb. 1896 (MICK); Fayette County, Fayette, {$35, Aug. 1895 (MICE). MAINE: Oxford County, Hartford, Parlin 13951 (Thomson). MISSOURI: Saline County, Emma, Demetrio, 6 July 1898 (MICE). OHIO: Ashtabula County, Orwell, Bogge, 1899 (MIGH, use). PYRENULACEAE Zahlbr. Engler & Prantl, Natuerl. Pflanzenfam. 1. Teil, Abt. I‘: 62. 1903. This is the earliest use of the name Pyrenulaceae which I have verified. Further research may bring to light earlier uses of the name e PLAGIOCARPA gen. nov. Ascocarp a perithecium surrounded by a stromatic shell. Ostiole borne on one side of the perithecium at the end of a short neck, spores hyaline, endospore very slightly thickened, transversally septate. Type species: Xerrucaria hyalospora Nyl. Apparently most closely related to Pyrenula, especially in ascocarp structure but differing in the hyaline spores with little 6? endospore thickening. The spores are quite similar in appearance to Arthopyrenia and Pleurotrema which differ in having pseudothecia. The type species is the only species known. It was consid- ered by Fink (1935) to be an Arthopyrenia. The material in his herbarium from Iowa and Minnesota under this name is, in fact, either A. quinqueseptata or A. quisquiliae. Even if Fink had not misinterpreted the species, Plagiocarpa hyalospora. could nOt be included in ArthOpyrenia since that genus is ascolocular in devel- 0pment while Plggiocarpa is ascohymenial. Also the exit pore in ArthOpyrenia is vertical not on one side. Degelius (l9hl) collected this species in Tennessee and noting the ostiole on the side of the perithecium placed it in the genus Pleurotrema, describing a new species, P. solivagum. This disposition is also incorrect since all all of the species of Pleurotrgma examined, including the type of the genus, are ascolocular although they do have their exit pores on the side of the pseudothecium. The type of asci and paraphyses seems to exclude it from any close relationship with Tryphethelium, which represents one ascohymenial line. The other ascohymenial pyrenolichen line, as represented by Pygenula, seems at present, to have Plagiocarpa‘s nearest relatives. Both Pyrenula and Plagia- gggpg” have a stromatic shell surrounding a well-developed exciple and to have similar paraphyses and asci. Plagiocarpa differs from Pygenula in having a tilted ostiole and in having hyaline spores with a relatively thin endospore, although the endospore is better develoPed than in the ascolocular genera. For these reasons a new genus is required. 68 Plagiocarpa hyalospora (Nyl.) comb. nov. Verrucaria hyalospora Nyl. Mem. Soc. Acad. Maine-et-Loire 4: 48. 1858. (Expos. Syn. Pyrenocarp.). Mem. Soc. Imper. Sci. Nat. Gherb. 5: 337, 1857 (nom. nud.). Pyrenula hyalosppra (Nyl.) Tuck. Gen. Lich. 273. 1872. ArthOpyrenia hyalospora (Nyl.) Fink, Lich. Flora 50. 1935. Pleurotrema solivagum Degel. Ark. Bot. 30A(3): 21. 1941. Nylander stated in his original description that his material came from Tuckerman's herbarium. Tuckerman (1872) says that Nylander found this species mixed with some other material which Tuckerman had sent. I have examined all of Tuckerman's material of this species but have not seen any material from Nylander's herbarium. Therefore no type has been selected. All of Tuckerman's material is uniform and correctly determined. An isotype of Pleurotrema Bolivagun (US) has been examined. Degelius listed the substrate as 22353, it seems to me to be Acer saccharum. Description. Thallus endoPhloeodal, pale gray or whitish, KOH+ yellow becoming yellow-brown, smooth. Trentepohlia usually abundant and well-develoPed, forming short