MORPHOLOGICAL AND ANATOMICAL STUDIES IN $QME GENERA OF THE PANNARIACEAE Thesis gov fire Deg?“ of M. 5. MECEEEAN STATE URIVERBETY Carol Maya lata Michael 1964 THESE LIBRARY Michigan State University —— ABSTRACT MORPHOLOGICAL AND ANATOMICAL STUDIES IN SOME GENERA OF THE PANNARIACEAE by Carol Maya lata Michael This study is primarily on the morphology and anatomy of some species of each of the following genera: Coccocarpia, Erioderma, Hydrothyria, Massalonqia, Pannaria, Parmeliella, Placynthium and Psoroma. These genera were studied to find some anatomical characters which might be used in the taxon- omy of the Pannariaceae. Little anatomical work has been done recently on this family, hence a need was felt for this study. Some species from the West Indies and some from the United States were utilized for this study. There has been no uniformity in the terms used for the different tissues in the thalli and apothecia and it is hoped that the terms used here can be of help as far as uniform terminology is concerned. First, the anatomical characters of the apothecium and then those of the thallus are used in suggesting any Carol Maya lata Michael change in the present circumscription of the Pannariaceae. In addition, phylogenetic advancement in the family is re- vealed by the increasing complexity of some fungal tissue, e.g., from lower pseudocortex in Hydrothyria venosa to a true lower cortex in Massalonqia carnoga. Relationships between the genera are evident. Algal characters can be used to a great advantage along with the fungal characters in identifying the genera and sometimes even the species, but they are of no taxonomic value by themselves. The apothecial characters are very interesting. Although the apothecia in Pannariaceae were formerly thought to be either lecanorine or lecideine, they appear to me to be lecanorine or mycolecanorine. Since anatomy and morphology of Placvnthium nigrum is quite different from the others, it is suggested that this species be removed from the Pannariaceae and be placed in a family of its own. It is also suggested to leave Erioderma physioides as a species of Erioderma and not be made the type species of new genus Malmella. MORPHOLOGICAL AND ANATOMICAL STUDIES IN SOME GENERA OF THE PANNARIACEAE BY Carol Maya lata Michael 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 1964 ACKNOWLEDGMENTS It is with deep sincerity and grateful apprecia- tion that I thank my major professor, Dr. Henry A. Imshaug, not only for the use of his collection which formed the basis of this investigation, but also for his untiring help, guidance and inspiration, without which this present work would not have been accomplished. I also want to gratefully acknowledge Dr. Everett S. Beneke and Mr. Irwin M. Brodo for their advice and val- ued assistance during the absence of Dr. Imshaug. It is with deep appreciation that I want to thank the members of my Guidance Committee, Dr. William B. Drew, Dr. Everett S. Beneke, Mr. Irwin M. Brodo, Dr. Roland L. Fisher, Dr. Irving W. Knobloch, and Dr. John L. Lockwood, for their suggestions and advice concerning the present study and for willingly reading and correcting the final manuscript of this thesis. Thanks are due the Michigan State University Library (Reference Department) for the assistance they so generously [provided in obtaining needed publications from other libraries. ii My special thanks to Conservatoire Botanique, Geneva, Switzerland, for loaning the type specimen of Erioderma polycarpa and Herbarium of University of Mich- igan for loaning the other species of Erioderma. Sincere thanks are due Mrs. Judy Burgess, Mr. Irwin Brodo and Mr. Clifford Wetmore, my colleagues at the Lichenology Laboratory of Michigan State, for their helpful suggestions, encouragement throughout my inves- tigation, and generous help in the translation of foreign publications used during the course of my study. To them as well as to all the staff of the Botany Department I am grateful; their friendship and help in more ways than one have made my stay at Michigan State a most pleasant and rewarding experience. Last but certainly not the least, I wish to thank my husband Raj, for his faith, encouragement, numerous helps and endless patience. iii CONTENTS Page ACKNOWLEDGEMENTS. . . . . . . . . . . . . . ii LIST OF PLATES. . . . . . . . . . . . . . . v Chapter I. INTRODUCTION. . . . . . . . . . . . . . . . 1 II. HISTORY OF THE FAMILY PANNARIACEAE. . . . . 4 III. MORPHOLOGICAL AND ANATOMICAL CHARACTERS . . 14 Methods of Study Thallus Apothecia IV. THE PHYCOBIONTS . . . . .. . . . . . . . . 36 V. LICHEN SUBSTANCES . . . . . . . . . . . . . 39 VI. KEY TO THE GENERA STUDIED . . . . . . . . . 41 VII. THE GENERA STUDIED. . . . . . . . . . . . . 44 VIII. DISCUSSION. . . . . . . . . . . . . . . . . 118 IX. SUMMARY . . . . . . . . . . . . . . . . . . 131 X. LITERATURE CITED. . . . . . . . . . . . . . 134 XI. PLATES. . . . . . . . . . . . . . . . . . . 138 iv Plate' II III IV VI VII VIII IX XI XII LIST OF PLATES Page COCQOCarpia_§ayana. Cross section of the thallus. . . . . . . . . . . . . . . . . 138 Coccocarpia Gayana. Longitudinal section of the apothecium. . . . . . . . . . . . 140 Cgccocarpia pellita. .Longitudinal section of the apothecium (diagram). . . . . . . 142 Erioderma microcarpa. Longitudinal section of the apothecium (diagram). . . . . . . 144 Erioderma physioides. Cross section of the thallus. . . . . . . . . . . . . . . . . 146 Erioderma physioides. Longitudinal section of the apothecium (diagram). . . . . . . 148 Erioderma physioides. Longitudinal section of the apothecium (drawing). . . . . . . 150 Erioderma polycarpa. Longitudinal section of the apothecium. . . . . . . . . . . . 152 Erioderma Wrightii. Cross section of the thallus. . . . . . . . . . . . . . . . . 154 Erioderma Wriqhtii. Longitudinal section of the apothecium. . . ... . . . . . . . 156 Hydrothvria venosa. Cross section of the thallus. . . . . . . . . . . . . . . . . 158 ,Hydgothyria venosa. Longitudinal section of the apothecium. . . . . . . . . . . . 160 Plate XIII XIV XV XVI XVII XVIII XVIX XXII XXIII XXIV Page Massalongia carnosa. Cross section of the thallus. . . . . . . . . . . . . . . . . 162 Massalongia carnosa. Longitudinal section of the apothecium. . . . . . . . . . . . 164 Pannaria lurida. Cross section of thallus. 166 Pannaria lurida. Longitudinal section of apothecium . . . . . . . . . . . . . . . 168 Pannaria pezizoides. Longitudinal section of apothecium (diagram). . . . . . . . . 170 Parmeliella microphvlla. Cross section of thallus. . . . . . . . . . . . . . . . . 172 Parmeliella plumbea. Cross section of the thallus. . . . . . . . . . . . . . . . . 174 Parmeliella plumbea. Longitudinal section of the apothecium. . . . . . . . . . . . 176 Placynthium nigrum. Cross section of an areole of the thallus. . . . . . . . . . 178 Placynthium nigrum. Longitudinal section of the apothecium (diagram). . . . . . . 180 Psoroma hypnorum. Longitudinal section of the apothecium . . . . . . . . . . . . . 182 Psoroma pallidum. Cross section of the thallus. . . . . . . . . . . . . . . . . 184 Collema. Types of excipular tissue . . . . 186 vi INTRODUCTION The purpose of this investigation of the family Pannariaceae is 1) to search for new characters that might be used in the taxonomy of the family, 2) to evaluate the advisability of recognizing Placynthiaceae as a family. and 3) to evaluate the advisability of recognizing MEI: mella as separate genus. The genera studied were: Coccocarpia, Erioderma. lflygrothyria, Massalongig, Pannaria, Parmeliella, Plagyn— thium and Psoroma. Representatives of available species of these genera were studied both morphologically and ana- tomically. This paper deals with genus Erioderma in the greatest detail and a key to the species studied is pro- vided; also a key to the genera, based on the species identified, is included. The specimens for this study were obtained from Dr. H. A. Imshaug's collection of West Indian lichens and from the general lichen herbarium at Michigan State University. The type specimen of Erioderma ,microcarpa was borrowed from Conservatoire Botanique. Geneva, Switzerland, and the other species of Erioderma seen for this study were obtained from University of Michigan herbarium. Most of the species studied are tropical ones. The Pannariaceae has been and is in great need of study. Many changes have taken place since this family was established by Tuckerman (1872). New genera have been included and original genera excluded. This has been due to a lack of complete information about any particular genus, and, of course, because of different ideas about morphological and anatomical characters. This study is necessarily of a preliminary nature, but some new charac- ters have been noted which have not been mentioned before and it is hoped that they will be of some help towards more extensive work. All the original descriptions of the genera, as well as those of species of Erioderma have been obtained and referred to when necessary; a historical survey of the Pannariaceae is also included. The description and the plates of the genera and their species are arranged alphabetically. In the last section detailed drawings and diagrams are given. Drawings of both thallus and apothecium of each species are included except a few cases where only one or the other is provided. II HISTORY OF THE FAMILY PANNARIACEAE The composite nature of the lichen thallus was not known until 1867 (Schwendener). Prior to this time it was impossible to consider the algal component in the taxonomy of the lichens. When the Pannariaceae was des- cribed for the first time by Massalongo (1855), and later by others, only the morphological characters of the thal— lus and apothecium seemed to be important. Massalongo (1855) divided lichens into several orders, order No. VII being the Parmeliaceae. He divided the Parmeliaceae into many tribes and one of them was Pannariaceae. The genera he included in this tribe were. Pannaria (this was under Lecidiaceae according to Bory. 1828), Capodium, Massaloggia, Polyzosia and Sporoaconia. He put Psoroma in a separate tribe (Psoromae) along with Fglgenia, Gyallolukia, Solengspora. Niospora and Acaro- spgrg. Before this Psoroma was often included in the Lecanoraceae. The tribes were mainly separated on the basis of whether they were foliose or crustose. Sometimes it was hard to separate one growth form from the other. and therefore, foliose species were occasionally put with crustose ones and vice versa. In the same year (1855), Nylander described Pan- nariei as a subtribe of Lecanorei and included the fol— lowing genera: Erioderma, Pannaria, Coccocarpia,‘gg£a and Qichogema. Koerber, also in 1855, described the Pannariaceae as a subfamily of Lecanoreae including Pannaria and Massa- lgggia. The genus Psoroma was put in the next family. Placodinae. I Mudd (1861) considered the subtribe Pannariaceae in the tribe Lecanoreae and included Pannaria, Massalongia and Ampgilgma in it. Nylander (1861) once again described Pannariei as a subtribe of the tribe Pyxinae and included Heppia and Pannaria. Reinke (1895) described Pannarieen as a family under the order Cyanophili. Within this category he in- cluded Plagynthium, Leptogidigm,.gglychidium, Heppia. Hetgrina, Leprocollema, Parmeliella, Stictina, Sticta, Ricagglia, Pannaria, Lepidocollema, Psoroma, Erioderma, and Hydrothyria. Before this, some species of Erioderma belonged to the Peltigeraceae, some to Stictaceae and some to the genus Lichen. The genus Hydrothyria was put under Collemaceae by Russell (1853). Zahlbruckner (1907) introduced some better ideas about the classification of lichens and based them on more natural character. He recognized Pannariaceae as a family and included Psoromaria. Psoroma. Lepidocollema. Lepidgleptogium, Pannaria. Hueella. Pagmeliella.gPlacyn- sh;2_. Erioderma. Coccggagpia and Hydrothyria. He gave the following key for their identification. A. Thallus with dark green phycobiont. I. Apothecium biatorine ..... . ................ Psoromaria II. Apothecium lecanorine ...................... ...Psoroma B. Thallus with blue-green gonidia. I. Lower surface of the thallus indistinctly veined; prothallus and rhizoids usually well-developed; spores one-, seldom two- celled, colorless. a. Upper cortex narrow. indistinct; algal layer almost taking up the entire breadth of the thallus...... ..... ..Lepidocollema b. Upper cortex well-developed, distinct. 1. Upper cortex arising from the hy- phae which are perpendicular to the surface. (a) Upper surface of the thallus naked. (1) Thallus with Nostoc. [a] Apothecium lecanorine. [1] Upper cortex paraplec- tenchymatous, lower cortex lacking...... .....Lepidoleptogium [2] Upper and lower cor— tices well-developed. a) Spores one-celled .........Pannaria b) Spores two-celled ..........Huee11a (2) Thallus with Scytonema. [a] Apothecium biatorine or lecidiene. [l] Spores one-celled... ......... Parmeliella [2] Spores two-celled... ......... Placynthium (b) Upper surface of the thallus shaggy ..................... ..... Erioderma 2. Upper cortex arising from the hyphae which occur horizontally.....Coccocarpia II. Under surface of the thallus veined; prothallus and rhizoids lacking; spores parallel, 4-celled .......... ............... ..... ...... Hydrothyria Zahlbruckner (1907) considered the Pannariaceae as closely related to the Heppiaceae and the Stictaceae. The structures of the thallus and the apothecium of members of the Pannariaceae are intermediate between the two families and the Pannariaceae is united to both by the intermediate genera, Lepidgcollema of the Heppiaceae and Lgbaria of the Stictaceae. This made the correct placement of the Pannar— iaceae difficult. These three families perhaps represent a monophyletic series, the origin of which Reinke (1895, p. 344) considered as the genus Parmeliella. Reinke (1895) also pointed out the close relationship between the Pannar- iaceae and the Peltigeraceae, which is clearly indicated by genus Hydrothygia. Zahlbruckner (1907) later suggested that gagggma should be included in the Pannariaceae in spite of its having dark green algae. Before this Psoroma was placed in the Lecanoraceae as a section of Lecanora. Hue (1912) reduced the whole family Pannariaceae to one genus, Pannaria. His sections and subsections of the genus corresponded more or less closely to the genera of the other workers. According to Smith (1921), the Pannariaceae and the Heppiaceae are intermediate between families, ‘with less developed. gelatinous and homoiomerous thalli and those with advanced and entirely heteromerous thalli. She saw an evo- lutionary development within the family from genera with blue-green phycobionts (the large majority) to two (Psoroma and Psoggmaria) having green phycobionts. Vainio (1923) included Megalospora. Physcidia and Heterodea in the Pannariaceae. They were shifted to Parmel- iaceae by Zahlbruckner in 1926. Zahlbruckner (1926) included the same genera in the Pannariaceae as he did in 1907; they were Hydrothyria,‘Lg- pidocollema, Masgalongia, Lepidoleptogium, Placynthium, Parmeliella. Pannaria. Hueela. Pgorgma. Psoromaria. Cocco— carpia. and Erioderma (listed here according to zahlbruck- ner's phylogenetic arrangement of the genera). Dodge (1933, p. 438) wrote, "In many ways the fam- ily (Pannariaceae) seems very homogeneous, so that genus distinctions are often rather arbitrary." He recognized the following seven genera from tropical America: Psoroma. Lepidogollema, Pannaria, Malmella. Parmeliella, Erioderma. and rogocarpia. He formed the new genus Malmella by com- bining one species of Erioderma (g, physioides) and one spe- cies of Pannaria (g. rubiginosa f. cinerascens) and adding three new species. 10 Gyelnik (1940) included the following genera in the Pannariaceae: Massalongia. Hueela. Placynthiella, Parmeli- .gllg, Pannaria, Molleropsis, and Vestergrenopsis (the last two being gen. nov.). However, he put the genus Psoroma in the Stictaceae. He gave more importance to spore characters in separating the different genera and used thallus charac- ters-—heteromerous or homoiomerous. He removed the genus Placynthium from the Pannariaceae and put it in the Lichen— aceae. He considered the Pannariaceae as closely related to the Heppiaceae and Lichenaceae. Rasanan (1943), like Zahlbruckner (1926), placed the Pannariaceae between the Heppiaceae and the Stictaceae. He did not make major differences in the previous circumscrip- tion of the family Pannariaceae but included 7 more genera than did Zahlbruckner. The nineteen genera that he included in this family were: Hydrothyria, Coccocarpia (and its two sections Subimbricaria and Umblicularia), Erioderma, Theli— dea (Which Zahlbruckner considered as a genus of doubtful position). Psoromaria. Psoroma. Hueella. Massalonqia. Pan- naria, Placynthium, Amphidium (syn. Steinera, Which Zahl- bruckner had in Lichenaceae), Placygthiopsis. Huillia, Par- meliella, Lepidocollema. Lepidoleptogium, Megalospora. and ll Physcidia and Heterodea (the last three genera having been put in the Parmeliaceae by Zahlbruckner). Dahl (1950) formed the new family Placynthiaceae to include Placynthium, formerly in the Pannariaceae, Porocyphus. formerly in Ephabaceae. Pterygium. formerly in Lichenaceae (Zahlbruckner 1926). and Vestergrenopsis, formerly in Pannar- iaceae (Gyelnik 1940). When Dahl (1950) was describing 2939— gyphus from Greenland he was aware that Placynthium and Eggp- cyphus were morphologically and anatomically much alike. He also Observed that Placynthium and Pterygium were much too similar to be put in two separate families. According to him the anatomy of the thallus lobes of vestergrenosis igigi- 33g and those of Placynthium pannariellum are almost alike. He suggested that instead of putting the four closely related genera, Porocyphys, Placynthigm, Pterygium. and Vesterqrenop— gig, in three different families. they should be put together in one separate family which he called Placynthiaceae. Dahl (1950) presented two major differences between Pannariaceae and Placynthiaceae. He studied the algae in both families by setting them free in a quetsch (crush-mount) preparation. In the Pannariaceae he observed that the algae are true Nostoc (typical rosary-like chains with smooth 12 globose cells) with no distinct orientation of chains within the thallus. He commented that very often in some species of Pannariaceae scytonemeiform (Scytonemgelike) Nostoc occurs. Dahl calls the algae found in Pannaria and other