chains. Ascocarps scattered, dark brown, not shiny, subglobose, initially completely immersed in the bark but commonly almost entirely free in age. Oatiole displaced to one side, not vertical, with a short neck; occasionally vertical or not projecting. Involucrellum entire, dark brown, c. 80u thick above, thinner below, 20-40u. Exciple thin c. lO-ZOu, pale brown to almost colorless, often entirely free from the involucrellum and shrinking away from it on drying except at the ostiole. Nucleus IKI+ greenish-blue becoming red-orange if the iodine concentration is high enough. Paraphyses septate, very rarely branched but 69 apparently never anastamosed. Asci cylindrical or somewhat obclav- ate, not much thickened at the tip, dimple present, often cross- shaped in Optical section; tip expanding in KOH for a distance of 20-30n back from the end, forming a narrow canal; 65-90 x 20p. Spores eight in the ascus, roughly biseriate to irregularly arranged, hyaline, ellipsoid, 3-septate, with the end cells smaller than the median cells, not constricted at the septa, endospore slightly thick- ened, lumina spherical to almost cylindrical; no gelatinous outer layer; 18-25 x 9-llr. Plagiocarpa hyalospora is most frequently found on Acer saccharum but also occurs on gilig americana, Betula $2222.39d ngyg. I have seen a single collection on lignum. Plagiocarpa is quite common around the Great Lakes occuring east to New England and south- to Tennessee. A single collection with slightly smaller ascocarps and spores, collected by B. A. Imshaug in the West Indies, is very tentatively included. Exsiccati examined. Lojka ZOO (MICE); Merrill I. 102 (MICH, US). Specimens seen. CANADA ONTARIO: Carleton County, Ottawa, Macoun, 18 Apr. 1896 (US); Hastings County, Belleville, Macoun 5,41, 1894 (FR); Middlesex County, London, Drummond 28, 1869 (FE); Nipissing District, Lookout trail, Algonquin Prov. Park, Imshaug 26822 (MSG); Parry Sound District, Magnetawan, glitt, 29 July 1920 (US). UNITED STATES * MAINE: Knox County, Rockland, Merrill, 15 Aug. 1909 (MICE, MSG). MARYLAND: Baltimore County, Catonsville, Eliiifieé’ 14 Jan. 1911 (US). 70 MASSACHUSETTS: Bristol County, New Bedford, Willey, 1868 (FH), Willey, Dec. 1885 (US); Hampshire County, Amherst, Tuckerman, 1857 (PH). MICHIGAN: Alger County, Au Train Lake, A. H. Smith 2103 (HIGH), Kingston Lake, Harris 618, 662 (MSG); Berrien County, Warren Woods, Harris 216A, 212_(MSC); Cheboygan County, gorge near Douglas Lake, Nichols, June-Aug. 1923 (HIGH); Ingham County, East Lansing, Harris filQ (MSG); Iosco County, Iargo Springs on Au Sable River north of Tawas, Harris 829, 836, 8 7, 858, 840, 841, 842, 84; (MSG); Iron County, Teepee Lake south of Kenton, Harris 624, 696, 727, 728, 750A (MSG). NEW HAMPSHIRE: County unknown, White ths., Tuckerman, 1847 (FE); Grafton County, Warren, Willey, 1882 (US). NEW YORK: County unknown, Trenton Falls, Willey, 1880 (US); Genessee County, Bergen swamp, Muenscher & Brown, 17 Nov. 1946 (HIGH); Niagara County, Goat Island, Tuckerman, Aug. 1848 (FE); Otsego County, Richfield Springs, Willey, 1884 (US). OHIO: Franklin County, Georgesville, m, 12 May 1894 (HIGH). SOUTH CAROLINA: Chester County, Chester, EEEEE! 30 Jan. 1886 (MICH, use). m. 9 Apr. 1881+ (us). ’ TENNESSEE: Sevier County, Cherokee Orchard, Degelius, 11 Sept. 1939 (US). WISCONSIN: Sawyer County, Flambeau River State Forest, H512 £921 (WIS). WEST INDIES ST. LUCIA: Quarter of Soufriere, southwest slepe of Gros Piton, south of Soufriere, Imshaug 29652 (MSG). 71 PYRENULA Massal. Ricerch. Anton. Lich. 162. 