genera of Pannariaceae "nostociform" whereas in Placynthiaceae he calls it "scytonemeiform," because in the latter the algal cells are larger. irregularly globose, and in irregular chains. The second important difference between the Pannar- iaceae and the Placynthiaceae that Dahl observed was that in the latter there is a distinct tendency toward forming minute areoles in the thallus; this does not occur in the Pannaria- ceae. Dahl further reported that though the external morph— ology of Coccocarpia in some respects approaches that of the other members of the Pannariaceae, he thought that, according to Bornet's (1909) figures, it has scytonemeiform algae. Dahl does not say clearly whether Coccocarpia should be included in the Placynthiaceae, but if keyed out according to his key it does come under this family rather than under Pannariaceae. By recognizing Placynthiaceae as a separate family Dahl has made Placynthium the type genus and has included Pterygiopsis, Porocyphus, Pteridium and Vesterqrenopsis, 13 within the family. He also thought that Leptogidium, Lichen- 'legm. Lichenella, Homospella. and perhaps Asirosiphon may belong to this family as well. Dahl also suggested that Erioderma should be placed in a family of its own, but he did not propose the name. This family. he said, should have a similar relationship to the Placynthiaceae as Peltigeraceae has to Pannariaceae. He came to this conclusion because Egigderma has scytoneme- iform algae but since the thallus differs considerably from that of members of the Placynthiaceae, he did not recommend including it in that family. Thus Dahl (1950) put the following genera in the fam- ily Pannariaceae; Hydrothyria, Lepigocollema, Lepodolepto- 332m, Massalongia. Parmeliella, Pannaria, Hueella. Psoroma. Psromaria, and Placynthiella (Gyelnik 1940). Recently Choisy (1960) recognized Pannariaceae and Placynthiaceae as separate families. His work was mostly concerned with the genera from these families found in a particular region of France. Hale (1961) included Placynthigm in the Pannariaceae. According to him most of the species belonging to the Pannar- iaceae are crustose or squamulose and very rarely are subfo- 1iose and foliose. III MORPHOLOGICAL AND ANATOMICAL CHARACTERS Methods of Study: The morphological and anatomical characters of both the thallus and apothecia were studied. Microtome sections were made in the following manner. The specimen was soaked overnight in "Photoflo" (a concentrated liquid wetting agent). The material was mounted on the freezing disk of the micro- tome in a drop.of gum-Arabic. Sections were cut at approxi- mately 15 p thickness and were picked up with the help of a camel-hair brush, deposited in a watch glass containing water. and selected. The selected sections were stained in acid fucshin (10-15 minutes) and were passed through an alocohol series (from 70 percent to 100 percent) for 10—15 minutes in each of the series. Finally they were put in butyl alcohol for 15-20 minutes and mounted in diaphane. If the material was too small (e.g., Placynthium nigrum) it was embedded in agar and the agar blocks were then mounted on the freezing disk. The sections thus made were studied and drawn by means 14 15 of camera-lucida. Both algal and fungal components were noted and compared among the species. The orientation of the hyphae and algal filaments was carefully considered for Possible use as a generic or specific character. Thallus: The lichen thallus is composed of both fungal hy- phae and algal colonies, each with a definite growth form. The thallus characters in the family Pannariaceae have not been uniformly described by lichenologists and different terms have often been used for the same tissue. One of the objectives of this study was to compare the tissues in the various species in the Pannariaceae and describe them by using a uniform terminology. ‘Mgrpholoqv of the Thallus: The thallus in the Pannariaceae varies from foliose to almost crustose. The best example of a foliose thallus is that of Erioderma Wrightii in which the lobes are excep- tionally large for the family as a whole; they are 1.0-1.5 cm broad. Other species of Erioderma are also foliose but their lobes are not so large as those of g, Wrightii. 16 Additional examples in the family having foliose thalli are species of Parmeliella, Pannaria lurida. Massalongia.g§£n- .ggg, and Hydrothyria venosa. These species have thallus lobes 0.6—1.0 cmbroad. Species with thallus lobes smaller than would be found in typical foliose thalli are termed "subfoliose." The lobes of the subfoliose species are 3.0-5.0 mm broad. These species are Pannaria leucosticta, Psoromahypnorum, and a few others belonging to the genera. Pannaria, Parmel- uigllg and Psoroma. Species with thallus lobes 1.5-3.0 mm in size are called squamose in this study, examples are: Parmeliella lepidigta, Parmeliella microphylla. and Pannaria pezizoides. The thallus of Placynthium nigrum is basically crus- tose, being broken into small areoles. These areoles are 1.0-1.5 mm wide. Sometimes the defining line between foliose and squamose is very fine and it is often hard to say Whether a given thallus is one or the other. The color of the thallus varies in different genera. even between the species. It can be yellowish—green, brown- ish. or black. 17 In foliose species the primary lobes divide dichoto- mously to form secondary lobes or lobules. Lobes are mostly contiguous, and sometimes imbricate with swollen tips as in Massalgngia carnosa or with crenate tips as in Pannaria £3- biginosa. The thallus surface is either tomentose or glabrous. Sometimes the sides of the apothecia have the tomentum though the thallus is without any, as in Hydrothyria venosa and in Massalongia carnosa. Hairs are septate or non-septate. and branched or simple, arranged densely or sparsely and are colorless. Either isidia or soredia or both may be present. Isidia are mostly marginal and are concolorous with the thallus or are either white or grayish, and globular or cylindrical in shape. In Massalongia carnosa they are con- colorous with the thallus and are marginal and corolloid. Isidia were seen in cross-section in Coccocarpia giliata: they had 1—2 layers of cortical cells and 2-6 algal cells. They do not occur as often as soredia in this family. Soredia occur on the upper surface, they are fari— nose and are either marginal or scattered in groups. They are small microscopic bodies consisting of few algal cells 18 accompanied by few gelatinized hyphae, are non-corticate and often bluish-white. All the genera studied here have sorediate species except Placynthium and Massalongia. Rhizinae occur on the under side of the lower mar- gins of the thallus. They are usually quite dense and are black, brown. or even pale. Hyphae in the form of cords extend from the lower part of the medulla, pseudocortex or lower cortex, to the substratum and anchor the thallus firmly. They are either on the entire underside, or just on the lower margins, or on the underside exclusive of the margins. Lichenologists, e.g., Smith (1921) and Hale (1961). have distinguished between a hypothallus and a prothallus. although they both are non—lichenized tissues within the lichenized thallus. Tulasne (1852) recorded the thallus growth in 2g;- rucaria muralig, which is an epilithic crustose species. He observed that the spore on germination produces a deli— cate branching, septate hypha which produces radiating secondary hyphae on all sides. Zukal (1895) called this beginning of a thallus a prothallus or protothallus. The hyphae of the prothallus adhere to the substratum and form 19 a kind of plectenchymatous tissue. Algae come in contact with these fungal hyphae and the contact between the two increases their growth and cell division. Zukal (1895) also observed that the hyphae in contact with algae become more and more thin—walled and produce many new branches. Some of these new branches grow upwards and form the cor- tex and some grow downwards and build up the medulla. but the hyphae at the circumference continue to advance and may start new areas for the algae to come in contact with: these areas he called the prothallus or protothallus. Smith (1921) said that the prothallus becomes over~ grown and obscured by the vigorous increase of the first formed lichenized tissue and can be seen as a white or dark line bordering the thallus. Earlier, Schwendener (1866) stated that the lichens which develop from the spores have a prothallus (protothallus) and those which arise from so— redia do not have this first thalline tissue. Hale (1961) explained that the hypothallus is a thin layer of non-lichenized hyphae which projects beyond the main thallus and appears as a thin black rim, and accor- ding to him, its function is unknown. Smith (1921) recog- nized the hypothallus as formed by an unusual development 20 of hyphae from the lower medulla or lower cortex. These hyphae become dark-brown and are divided into thick—walled cells. Frequent branching and anastomoses of these hyphae result in a Cushion-like structure as in Angie colpodes and Pannoparmelia anzioides (Fig. 51, Smith, 1921). Fink (1935) said that hypothallus is the first growth of hyphae before any differentiation has taken place, and it often persists as a dark layer below or surrounding a few lichen thalli. The hypothallus is.a special tissue which forms a rather thick, cushion—like structure of thick-walled, branched, and cellualar hyphae, as in Pannoparmelia anzioides (Fig. 51, Smith, 1921). A dark line bordering the thallus can be very distinctly seen in Parmeliella plumbea and Ear- meliella pannosa. Underneath the thallus, a mat of rhizinae is also formed in these species. This dark line of hyphae bordering the thallus and forming a mat underneath, belong to the prothallus (Dodge. 