1852. Ascocarp a perithecium surrounded by a stromatic shell, spores brown, transversally septate, endospore much thickened so that the spore lumina are lenticular or spherical. gygenula Massal. was published as an emendation of Pyrenula Ach. (Lich. Univ. 64, 314. 1810) but does not include any of Acharius' original species with which Eygenula Ach. would have to be typified. None of these original species were even pyrenocarpous. Therefore, it would seem best to conserve Pyrenula Massal. against Pygenula Ach. Pyrenula neglecta sp. nov. Thallus endophloeodal, yellowish-green to olive-green, smooth, often shiny; upper layer c. 0.1mm thick consisting of bark cells intermingled with hyphae. Trentepohlia abundant, very well-deveIOped, forming a very definite layer. Ascocarps dark brown to black, dull to shiny, scattered or occasionally clustered and fused, O.4~O.6mm in diameter, initially covered by bark and thallus but finally almost completely emergent. Ostiole evident, c. 4GP across, flush with the ascocarp surface or often raised in a small papilla. Involucrellum dark brown in section, e. 50-60u thick, mostly dimidiate, reaching about to the base of the perithecium, often containing bark cells. Exciple brown, 10-15? thick, often separating from involucrellum and bark. Nucleus containing an abundance of oil draplets, IKI+ blue-green at the base. Paraphyses simple, c. In thick. Asci cylindrical to somewhat clavate, 70-85 x 15—18P. Spores eight in the ascus, mostly uniseriate, pale olive-brown, 3-septate, not 72 constricted at the septa; lumina oval in Optical section, the ter- minal lumina smaller and pressed directly against the outer hyaline spore wall; usually a least one half as wide as long, 1A.20 x 7-12P. This widespread species has previously been considered to be identical with the European species, Pyrenula nitida (Weig.) Ach. On careful examination the two species are quite different. The most obvious differences are summed up in the following chart. Eyrenula nitida Ostiole tends to be depressed Involucrellum contains crystals Brown layer around base of nucleus KOH+ red Nucleus IKI- Nucleus with very little oil, if any Spore width/length ratio mostly less than 1:2 Terminal lumina separated from outer spore wall by endospore at tip Lamina usually angular, diamond-shaped in pptical section gyrenula neglecta Ostiole tends to be raised Involucrellum without crystals No KOH+ layer around base of nucleus nucleus IKI+ bluish-green at the base Nucleus with much oil Spore width/length ratio mostly more than 1:2 Terminal lumina directly against outer spore wall at tip Lumina not angular, oval in optical section Several other species from Southern Ohio and further south have also been called gyrenula nitida but none of them are comparable to the EurOpean species, which apparently does not occur in North Amer- ica. The fact that American lichenologists have been confused by Ezrenula neglecta is indicated by the fact that much material of this species is also found under gyrenula laevigata or its synonym, P.+glabrata. 73 Ezrenula neglecta is almost always found on hard barks of such trees as Faggs grandifolia, Quercus borealis, g. veutina or Acer saccharum. On the basis of specimens so far examined the species ranges from coast to coast along the U.S.