1933, called this hypothallus). In a cross section of the thallus the mat of rhizoidal tis- sue is very clearly seen (Plate 19). This tissue is formed of parallel, conglutinate hyphae which do not form a thick cushion—like, arachnoid tissue. Although the tissue appears 21 Although the tissue appears hypothalline in cross-section. it would seem to be prothalline in origin. Anatomy of the Thallus: Wallroth (1825. l. in Smith, 1921) was the first to study and compare different lichen thalli from an ana- tomical standpoint. He called lichens in which algal cells are distributed uniformly along with the fungal hyphae through the entire thallus "homoiomerous," and those in which they are confined to a distinct zone "heteromerous" Smith (1921, p. 21). The Pannariaceae along with Heppia- ceae form a transition between the gelatinous and mostly homoiomerous thallus and the true heteromerous thallus (Smith. 1921, p. 286). In Parmeliella microphylla and Placynthium nigrum (Plates 18 and 21 respectively) the algae occupy the entire thickness of the thallus. In these two cases a cortical layer surrounds the part of the thallus in which the alga and fungus are distributed without any definite layers. 1. The upper cortex in the Pannariaceae is paraplec- tenchymatous. Lindau (1899) introduced term "p1ectenchyma," by which he meant any tissue of hyphae. He also introduced 22 the terms parenchymatous plectenchyma or paraplectenchyma and prosenchymatous plectenchyma or prosoplectenchyma, the former denoting the tissue with more or less isodiametric cells (the more typical "pseudoparenchyma") and the latter denoting that with more elongated cells. The hyphae in the upper cortex are oriented in dif- ferent ways. They are parallel to the surface of the thal— lus in Erioderma physioides (Plate 5), Hydrothyria venosa (Plate 11), Pannaria lurida (Plate 15), and Parmeliella microphylla (Plate 18) and they are almost parallel to the surface of the thallus in Coccocarpia Gayana (Plate 1). The hyphae are vertical to the surface of the thallus in Parmel- iella plumbea (Plate 19), Parmeliella pannosa, and Massalon- qia carnosa (Plate 13). The hyphae of the upper cortex are irregularly arranged or diffuse in Erioderma Wrightii (Plate 9), Erioderma polycarpa and some other species of Coccocarpia. The upper cortex varies in thickness from one to many cells. In Placynthigm nigrum (Plate 21) it is just one cell thick and completely surrounds the areole of the thallus. It is 2-3 cells in Parmeliella microphyllg (Plate 18) and the hyphae are thick-walled. The layer is 1-2 cells thick in Massalonqia carnosa (Plate 13) and the hyphae here too are rather thick-walled. 23 In all the species of Erioderma that are studied here. except g, limbatum, the upper cortex is covered with a tomentum. These hairs are cylindrical, either straight or curved. They are mostly unbranched and occasionally septate, and are either dense or sparse. The tomentum is made up of the thickened ends of more or less vertical hy- phae. 2. The algal layer comes next to the upper cortex. This layer is usually narrower than the medulla, although in Parmeliella microphylla (Plate 18) and Placynthium gi- ggum (Plate 21) it is the widest layer. In these species the medullary layer is absent and the whole thickness of the thallus is taken up by the algal layer. In Psoroma pallidum (Plate 24), the algal layer is very narrow and the algal chains are not close together. In some species they are very densely packed. e.g., Parmeliella microphyllg (Plate 18). The algae in the genera studied here are Egg- .tgg, Scytonema in the cyanophyta. and Dactylococcus or £93: comyxa in the chlorophyta. 3. Below the algal layer is the medulla. It is usually a thick layer of loosely interwoven thin-walled hyphae, which are generally irregularly arranged. They are periclinal in 24 Coccocarpia Gayana (Plate 1), parallel to the surface of the thallus in Hydrothvria venosa (Plate 11), almost paral- lel to the surface in Erioderma physioides (Plate 5) and, parallel. clearly septate and branched in Massalonqia‘ggr- Iggy; (Plate 13). In Erioderma physioides (Plate 5) the medullary hyphae are accompanied by special kinds of cells called spheroidal cells by Asahina (1952). He saw them for the first time in Parmelia sublaeviqata. Culberson (1961) re- ported them in Parmelia galbina and called them moniliform cells. These cells are in the form of beaded filaments and take a pink stain with acid fucshin. The significance of these cells in the medulla is not yet clear. 4. All the species of family Pannariaceae that are in— cluded in this study except Massalonqia carnosa (Plate 14), are without a true lower cortex. The above mentioned spe- cies has a true lower cortex since it is exactly like the upper cortex (Degelius, 1954). In some species there is a tissue below the medulla which looks like a cortex but does not resemble the upper cortex. This lower layer is called a pseudocortex by Degelius (1954). There are some doubts and arguments about the presence of a true cortex in different species of the Pannariaceae. 25 In Hydrothyria venosa (Plate 11), the lower part of the medulla has septate conglutinate hyphae and is best termed a pseudocortex. In some species of Coccocarpia the lower pseudocortex can be seen, Dodge (1933) called this tissue the lower cortex. The lower pseudocortex is an indistinct but easily recognizable layer where it is present. Placvnthigm,nigrum (Plate 21) has a cortical layer just one cell thick over all the exposed surfaces of the areole of the thallus as the outer most layer surrounding the areaole. In Parmeliella microphylla (Plate 18) a lower pseudo cortex is present. The difference between this and the upper cortex is quite clear because the tissue in the lower pseudocortex is loose as compared to the upper cor- ~ tex. 5. Rhizinae arise from the under surface of the thallus and are either black or brown. They extend from the lower part of the medulla, lower cortex, or the lower pseudocor- tex. The rhizinae are simple or branched, septate or non— septate, in groups or single; they often anastomose. There is thick prothalline mat formed by the rhizoidal hyphae in Parmeliella plumbea (Plate 19), Parmeliella pannosa and 26 Cogcggarpia pellita. Presence of this tissue is not a diag- nostic character for any genus. Apothecia: There has been much confusion concerning the usage of the terms for different tissues in the apothecium of the family Pannariaceae. The terms I have used here are those which were adopted by Degelius (1954).. figrpholoqv of the Apothecia: When apothecia are present, they are found on the upper surface of the thallus. They are either abundant or rare and can be either marginal or scattered. In most of the species they are sessile, although in all the species of Erioderma they are stipitate. In many species of £2999- carpia they are adnate to the thallus. The apothecia are usually circular and flattened (disk—shaped). Sometimes they are slightly convex as in Coccocarpia pellita (Plate 3) or slightly concave as in Psoroma hypnorum (Plate 23). The apothecia may become ir— regular as a result of crowding. 27 The sizes of the apothecia differ considerably. even in the same species. However the average mature apo— thecia are 1.0—1.5 mm in diameter in most of the species except Placynthium nigrum where they are 0.3-0.5 mm in diameter. The development of the apothecia can be either an- giocarpic or gymnocarpic. Corner (1929) explained these types, and also the intermediate type, the hemiangiocarpic development. He presented a series of diagrams showing the angiocarpic development of an apothecium in the Discomycetes. Gaumann (1952) labeled the same_series of diagrams as those of a hemiangiocarpic apothecium. Whatever the case may be according to the definitions, in the angiocarpic develop— ment the hymenium is formed within a mass of interwoven hyphae. called the primary sheath; in the gymnocarpic de- velopment the hymenium is superficial from the beginning. The species of Erioderma (E. limbatum, g. physioides, g. polycarpa, and‘g. migrgcarpa) have angiocarpic or hemian- giocarpic type of development. Massalongia has hemiangio- carpic type of development of its apothecia (Henssen, 1963), i.e.. one in which hyphae do not form a closed sheath. 28 The margins of the apothecia are usually lighter in color than their disks. The apothecial disk is brown to black in color and is not pruinose generally. Anatgmy of the Apothecia: Dughi (1952) postulated some new theories about apothecial anatomy. He emphasized that an apothecial cor— tex is different from a thallus cortex in that the former is produced by the exciple. He divided the different types of apothecia into six categories according to their anatomy. 1. Lecideine apothecia: In which the exciple is uniform and there are no algae in the margins. 2. Superlecideine apothecia: In which the exciple becomes cortical in character. 3. Lecanorine apothecia: In which there are algae in the margins. A cortex and medulla are often developed. 4. Mycolecanorine: In Which there are no algae in the margins, but cortex and medulla are de- veloped. 5. Cryptolecanorine apothecia: In which the exci- pular hyphae come into the thallus for symbiosis- immersed lecanorine. 29 6. Pseudolecanorine apothecia: In which the ex- ciple is uniform but some algae accidentally occur in the margins. Apothecia of the Pannariaceae have been called both lecanorine and lecideine. I do not think that any species of this family that I have examined has lecideine apothe- cia. All the species that I examined have the amphithecium as the outer most margin in the apothecia (though this is questionable in Cocggcarpia). The amphithecium has a dis- tinct cortex and medulla and is with or without algae in different genera. Therefore. according to my observations. -the apothecia in the Pannariaceae are lecanorine (with al- gae in the amphithecium). and mycolecanorine (without algae in the amphithecium). Among the genera I have studied, Pannaria, Massalon- ELLQ (only M, carnosa studied), and Psoroma have lecanorine apothecia and Erioderma, Parmeliella, Hydrothyra, and £1;- cynthium (only g. nigrum studied), have mycolecanorine apo- thecia. In the mycolecanorine apothecia the hyphae are not receptive to the establishment of a symbiotic relation with the algae (Imshaug. 1957). 30 1. Amphithecium is the outer most layer of the apothe— cia in the genera of the family Pannariaceae that were stu- died. The amphithecium may be with or without algae, making the apothecia lecanorine or mycolecanorine. The genera.hav— ing lecanorine apothecia are Massalonqia, Pannaria, and Psoroma (Plates l4, l6, and 23). In Massalongia carnosa (Plate 14) and Pannaris lurida (Plate 16) the amphithecium is clearly divided into cortex and medulla. The apothecia in Erioderma (Plates 4, 6, 7, and 8), Hydrgthyria venosa (Plate 12), Parmeliella plumbea (Plate 20). and Placynthium Sggflagg (Plate 21) are mycolecanorine. The hyphal tissue in these species is rather compact on the outside (forming a cortex) and loose towards inside (forming the medulla). In Hydrothyria venosa (Plate 12) and Massalongia carnosa (Plate 14) a tomentum is seen on the sides of the apothecia, coming from the outer most layer of the amphi— thecium. These hairs are formed by vertical or horizontal. septate or non—septate, colorless hyphae. The amphithecium is usually paraplectenchymatous and quite often the develop- ment of this tissue from the exciple can be seen very well from the orientation of the amphithecial hyphae. 31 2. The exciple is a very thin; pale tissue between the hymenium and the amphithecium and usually is not prominent. It is usually euthyplactenchymatous and less frequently par- aplectencymatous. Euthyplectenchyma is a hyphal tissue with- out a ”cellular structure." In agreement with Degelius (1954) I consider this tissue to be loose or dense and made up of thin (filiform) or broad hyphae, which are more or less con- glutinate. They may be parallel or not and have short or usually more or less long cells. Different kinds of euthy- plectenchyma may be distinguished. e.g., a loose euthyplac— tenchyma of non-parallel hyphae, or dense one of parallel hyphae. The euthyplectenchymatous exciple runs parallel to the surface of the apothecium in the center and is per— pendicular to it in the sides. Sometimes it is quite irreg— ular. The limit of the exciple toward the amphithecium is quite sharp, but not so toward the subhymenium above. thus making it difficult to distinguish the subhymenial tissue from the exciple. There are two other kinds of excipular tissues found in the Collemataceae which are discussed by Degelius (1954): subparaplectenchyma and euparaplectenchyma (Plate 25, redrawn from Degelins, 1954). Most of the species that 32 are studied here have an exciple of loose or dense euthy- plactenchyma and few have of paraplectencyma. Coccocarpia Gayana (Plate 2), Coccocarpia pellita (Plate 3), Erioderma Wrightii (Plate 10), Hydrgthyria venosa (Plate 12):.fléééé' longia carnosa (plate 14), Placynthium nigrum and Psoroma hypnorum (Plate 23) have a loose euthyplectenchymatous ex- eXciple and the hyphae are parallel or almost parallel. In Pannaria lurida (Plate 16) the exciple is paraplectenchyma- tous. It is of irregular euthyplectenchymous tissue in Erioderma physioides (Plate 7), Erioderma polycarpa (Plate 8) and Parmeliella plumbea (Plate 20). The thickness of the exciple varies in different species. In some it is quite wide below the hymenium, and the lateral margins extend right up to the surface of the apothecium as in Placynthium nigrum (Plate 21); in.§yg£_— thyria venosa (Plate 12), the lateral margins of the ex- ciple can not be distinguished from the amphithecium, but below the hymenium it is quite thick and distinct. In Q9- ccoggrpia (Plate 2 and 3) it is not easy to separate the exciple from the amphithecium; and, therefore, the labels in the plates for these layers is necessarily approximate and perhaps tentative. 33 Sometimes the excipular tissue has pigmented hyphae as in Erioderma physioides (Plate 7). 3. The subhymenium is a very thin layer between the hymenium and the exciple but above the hypothecium when the latter is present. In most cases the subhymenium is indistinct from the excipular tissue. It looks somewhat clearer in Erioderma Wrighti; (Plate 7) though. Asci and paraphyses develop from this layer. 4. Some authors (Hue, 1906; Pink and Richards, 1915; 'Vainio. 1921, etc.) have called the central part of the ex- cipular tissue as the hypothecium and its lateral margins as parathecium. But as Degelius (1954) explains. it is a special layer of thick—walled hyphae below the subhymenium and above the exciple. This layer is not seen in any of the species that have been examined. 5. The hymenium (thecium) consists of asci and para- physes. The thickness of this layer differs in different species, usually varying between 30 and 100.n. The hymen- ial layer is colorless, except the tips of the paraphyses and the gelatine which is brown and sometimes even blaCk. This pigmented area forms the epithecial gell or epithecium. The iodine reaction of the hymenium of the species examined is I + blue. 34 6. The asci are shorter than the paraphyses, but their lengths and widths vary in different species. They are mostly clavate, e.g., Massalongia carnosa (Plate 14), Egg;- ‘Qma‘hypnorum (Plate 23), Parmeliella plumbea (Plate 20). Pannaria lurida (Plate 16) and Coccocarpia (Plates 2 and 3). The asci are cylindrical in Plagynthium nigrum (Plate 21) and Hydrothyria vengsa (Plate 12). They are oblong in most species of Erioderma (Plates 7 and 10), and in Erioderma polycarpa (Plate 8) the tips of the asci are pointed. 7. There are eight spores in each ascus arranged in one row (monostichous) or in two rows (distichous). Some- times both arrangements can be seen in the same apothecium. e.g., Parmeliella plgmbea (Plate 20), although one type usu- ally dominates, as is seen in the species of Erioderma (Plates 7 and 10). The spores are usually non-septate. They are 2-celled in Coccocarpia Gayana (Plate 2) and 31g- cynthium nigrum (Plate 22). In Erioderma Wrightii (Plate 7) and Massalongia carnosa (Plate 14), the spores are usu- ally two-celled but three-celled spores can sometimes be seen. For Massalongia the presence of two-celled spores is a generic character. The spores in Coccocarpia Gayana (Plate 2), Erioderma Wrightii (Plate 7) and Parmeliella 35 plumbea (Plate 20), have a thick epispore, though it is not so thick in the latter two species as in the first one. The spores in all the species are hyaline. The shape of the spores does not vary a great deal in the family. The spores are mostly ellipsoid to oval with rounded or pointed ends. In Magsalongia carnosa (Plate 14) and Placynthium nigrum (Plate 21). the spores are elongate elliptical to fusiform with somewhat pointed ends. 8. The paraphyses are numerous and develop from the subhymenium. They are club—shaped to filiform, e.g., figs: salongia carnosa (Plate 14) and Psgzgmg_hypngrnm_(Plate 24) respectively. They are unbranched in all the species stu- died. They are longer than the asci and almost reach the surface of the hymenium. Sometimes they are septate as in Coccocarpia Gayana (Plate 2). The tips of the paraphyses are thickened and take a dark brown or black pigmentation. The pigmented tips of the paraphyses are responsible for the color of the epithecium (area formed by the tips of the para- physes). The epithecium can be seen in most apothecia with- out any difficulty. This region is very thick in Erioderma limbatum and is absent in Placynthigm nigrum (Plate 22) and Psoroma hypnorum (Plate 24). IV THE PHYCOBIONTS Most of the genera in the family Pannariaceae have blue-green algae (Nostgc, Scytonema, and maybe Ulothrix). However Psoroma has green algae, (usually called Dactylo- coccus but probably Coccomyxa). In thalli where Nostoc or Scytonema are found, their characteristic