-Canada border, south in the Eastern United States to Florida, although it seems to be rare south of Washington, D.C. Specimens collected by H.A. Imshaug in the Canary Islands may belong to this species also. Exsiccati examined. Cummings I. 150 (HIGH, MSG, WIS); Macoun I. 68 (MICE, WIS); Macoun II. 228 (MICE, Thomson); Merrill I. 186 (HIGH); Rel. Far. 45% (WIS); Rel. Tuck. 131 (MICK, Thomson, WIS). Specimens seen. CANADA ONTARIO: Carleton County, Ottawa, Macoun, 26 Apr. 1891 (MICE, WIS), Hacoun,6 Sept. 1891 (MICE, Thomson); Simcoe County, Blue Mtns. between Singhampton and Duntroon on hwy. 24, Inshaug 26226, gégéé (MSG). UNITED STATES ILLINOIS: Cook County, Calkins £88 (Thomson). IOWA: Clayton County, Pikes Peak, bluffs along Mississippi River near McGregor, Imshaug §Z§2§ (MSG); Delaware County, Maquoketa Diver valley, Backbone State Park north of Dundee, Imshaug 27928 (MSG): Fayette County, Fayette, £13k, Apr. 1894 (MICE); Jackson County, Maquoketa Caves Park northwest of Maquoketa, Imshaug 280§§, g§929,(msc). MICHIGAN: Alger County, Au Train, L233 ;2_2, izgz (MICE); Bennie County, Bennie State Park, Imshaug gzfilg (MSG); Charlevoix County, Beaver Island, old apple orchard, Imshaug 27588, Sandy Bay, Imshaug‘gZéfiz (use); Cheboygan County, Bois Blane Island, lgghggg 2292 (MSG), Colonial Point, Douglas Lake, Nichols, JuneaAug. 1923 (MICK), 74 gorge near Douglas Lake, Imshaug 21}? (NSC); Crawford County, Hart- wick Pines State Park, Imshaug 2158: (MSG); Emmet County, 4 miles south of Mackinaw City, §1ggk, 18 July 1957 (WIS); Boughton County, Lake Superior near Calumet, Imshaug 2982; (MSC); Luce County, Upper Tahquamenon Falls, Imshaug 12829 (MSG); Mackinac County, Mackinac Island, Fort Holmes, Imshaug gzggg (MSG). WISCONSIN: Door County, Valmy at Cave Point, Culberson 2§1Q (WIS); Grant County, T6N, R6W, sect. l9, §g1g 1122 (WIS); Green Lake County, Tth, 212E, sect. 26, £313 1918 (WIS); Sauk County, Baraboo Range, Thomson 1291 (Thomson, WIS); Vernon County, T13N, R6W, sect. 32, Hale 1226 (WIS); Washburn County, near Long Lake, Culberson £229 ms). gyrenula agawae sp. nov. Thallus endophloeodal, indicated only by greenish color under upper layer of bark or by a grayish blotch in age. Trentepohlia abundant, well-developed. Ascocarps scattered, dark brown, usually covered by thin layer of bark, flattened subglobose, 0.2-0.4mm in diameter. Involucrellum mostly entire, continuous with clypeus which extends c. 0.1mm beyond the involucrellum, containing numerous bark cells. Exciple brown, c. 20-50? thick. Nucleus IKI-, without oil. Paraphyses unbranched, septate, c. In thick. Asci cylindrical- clavate to clavate, 50.70 x lO-lSy; unchanged in KOH. Spores eight in the ascus, uniseriate to biseriate, gray-brown, ellipsoid to lemon-shaped, B-septate, not constricted at the septa; lumina ellip- soid to somewhat diamond-shaped in Optical section, terminal lumina not directly against outer spore sell but separated from it by a layer of endospore; 17-20 x 9'10P’ 75 This species is known only from a single collection on Betula 12135. It differs from our only other northern species of Pyrenula, P. neglecta, in having a poorly deve10ped thallus, distinct clypeus, lack of oil in the nucleus, IKI- nucleus and position of the terminal spore lumina. Specimens seen. CANADA ONTARIO: Algoma District, Lake Superior Prov. Park, % mile north of Doc Grieg Lake, Harris 262 (MSG). TRYPHETHELIACEAE Zahlbr. Engler & Prantl, Natuerl. Pflanzenfam. I. Teil, Abt. I‘: 69. 1903. This is the earliest use of the name which I have verified» Further research may bring to light earlier uses of the name. TRYPHETHELIUM Spreng Anleitung zur Kenntn. der Gewaechse, 3: 309. 1805. (conserved over Bathelium Ach. Method. Lich. 111. 1803) Tryphethelium virens Tuck. in W. Darl. F1. Cest. ed. 3. 453. 1853. Description modified from Johnson (1959). Thallus endoploeodal, smooth, greenish, yellowish-gray or yellow-brown to orange-brown; upper layer of fused hyphae and bark cells, cortex-like, 45-90p thick. Trentepohlia abundant, well-develOped, in a layer 15-23” thick. Perithecia aggregated and imbedded in a pseudostroma up to 7mm in length, slightly elevated, more-or-less concolorous with the thallus (especially when young) but becoming lighter or darker with 76 age; outer layer of bark cells up to 55? thick, with perithecial initials forming below this layer, raising the bark layer as the perithecia enlarge, sometimes altering the bark cells to a blackish color or interwoven hyphae becoming carbonaceous. Perithecia 0.4- 0.6mm high by 0.2-0.5mm across. Ostioles seen from above as black dots c. 0.1mm in diameter. Exciple carbonaceous, 30-60? thick. Paraphyses c. 1P thick, anastamosed, forming a broad net. Asci clavate to clavate-cylindrical, 100—125 x 15-l7r. Spores eight in the ascus, irregularly biseriate, hyaline, 7-9-septate, not constrict- ed at the septa; lumina elliptical or diamond-shaped in Optical section, 38-52 x 7-10P° Pycnidia large, embedded in stroma, labyrin- thiform, with long thin conidiOphores; microconidia hyaline, rod- like, c. 5 x In. In the Great Lakes region the usual substrate Of this lichen is [3523 but it has been also found on Carpinus. The species is of special interest as it is the only repre- sentative of a very large trOpical group,which is highly developed along stromatic lines. It apparently reaches its northern limit at about the Straits Of Mackinac, east to Vermont and New Hampshire, west to Wisconsin, reaching its southern limit in Oklahoma, Missis- sippi and Florida. Exsiccati examined. Cummings I. 253 (MSG); Cummings II. 193 (MICH); Merrill I. 69 (HIGH); Rel. Far. 456 (MICE); Zahlbr. 231 (MICE). Specimens seen. CANADA ONTARIO: Parry Sound District, Indian Docks, Parry Sound, Gain 26671 (WIS); Simcoe County, Blue Mtns. between Singhampton and Duntroon on hwy. 24, Imshaug 2627 , 26992 (MSG). 77 UNITED STATE MICHIGAN: Benzie County, Bennie State Park, Bayliss 41 (MSG); Cheboygan County, Pine Grove campground east of Wolverine, Harris 282, 783 (MSG), Douglas Lake, Nichols, June-Aug. 1923 (MICE); Eaton County, near Vermontville, Imshaug 12§2_, 22122 (MSG); Ingham County, Baker Woodlot, East Lansing, Harris fig, 22 (MSG). OHIO: Lake County, Painesville, {121, £42, 6 Oct. 1915 (MICE). WISCONSIN: Door County, near Jacksonport, Culberson 2709, 2912 (WIS). SUMMARY This survey of the corticolous pyrenocarpous lichens in the Great Lakes region treats nine genera and 23 species. Spores are illustrated for all the species treated. .Their morphology and ecolOgy are discussed briefly. Molestia and Plagiocarpa are described as genera new to science. Arthopyrenia guisquiliae, A. thomsonii, A. willeyana, Pyrenula a awae, and E: neglecta are described as species new to science. Arthopyrenia myricae.(Nyl.) Zahlbr., Leptorhaphis atomaria (Ach.) Snatala, L. parameca (Massal.) Koerb., and Polyblastiopsis meridionalisfigkyfireported as new to North America. Microthelia wallrothii (Hepp) Rehs is reported as new to the Great Lakes region. Eyrenula nitida (Weig.) Ash. and ArthOpyrenia alba (Schrad.) Zahlbr. are excluded from the flora. The study was based on field work and examination of material from the Farlow Herbarium, Michigan State University, Smithsonian Institutuin, University of Michigan, University of Wisconsin and the personal herbarium of J. W. Thomson. 78 LITERATURE CITED Arnold, F. 1863. Die Lichenen des Fraenkischen Jura. Flora. 46: 601—604. . 1871. Die Lichenen des Fraenkischen Jura. Flora. 5E: 482-490e Butler, E. T. 1940. Studies in the Patellariaceae. MchIOgia. 32: 791-32}. Clements, F. E. 1909. The genera of fungi. Minneapolis. 