forms are not seen many times and correct identification is only possible by making separate cultures of the gonidia. The cells of Ngstoc are usually in the form of short chains (trichomes). The chains which can be from 2 cells to 10 cells long, are arranged irregularly in the thallus and sometimes form lumps or groups. Usually they are vertically arranged but occasional horizontal chains are also seen, as in Hydrothyria venosa (Plate 11). The cells of Nostoc are normally globose, but sometimes become irregular in shape by being compressed together, as in Hydrothyria venosa (Plate 11). Besides Hydrothyria and some species of Erig— derma, Pannaria, and Parmeliella have Nostoc. Distinct 36 37 chain—sheaths are absent but individual cells are surrounded by a transparent gelatinous covering. Scytonema, seen in the form of coiled chains, is found in Coccocarpia and in some species of Erioderma,lMg§- salongia and Placynthium. Sometimes the coiled chains can be seen enclosed in thin sheaths as in Massalonqia carnosa (Plate 13). Horizontal chains of Scytonema like the chains of Nostoc in Hydrothyria venosa (Plate 11), can be seen in Cgccocarpia Gayana (Plate 1). The cells of Nostoc and Scytonema appear yellowish- green in contrast to those of Dactylogoccus (or Coccomyxa) in Psoroma, which are bright green. The abundance of the algal chains differs in differ- ent species. In Parmeliella microphylla (Plate 18) the al- gal layer occupies the whole of the central part of the thallus so that the medulla is absent. On the other hand the algal layer in Psoroma palligum is very thin (Plate 24). In all the species of Erioderma examined. the algal cells are not densely packed and the layer is quite thin. ‘Mggga- longia carnosa (Plate 13) also has a somewhat loose, thin algal layer. Hydrothyria venosa (Plate 11) and Coccocarpia Gayana (Plate 1) have quite a thick algal layer and the al- gal filaments are closer together. 38 No work on the culture and identification of the al- gae was done in this study. therefore all the observations mentioned here are based on previous works. Additional re— search work on the identification and form details of the algae in Pannariaceae is greatly needed. LICHEN SUBSTANCES Not many chemical substances have been reported from the Pannariaceae because not much work has been done on the chemistry of this family. Hesse (1901) reported the presence of hydroxyroccellic acid and pannaric acid from Pannaria languigosa. Yosioka (1941) did not find the above acids but found a new chlorine—containing neutral substance in some species of Pannaria and called it pan— narin. Asahina (1954) reported pannarin from the follow— ing species of Pannaria: ‘3. coeruleobadia, g. fulvescens. .g. lanuginosa, g. lurida. g. rubiginosa, and g. subluridg. The above species have a PD+ orange—red thalline reaction due to the presence of this chemical. Asahina (1954) also reported psoromic acid from Psoroma crasgum. The thallus of this species gives a PD+ (yellow) color. Runemark (1956) has also reported psoro- mic acid from Psoroma. 39 40 More information on the chemistry of the Pannaria- ceae is needed before chemical differentiation can be a useful tool in the taxonomy of the family. The only chem- ical test made in this study was the hymenial iodine reac- tion. In all species the hymenimnwas I+ blue. VI KEY TO THE GENERA STUDIED Phycobiont ChlorOphyta, prdbably Qggggmyxa, Thallus subfoliose, greenish in color; lobes corolloid; apothecial disk concave, brown in color. margin concolorous to the thallus; spores one-celled. monostichous or disti— chous...... ..... .................................Psoroma Phycobiont Cyanophyta............................2 2. Aquatic. Thallus foliose (but becoming almost fruticose as itgnnws away from the substrate); filaments of Nostoc oc- casionally horizontal; apothecia with some tomentum on the sides...............Hydrothyria 2. Non-aquatic.................... ...... ....3 Stipe develOped. Thallus foliose, pale green to yellowish-green, upper surface mostly to- mentose; algae Scytonemg or Nostoc; apothecia mycolecanorine (no algae in the amphithecium...Eriode;ma 41 42 Stipe not developed........................4 4. Apothecia mostly adnate, sessile or constricted below; medulla of peri- clinical, mostly conglutinate hyphae. Alage Scytggema... ...... .................Coccocg;pia 4. Apothecia not adnate; medulla of loose irregular hyphae or absent.......... ..... 5 Thallus squamulose areolate, areoles granulose; both thallus and apothecia black in color; med- ulla absent (algal layer occupies the whole thickness of the thallus); apothecia mycole- canorine; spores 1-2 celled; algae figufigunama................. ..... ..............Placyn§hium. Thallus squamulose to foliose (0.3—1.5 mm wide .............. . ....... ...... ............. 6 6. Apothecia mycolecanorine; thallus squa- mulose to foliose, brownish to grayish in color, apothecia black in color. oc- casionally brown; sometimes prothallus developed and seen as a dark line bor- dering the thallus; algae Nostoc... ..... .Parmeliella 6. Apothecia lecanorine.............. ....... 7 43 Spores mostly 2—celled, rarely 3-celled; thal- lus foliose, brownish in color; lobes imbricate; isidia corolloid; apothecia brown, quite small as compared to the thallus lobes; algae §gy£_— nega.........................................Massalongia Spores 1-celled; thallus squamose to foliose. pale green to gray in color; apothecia with cortical tissue clear in the amphithecium; algae Nostoc............... ....... ..............Pannaria VII THE GENERA STUDIED Cogcocarpia Pers. in Gaudich. In Freycinet, Voy. Uranie, Bot. 206, 1826. Type Species: ‘9, molybdea Pers. apud Gaudich. Species §tudied: .Q. ciliata, g. cropia, Q. Gayana,‘g. pellipa. L Descpiption of the Genus based 9n the Species Studied: Thallus: It is squamose to foliose and is quite often isidiate. Color of the thallus is greenish-yellow. olive gray or gray. The upper surface is without any tomentum. The lobes are 1-4 mm wide. Dark or light rhizinae can be seen on the lower surface of the thallus leaving the margins free. They are quite abundant and arise in groups rather than being solitary. The upper cortex is paraplectenchymatous. The hyphae are loose and are arranged irregularly or are 44 45 parallel to the surface of the thallus. The upper cortex is 10-30;; thick. The algal layer is 50-70,» wide and the phycobiont is §cypogema. There are vertical filaments along with oc- casional horizontal ones. The Scypogema cells have a sheath around them. In some species the upper part of the medulla has loose hyphae whereas toward the lower part the hyphae are more conglutinate. This lower most layer is called a pseudocortex because it is not like the upper cortex and the hyphae are septate, conglutinate, and form a sort of cortical tissue. Dodge (1933) called it the lower cortex. From the lower surface of the pseudocortex arise tufts of rhizinae. These are septate and very prominent. and in some species form a dark, thick mat. The tufts are 3—10‘p in width. Apothecia: The apothecia are scattered on the upper sur- face of the thallus. They are sessile or adnate, the latter being more frequent. Sometimes they are constricted below. The color of the disk varies from black to brown, or tan. The margins are concolorous with the disk. The diameter of the apothecia is 2-4 mm. 46 There has been some problem in determining whether the outer most layer is the amphithecium or exciple. That is why it is hard to say witfircertainty whether the apo- thecia are lecidiene or mycolecanorine. There are no al— gae in the outer most layer. Sometimes numerous hairelike appendages arise from the sides of the apothecia. These are cilia or may be considered a dense tomentum. The outermost tissue is paraplectenchymatous. It is quite loose and irregular in some species but in some it is of isodiametric, almost equal sized cells. The subhymenium is not distinct from the exciple or the amphithecium, whatever the outer most layer should be called. The hymenium is 80—100,u thick and has linear. oblong or clavate asci. Each ascus has 8 monostichous or distichous, one or two~celled spores. The paraphyses are also linear or clavate and are septate or non—septate. Sometimes their tips are thickened within forming an epi— thecium. 47 Coccpcarpia ciliata Wainio Annal. Acad. Scient. Fennic., Ser. A, 15(6):26. Material Studied: The material studied was misplaced. and could not be found to get the collection data. Qetailed Description: Thallus: The thallus is foliose with lobes about 5 mm broad. The upper surface is grayish. There are numerous whitish isidia scattered on the upper surface. Dark bunches of rhizinae arise from the lower surface of the thallus frequently leaving the margins free. The upper cortex is of loose paraplectencyma which is parallel to the surface of the thallus. This layer in this species is not very thick (only 2-3 celled or about 10 p). The upper cortex also continues in the isidium. The algal layer is formed by very short chains of Scytonema which are not too compact. The layer is 30—40 p.thick and the Scypongma cells are 3-5‘p in dia— meter. I could not see any horizontal filaments of the phycobiont in this species as I did in Q. Gayana and .g. pellita var. prolificaps. 