227p. Curtis, J. T. 1959. The vegetation of Wisconsin. Madison. 657p. Degelius, G. 1940. Contributions to the lichen flora of North America. I. Lichens from Maine. Ark. Bot. 3OA(1): 1-62. . 1941. Contributions to the lichen flora of North America. II. The lichen flora of the Smokey Mountains. Ark. Bot. 301(3): 1-80. Fink, B. 1902. Contributions to a knowledge of the lichens of Minnesota. VI. Lichens of Northwestern Minnesota. Minn. Bot. Stud. 2: 657-709. . 1913. The nature and classification of lichens. II. The lichen and its algal host. Mycologia. 5: 97-166. . 1935. The lichen flora of the United States. Ann ”bore . x + 426Pe Fuckel, L. 1870. Symbolae mchlOgicae. Jahrb. Nassauischen Ver. Naturk. 23/2“: 1-459. Johnson, G. T. 1940. Contributions to the study of the Tryphetheliaceae. Ann. Mo. Bot. Gard. 27: 1-50. . The Tryphetheliaceae of Mississippi. Mycologia. 51: 74l~ 750. Keissler, K. 1938. Pyrenulaceae bis Mchporaceae, Coniocarpinae. In L. Rabenhorst, Kryptogamenflora, Leipzig. 9(1 Abt. 2 Teil) 346p. Koerber, G. W. 1855. Systems lichenum Germaniae. Breslau. xxxiv e 458p. Luttrell, E. S. 1965. Paraphysoids, pseudoparaphyses and apical paraphyses. Trans. Brit. Mycol. Soc. 48: 135-144. Magne, F. 1946. Anatomic et morphologie comparees des asques de quelques lichens. Rev. Bryol. Lichenol. 15: 203-209. 79 Magnusson, A. H. 1936. FUrteckning over Skandinaviens Vaxter, 4. Lavar. Massalongo, A. B. 1852. Ricerche sull' autonomia dei licheni crostosi. Verona. xiv + 207p. . 1854. Geneacaena lichenum. Verona. 24p. Morgan-Jones, G. and T. D. V. Swinscow. 11965. On the genus Microglaena Koerb. Lichenologist 3: 42-54. Mueller, E. and J. van Arx. 1962. Die Gattungen der didymosporen Pyrenomyceten. Beitr. Kryptog. Schweiz. 11(2): 1-922. Mueller, J. 1862. Principes de classification des lichens et enumeration des lichens de Geneva. Mem. Soc. Phys. Hist. Nat. Geneva. 16: 343-433. Nannfeldt, J.A. 1932. Studien ueber die Morphologie und Systematik der nicht-lichenisierten inOperculaten Discomyceten. Nova Acta Reg. Soc. Sci. Upsal. IV. 8(2): 1-368. Nylander, W. 1852. Observationes aliquot ad SynOpsim Lichenum Holmiensium. Bot. Notis. 1852: 175-180. . 1857. Enumeration generale de lichens, avec l'indication sommaire de leur distribution geographique. Mem. Soc. Imper. Sci. Nat. Cherbourg. 5: 85-146. . 1872. Addenda nova ad Lichenographium EurOpeam. Flora. 55: 353-365. Richardson, D. H. S. and G. Morgan-Jones. 1964. Studies on lichen asci. 1. The bitunicate type. Lichenologist. 2: 205-224. Riedl, H. 1962. Die Arten der Gattung Mchporellum Muell. Arg. sensu Zahlbr. Cat., nebst Bemerkungen sum System dothidealer Flechten. Sydowia, Ann. Mycol. ser. 2. 15: 257-287. . 1963. Vorstudien su einer Revision der Gattung ArthOpyrenia Mass. sensu ample, I. Sydowia, Ann. Myc. Ber. 2e 16: 263'274e Scheinpflug, H. 1958. Untersuchungen ueber die Gattung Didymosphaeria und einige verwandte Gattungen. Ber. Schweiz. Bot. Ges. 68: 325-385. Shoemaker, R. A. 1964. Staining asci and annellophores. Stain Technol. 39: 120-121. Singer, R. and I. J. Gamundi. 1963. Paraphyses. Taxon. 12: 147-150. Swinscow, T. D. V. 1962. Pyrenocarpous lichens. 3. The genus Porina in the British Isles. Lichenologist. 2: 6-56. 80 Swinscow, T. D. V. 1965. Pyrenocarpous lichens. 9. Notes on various species. Lichenologist. 3: 72-83. Tuckerman, B. 1872. Genera lichenum: an arrangement of North American lichens. Amherst. 283p. Vainio, E. 1921. LichenOgraphia Fennica. I. Pyrenolichenes. Acta Soc. Faun. F1. Fenn. 49: 1-274. Zahlbruckner, A. 1926. Lichens in Engler and Prantl, Die natuerlichen Pflanzenfamielen. ed. 2. 