48 The medullary tissue is not distinct from the lower most tissue (pseudocortex) which has linear and rectangular hyphae parallel to the surface. The lowermost hyphae continue downwards verti— cally, forming strands of rhizinae. The rhizinae are unihyphal. thin-walled, many—celled, and very conspicb' UOUS . Apophecia: The apothecia are scattered over the upper surface. They are black in color (both the disk and the margin). The apothecia are about 4 mm in diameter and they are adnate. The sides of the apothecial disk are tomentose. being covered with long conspicuous free hy— phae, hence the name "ciliata.” The apothecia are con- stricted below and at the point of the constriction are joined to the thallus. Sometimes the cilia continue up to the lower surface of the apothecia and almost look like rhizinae. The cilia are long. colorless. generally non-septate, and thin—walled. They are 2.5 p.thick and arise from the outer most layer of tissue. There is no distinction between the amphithecial and excipular tissue. The margin is paraplectencymatous and extends from below the hymenium downward. 49 The hymenium is 20.u wide and has oblong asci and clavate paraphyses. Each ascus has eight, one-celled. minute (about l‘p in diameter), round, hyaline. and thin- ‘walled spores. They are monostichous or distichous. The paraphyses do not have thick-walled tips hence there is. no epithecium. Coccpcarpia cronia (Tuck.) Wainio p, pronia (Tuck.) Wainio, Annal. Acad. Sc. Fennic., Ser. A, 6(7):103. 1915, Syn. Papmelia Cronia Tuck., Proceed. Amer. Acad. Arts &-i Sc., 1: 228, 1898; Syn. Lich. New England, 36. 1948. Matprial Studied: WEST INDIES. Grenada: Levera Bay. St. Patrick. Imshaug 16163. 1953 (MSC). Detailed Description: Thpllpp: The thallus is foliose, and the lobes are 2-3 mm wide. The upper surface is grayish-black and the lower surface is black due to presence of a thick mat (prothal- lus) formed by the rhizinae. Isidia are seen in the cen- ter of the thallus and soredia are scattered on the upper surface. 50 The upper cortex is 10-15 p wide and is formed of 2—3 layers of paraplectenchyma. The algal layer is composed of loose coils of Sc - tonema with cells mostly'lo‘p.in diameter. The layer is 35-40‘p thick and the phycobiont is more abundant than it is in Q. ciliapa. The medullary hyphae are thin-walled, irregular and colorless. The medulla is 20¢u thick and occupies a smaller area than in Q. ciliata. There is cellular tissue below the medulla; this has been called the lower cortex (Dodge. 1933), but since it is not like the upper cortex it is best called a lower pseudocortex. It is formed by thick-walled hyphae and is lO‘p wide. The rhizinae arise from the lower surface of this tissue, forming a tissue which appears to be prothal- lus. The rhizinae are long, septate. thick-walled, and form a kind of network. They are branched and arise in groups or are solitary. Appppepia: They are scattered on the upper surface and are blackish with concolorous margins. The disk is slightly convex and is 1.8-2.0 mm in diameter. The apothecia are 51 constricted below like.Q. Gayana. The surface of the apothecium, where not attached to the thallus, is covered with an abundant tomentum. The outermost layer (with no differentiation be- tween amphithecium and exciple) is composed of irregular. and thin—walled paraplectenchyma. This layer extends from below the hymenium to the bottom, and measures lOO‘p in thickness. The hymenium is.90,u thick and contains oblong asci with eight minute spores in each. The spores are one-celled, hyaline. without a thick epispore, and are either monostichous or distichous. The paraphyses are club-shaped and septate. They are unbranched and without thick-walled apices; hence, there is no epithecium as in g, Gayana (Plate 1). Coccocarpia Gayana Nyl. Synop. Lich.. 2:41. 1863. flaggrigl Studie : The material studied was misplaced and could not be found to give the collection data. 52 Dptaileg Description: Thallps: The thallus is squamulose, the squamules being 1.2,u thick. The upper surface is grayish-black and is isidiate. The rhizinae arise from the lower surface of the lower pseudocortex. The upper cortex is formed of loose. almost paral- lel paraplectenchyma (Plate 1). The layer is 20-30‘u wide and about 3-4 celled thick. The algal layer is formed of irregular algal fil- aments, which were identified by Dodge (1933) as Scytonemg. There are coils, short vertical filaments. as well as hor- izontal filaments (Santesson, 1944) (Plate 1). The algal cells differ quite considerably in size. Their diameters range between 5-10'p. The algal layer is 70‘p thick, the thickest of all species of Coccocarpia studied here. It is hard to distinguish between the medulla and the tissue below it, except that there are approximately 2-3 layers of tissue just below the algal layer which have very narrow hyphae grading into a few more layers having broader hyphae. These broad hyphae can be called the pseudocortex. These hyphae are thin—walled, conglutinate, septate, and are parallel to the surface of the thallus. 53 Rhizinae arise from the lower surface Of the pseu- docortex and come out in tufts; they are not so conspicu- ous as in Q. cronia. Apophepia: The apothecia are quite minute (about 0.9 mm in diameter). They are constricted below as in Q. cronia. The disk is black and slightly convex. It is not so difficult here to distinguish between the amphithecium and exciple (Plate 2, diagram) as it is in the preceding species since in the central part of the apothecia one can locate the excipular tissue by the ori- entation of the hyphae. The outermost layer is formed of thin—walled paraplectenchynatous cells. There are no algal cells seen in the upper parts (above the constriction) of the margins. The exciple is also of loose paraplecten— chyma. The hymenium'has clavate asci which are 8-10‘p wide near the centers. There are eight two-celled spores in each ascus. The spores are hyaline with a thick epi- spore (Plate 2). The paraphyses are clavate to filiform and are sep— tate but unbranched. There is no differentiated or pigmen- ted epithecium. The hymenium is 80 p.thick. 54 Cpccocarpia pellita (Ach.) Mflll. Arg. Flora 65: 320. 1862. .gagpglig ppllitg Ach., Lichenog. Univ. 468. 1810. Swartz Lich. Amer. 7, P1. 6, 1811. Mgpgrial Studied: WEST INDIES. Jamaica, Clarendon, gm— ahaug 13262, 1952 (MSC). Depgilpd pescription: ‘gppllpp: The thallus is foliose and is greenish-gray in color. The lobes are 3-4 mm wide and are not imbricate. The upper surface is glabrous. The rhizinae are on the lower surface leaving the margins free. The upper cortex is of loose paraplectenchyma. which is quite irregular and is 20 p wide. The algal layer has mostly vertical filaments. The cells are round and are much more uniform in Size than those in Q. Gayan . The cells are 2-5,p.in.diameter and the layer is 65 1: thick. The medulla is not very distinct from the lower pseudocortex. The latter has thin—walled. parallel con- glutinate hyphae. The layer is 60‘s thick and gives rise to rhizinae toward the lower surface. 55 The rhizinae are mostly seen in groups and are black and 2—4 p thick. ‘Appppppip: They are scattered on the upper surface and are adnate. The apothecia are black in color with con— colorous disks which are not prominent. They are 1.3— 1.6 mm in diameter and are slightly pruinose. The disks are slightly convex. It seems that in this species one can see the amphithecium and the exciple a little better than in the previous ones but there has been a controversy as to whether the outermost layer is the exciple of amphi— thecium. Both the "layers" are paraplectencymatous. and the only way one can tell one layer from the other is by the difference in the orientation of the hyphae. The hymenium is 40.p wide with club-shaped asci and filiform paraphyses. The eight minute spores in each ascus are round, hyaline, and without an epispore. The spores are monostichous or distichous. The paraphyses end in thick-walled tips forming a very well developed epithecium. 56 Erioderma Fée Essai Cryptog. Ecorc. Officini. 146. 1824. Typp Species: Erioderma polycarpum Fee, Essai Cryptog. Ecorc. Officini. 146, tab. XXXLV, fig. 2. 1824. Species Stpgieg: .p. limbatum, p. microcarpum. Q. physio- ides, and p. polycarpum, and p. Wrightii. ‘ fipperal stcriptipn of the Gepus baged on the Species Sppdied: .gppllpp: The thallus is subfoliose to foliose and is greenish-yellow to gray. The lObes vary between 3.0 mm to 1.5 cm. Thick black rhizinae arise from the entire' undersurface, the lower margins alone, or the undersur— face excluding the margins. All species except p. Igg- :ppppm have a tomentum on the upper surface of the thallus. The lower surface of the thallus is sometimes veined. The upper cortex is paraplectenchymatous, with the hyphae either irregular or parallel to the surface of the thallus. The tomentum is formed by vertical hy- phae originating from the uppermost layer of the cortex. The hairs are unicellular and colorless. The upper cor- tex is 3-many celled thick and sometimes appear to have a "cutiCle." 