8: 61-270. 81 APPENDIX I. BASIC ECOLOGICAL DATA Stand # 1. Boreal forest, dominated by Abies balsamea, Betula lutea and B. papyrifera. Lake Superior Prov. Park,‘% mile north of Doc Grieg Lake, Algosa District, Ontario. Tree species 1 piggete; in 93. Lichens ,s ,. Microtheliazmiculs emu": 2? O\ A 2’ a 3 3 Abies balsamea 13 10 2 1 26 1 Acer rubrum 1 1 1 Betula lutea 3 5 9 16 '33 3 l 4 Betula papyrifera 1 6 13 20 Picea glauca 2 9 ‘t 15 Sorbus americana l 1 Thuja occidentalis 2 2 4 Total 26 30 3o 20 6 82 Stand # 2. Wet-Mesic Northern forest, an almost pure stand of Acer rubrua and ragga ggandifolig. East side of road just north of Kingston Lake, Alger County, Michigan. T49N, 315w, sect. 32. Tree specges gingeter in cm. Lichens l Planocarpe hyalospora Microthelie micula 3 3 § § 3 3 an «n IR :8 Abies balsanea 8 4 8 Acer rubrun 33 11 4 4 52 l 1 Betula lutea 1 l l 2 5 1 1 Fag-us grandifolia 6 6 12 24 2 2 Pinus strobus 3 1 2 5 Tsuga canadensis 5 5 Total 51 12 12 25 4 83 Stand # 3. Northern Mesic forest, nixed hardwoods without any obviously dominant tree species. Porcupine Mountains, )1 mile east of parking lot, Ontonagon County, Michigan. T5113, R43w, sect. 15. Tree species ' Diameter in ca. Lichens f 3' a 5 73 E "" - g a a 3 n .4 D e w I. n a 3 a i a , 3. a a g g 0‘ e o 3 § 3 g 3 S a a a. *3. 24 N M t3 :2 g .3 .3 8 Abies balssmea 4 4 8 Acer saccharum 5 7 7 1 20 2 3 4 9 Betula lutea l 1 Betula papyrifera 11 1 12 6 6 Frazinus sp. 1, 2 1 1 1 Populus tremuloides 1 2 1 1 1 Quercus borealis 7 1 9' 12 Thuja occidentalis 1 1 1 3 Tilia americana 4 7 5 16 Tsuga canadensis 1 7 5 4 17 Total 39 31 21 9 17 84 Stand # 4. Swamp forest, dominated by Acer rubrum, Ulmus and Fraxinus. Big Cedar River State Forest campground, Menominee County, Michigan. T36N, R25I, sect. 28. Tree gpecies Diameter in cm. Lichens s 3 3 a a O 3 s . . a a“? 5.5. é a 2.: .. a a Abies baleamea 2 2 Acer rubrum 26 10 6 42 Acer eaceharum 2 2 Betula lutea 1 1 Betula papyrifera 2 2 lraxisus sp. 22 l 23 1 1 Pinus strobus 1 1 6 6 Ulmus americana 2 11 7 7 27 Total 56 23 13 8 7- 85 Stand # 5. Northern Mesic forest, mostly Acer saccharum. Pine Grove State Forest campground, Cheboygan County, Michigan. $53!, 31W, sect. 16. Tree species pigmeter in cm. ) Lichens § .1 3 e 3 .2 § #4 '3 '3 . g. a "" "a e g .. a a :1 3 s g; 0‘ <3 '3 E: :8 4% r4 5 g g a 3 3 a a 3 r! «n a: 63 a. :3 S! ta Acer saccharum 31 20 1 4 56 49 49 Betula papyrifera 1 1 ragus grandifolia 3 2 10 15 2 2 POpnlus grandifolia 1 Prunus 3p. 2 2 h 1 1 Tilis usricana 6 8 2 2 18 1 1 Tsuga canadensis 1 1. 2 Ulmus americana 1 l 1 3 Total 40 35 7 17 55 86 Stand # 6. Northern Heaic forest, nixed hardwoods with no clear doninanta. Iargo Springs on tho Au Sable River north of fauna, Iosco County, Michisnn. Tahfl, R6E, soot. 26. Tree opooice Qianotqr in on. 2123223 FF 6 8 5'8 3 ,3 1... Hg, 5 §'3 §% '3 :‘1 EM: 3 5% § 3 g; Mg: 3”§§§E3 ééa g a 3 g a: :2 Acer rubrun 1 1 2 Acer saccherun 11 1 12 1 1 1 3 Betula lutea 1 2 1 k Irexinue up; 9 33 2 1 15 1 1 1 3 Ostrye virginiene 9 5 1* Piano otrohua 2 3 .5 Populul grandidontata 1. 1 Quércu-alba 1 1 2 Quorcus bioolor 1. l QuOrcnn boroelin 1 2 1 2 6 l 1 Thuje oocidontalia 1 1 Tiliaenorioene 8 7 lb 827 6 6 Tonga cnnedonsie 2 1 3 Ulla: enericnne 2 2 rotel U» 22 1019 13 87 Stand # 7. Dry Southern Hardwoods, dominated by guercus velutina and g. alba. Near relict prairie northeast of Newaygo, Neiaygo County. T13N, 311w, west side sect. 19. Tree species L Diameter in cm. Acer’rubrun Malus sp. Pinua atrobus Papulun grandi - dentata Prunus carotina Quercus alba Quercus borealis Quercua velutina 92M. *3 ééaéa 4 h 6‘ 1 1 1 2 1 k # 9 13 h # No lichens found. 2# 6 1 £2 33 h k 15 20 2 37 Total 531.1 #2 88 Stand # 8. Soutern Wet-Mesic forest. Sanford voodlot, East Lansing, Ingham County. Tim, Rll, eeot. 18. Ire: enciee W. Lichens 3 a 5 ‘2» a :3 a g 5 3 a g. 5 3 . E 8 5 3 aga asngss a é Si :5. as 3: r3 .2 a icer rubru 5 h 4 13 icer eaooharu 9 5 2 5 21 1 1 2 Caryn up. 1 1 raga. grandifolia 4 1 2 11 2 2 Italian. up. 1+ 2 k 10 1 1 Oetrya Virginiana 1 1 Platanua occidentalie 1 1 1 1 Queroue alba F 4 1 5 Quercue borealis 1 1 2 Tilia americana 8 9 1 1 19 Ulmus americana 6 6 3 3 15 3 3 Ema 38 28 12 22 . 9 89 Stand # 9. Southern Dry-Mesic forest, oak—hickory woods. North of Crooked Lake, Washtenaw County, Michigan. T18, R3E, sect. 6. Tree species Diameter in cm. Lichens’ 3 n o e 8 ”8 a mi cu o a .. é o 3 a 3 .. I: n «a 4'3 a cu m 21 «4 -fi 11 o = e a O\ <3 Ox 1 P! 'E: E; E: £3 1H 3 ‘3‘ ‘3 a 3 5 5 5 E? *3 H N m .3 5‘3 £1 2 33 a. 5. Acer rubrun 3 1 1 5 Acer saccharun 1 1 Carya ovata 2 Caryn sp. 1} 7 3 2 16 Fraxinue sp. 1 3 “ 1. 1 Populus grandidentata 2 2 Prunus sp. 1 1 Quercus alba 5 11 13 8 3'7 15 1 1 13 21 Quercus bicolor 2 1 2 5 Quercus borealis 1 2 8 11 22 1 1 Sassafras albidum 3 3 Total ' 21 25 30 at» 23 9O Stand # 10. Southern Dry-Mesio forest, oak-hickory woods. Kellog Forest north of Augusta, Kalamazoo County, Michigan. T15, R9W, sect. 22. Tree species Diameter in cm. Lichens ‘3 o .,-| w c) a a be a a «H a 1: :2. a a: a .g c) a g :1 E a. 33 O\ (A 0\ r1 8' E; 0 r4 3 ‘3‘ 2? cs 3 5 3* ‘3 *3 .4 (N cn ;: £3 .2 ‘2 5g Es Acer rubrum 10 11 3 1 20 Acer saccharum 5 2 3 3 13 Carya ovata 1 1. l 1 Caryn sp. I 2 7 9 1 1+ 5 Cornus florida 1 1 Jugans cinerea l l Irunus sp. 5 3 1 7 12 Quercus alba 5 3 9 17 9 3 9 Quercus borealis 5 1 8 11+ 1 1 Sassafras albidum 2 l 3 Ulmus americana 2 l 3 Total 35 23 9 35 16 91 a ,7. M, w T:- “~- ~, r *wwv-z c- m; "7 .Mi‘"f)fl.z_u..ij A ii. . ijiflanuivflmN ULfijVLfiAibb I‘ll. 7 V I '3 .1.-‘ J11 . x, a .' ,.,‘I ' - ‘ ». “’ :\; bar Lasso fibulbu rs;a [i 33) . A1 Betula papyrifera Carpinus caroliniana Gleditsia triacanthos Platanus occidentalis Populus tremloides Juglans cinerea Acer saccharum Fagus grandifo ‘a Abies baisamsa Acer rubrum Fraxinus sp. Betula lutea Carya ovata Carya sp. ‘3 A r" Lh o pyr e 11 la finKii \ L...“ 4 4 . A. iii‘>'T-LCCZI':3 >< l... puncti— X X oruis A. spfiaeroidss A. thomsonii A. milieyana X X X X X X Bermatina X pyrenocarpa Leptorhaphis X atomaria L. contorta X L. epidermidis . X L. parameca X Microthelia X X X X X X micula %. wallrothii X Rolestia - X leUCOplaca Plagiocarpa X x hyalospora Polyblastiopsis X X fallaciosa P. lactea >4 >4 >4 >4 >4 >4 Pyrenula agawae X P. neglecta X Trypnetnelium X virens 92 Prunus sp. >4 Quercms alba Quercus borealis Tilia afiericana >3 Ulmus americana Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. l. 2. 3. 4. S. 6. 8. 9. Plate 2. Spores of Arthgpyrenia ArthOpyrenia finkii A. sphaeroides A. A. A. A. A. A. punctiformis willeyana guisquiliae guinquesgptata quingueseptata (with two longitudinal septa). myricae thomsonii 94 Plate 3. Spores of Leptorhaphis, Microthelia and PolzhlastiOPSis Fig. l. Polyhlastiopsis lactea Fig. 2. P. meridionalis Fig. 3. P. fallaciosa Fig. #. Microthelia wallrothii Fig. 5. Spore representative of Microthelia micula group. Fig. 6. Leptorhaphis contorta/L.gparameca Fig. 7. L. atomaria Fig. 8. L. gpidemidis 96 Plate he Fig. Fig. Fig. Fig. Fig. Fig. Fig. l. 2. 4. 5. 6. 7. Spores of Dermatina, Melanomma, Molestia, Plagiocarpa, Ezrenula and Trlphethelium Tryphethelium virens Plagiocarpa hyalospora Dermatina pyrenocarpa Melanomma sp. Molestia leucoglaca Eigenula agawae P. neglecta 98