57 Below the upper cortex is the algal layer. The phycobiont is either Nostoc or Scytonepa occuring in short chains or filaments. The thickness of algal layer is dif- ferent in different species. Below the algal layer is the medulla. This is formed by parallel, almost parallel. or irregular hyphae. These are mostly loose and sometimes are accompanied by special kinds of cells which are moniliform in appearance as in.§. physioiges. The medulla is exposed as there is no lower cortex. The thickness of the medulla ranges be- tween 30-180 it within the genus. Where rhizinae are present. they arise from the lower surface of the medulla. They are black or brown in color and are solitary or in groups. They are non- septate and thin—walled. Apothecia: The apothecia are conspicuous and are marginal or scattered. Eriodepga is the only genus in the Pannaria- ceae with stipitate apothecia. Usually the apothecial disk is brown in color with a light or dark margin. The sides of apothecia are sometimes covered with light tomentum. The apothecia are mycolecanorine. [The outermost layer is the amphithecium which is without any phycobiont.] 58 The amphithecium is paraplectenchymatous. The orientation of the hypahe can be easily followed and it can be estab- 1ished that the amphithecium is developed from the exciple. The proper margin or the exciple is between the amphithecium and the subhymenium. It is quite a thick layer of tissue in the center but its thickness varies in the lateral arms in different species. The exciple, which is quite distinct from the amphithecium is compose of loose and irregular euthyplectenchyma. Some of the hyphae in some species are even pigmented. The subhymenium is not distinct in any species ex- cept §.'Wrightii (Plate 9). In the rest of the species the subhymenium cannot be distinquished from the tissue of the exciple. The hymenium is between 40-100‘p wide. The asci are oblong, and each contains eight spores which are mono- stichous or distichous. The spores are usually one-celled, but two-celled spores occur quite often, and rarely even threewcelled spores can be found; the spores are ellipsoidal to subspherical, hyaline, and can be with or without an epispore. Sometimes they are roughened inside as in p. physipides (Plate 5). 59 The paraphyses are club-shaped and their tips end in an epithecial gell forming an epithecium. The epithe- cium appears quite prominent in some species since the thick—walled tip of the paraphyses take a dark stain. Key t9 the Species Stpdied of Epioderma 1. Upper surface glabrous. Margins sorediate.......limbatum 1. Upper surface tomentose.........................2 2. Rhizinae distributed over the lower surface except at the margins; lobes 1.0-1.5 cm wide.............................Wrightii 2. Rhizinae limited to margins; lobes smaller.....................................3 3. Lower surface not veined. Spheroidal moni- liform cells in the medulla...................physioides 3. Lower surface veined..........................4 4. Veins few, relatively inconspicuous, concolorous with the lower surface of the thallus; apothecia scattered..........microcprpa 4. veins yellowish, abundant, prominent: apothecia mostly marginal..... ....... ......polycarpa 6O Epioderma limbatum (Nyl) Dodge .E. limbatum, Ann. Missouri Bot. Grad. 20:456. 1933. E. Wrightii [Tuck.] var. limbatum Nyl. Flora 52:119. 1869. Matepial §tudiedz U. S. A., Michigan: Chippewa County west of Detour, Im§h§pg 3755, 1949 (MSC). WEST INDIES. Jamaica: Summit of Blue Mt. Peak, Imshaug 13829, 1953 (MSC); East Peak. Imshaug 14820, 1953 (MSC); summit of East Peak, Imshaug 14828, 1953 (MSC); Main Ridge Gap. Imshaug 14750, 1953 (MSC); summit of St. John's Peak, Imshapg 15164, 1953 (MSC); peak on Main Ridge, Imshaug 15210, 1953 (MSC); slope of High Peak. Imshaug 15227, 1953 (MSC); summit of Sugar Loaf Mt., Imshaug 15449. 1953 (MSC); second ridge below summit of Blue Mt. Peak, Imshaug 15554, 1953 (MSC) [studied the ma- terial in detail]; Knife Edge, near Sugar Loaf Mt., Imshaug 15498. 1953 (MSC). Depailed Description: Thallus: The thallus is subfoliose having lobes 0.3—0.5 cm wide. The lower surface has thick black rhizine except on the margins. 61 The upper cortex is paraplectenchymatous with a thick "cuticle" on the upper surface, and is 35 ’4 wide (about S-celled). The algal layer has irregular or slightly coiled filaments of Scytogema [?]. The layer is about 40‘p thick and the algal cells are 5-8,p in diameter. The medulla in this species is wider than in other species. It is composed of loose, irregular and thin hy- phae which become broader towards the lower surface. The layer is about 160‘» thick._ The hyphae in the upper part of the medulla are 4-5‘p wide and in the lower part are 6r7,p. The rhizinae are very abundant covering the entire lower surface except the margin. ‘Appfihpgia: The apothecia are stipitate, mostly marginal and are quite large (1.5-2 mm in diameter) as compared to the thallus lobes. The disks are dark gray to black with the margins eyen darker. The apothecia are mycolecanorine. The amphithe— cium is paraplectenchymatous with the hyphae running out- ward from the internal tissue (the exciple). The hyphal cells are isodiametric. 62 The exciple is composed of a loose irregular eu- thyplectenchyma. Some of the hyphae are thick-walled. The exciple is 90;: wide in the central part and 30 ,u in the lateral arms. The tissue of the stipe is developed below the exciple and inside the amphithecium. It is composed of loose and irregular paraplectxxmymatous tissue. The hymenium is 120‘p wide. The asci are clavate (tapering below). and are 8—10,u wide and 35—40‘u long. There are eight spores in each ascus, arranged monostich- ously or distichously with the latter dominating. The spores are mostly one-celled but are occasionally two- celled. They are without an epispore and are hyaline. They are subspherical measuring 2.5 x 2.5 - 3.0 x 2.5,u. The paraphyses are club-shaped and are 5—10 u wide near the apex. The tips of paraphyses are thickened inside resulting in an epithecium which is more prominent than in the other species. Erioderma micpocarpa Riddle Mycologia 4:134. 1912. The type specimen was collected by C. E. Cummings 189, Herb. Wellesley College. 63 Material Studied: DOMINICAN REPUBLIC. Cordillera Central, (La Vega): Wepmore 3459,;1958 (MSC);Cordillera Central (La Vega), summit of Alto de la Bandera, Imshaug ,2;§§1.l958,(MSO); Cordillera Central (La vega), Imshaug 23471.19581(MSC); Cordillera Central (La Vega), Imshaug 233§3.19583(MSC)- HAITI. Massif de la Hotte. Dept. du Sud., Imshaug 23226. 19.5.3.) (MSG). wss'r INDIES.- Jamaica: Halberstadt, Imshaug 15106.:1958 (.MSC). ' Detailed Description: Thallus: The thallus is foliose with the lobes 4—6 mm wide. The upper Surface is brownish and the lower surface) is whitish or very light. The upper surface is covered ‘with short tomentum. The lower surface is lightly veined. The veins are concolorous with the lower surface. Rhizinae are brown or light in color and they cover the lower surface except the margins. The upper cortex is paraplectenchymatous. The cells are loosely and irregularly arranged. The layer is 30‘p wide. 64 The phycobiont is .Scytonema, and is in the form of short chains. The layer is 40,u wide and the algal cells. which are 8-9 x Syu, can be seen enclosed in thin sheaths. The medulla is of loose hyphae. these are elongated and thin—walled. and are almost parallel to the surface of the thallus, making the medulla 60‘s wide. The lower cor- tex is absent and the rhizinae arise from the lower part of the medulla. Apophecia: They are scattered on the upper surface. The disk is brown with a lighter colored margin. The apothecia are stipitate and measure 1.0 mm in diameter. Longitudinal sections of the apothecia show that they are mycolecanorine. The amphithecium is of irregular paraplectenclma and the orientation of the hyphae is not very clear. The tissue is 20—120.u wide. The exciple is quite a wide layer (Plate 4), 30.” wide laterally and 100.u wide in the center. It is formed of loose euthyplectenchyma Which is vertical to the surface in the sides and parallel to the surface of the apothecium in the center. 65 The stipe is composed of irregularly arranged para- plectendflflmaand is between the exciple above and amphithe- cium surrounding it. The hymenium is about 80‘p wide. The asci are ob— long and are 40—50 )u long and 5-10 p wide. Each ascus has eight monostichous or distichous spores. The spores are ovoid in shape and are 8-12 x 5—8‘p.in size. They are mostly one-celled or rarely 2 or 3-celled. and are hyaline. and do not have a thick epispore. The paraphyses have club-shaped and thick-walled tips forming the epithecium. They are 5—8‘p wide near the surface. Erioderma phygioides Wainio Journ. of Botan. 34:70. 1896. The type specimen was collected in West Indies, St. Vincent, Boxwood by W. R. Elliot 150. Material Studied: U. S. A., Michigan: Mackinac County, Bois Blamdsland, Im- shaug 3261,1949 (MSC); Imshgpq,37;§.l949 (MSC); Chippewa County, Imshaug 3340. 1949 (MSC). 66 WEST INDIES. Jamaica: Summit of Blue Mt. Peak, St. Thomas, Imshauqil3047, 1952 (MSC); summit of Blue Mt. Peak, St. Thomas. Ipphauq 13840, 1952 (MSC); Bellevue to Mt. Rosanna. St. Andrew, Imshaug 14476, 1952 (MSC); summit of Mossmans Peak, St. Thomas, Imshaug 14686. 1953 (MSC); trail of Main Ridge Gap, St. Thomas. Imshaug 14591, 1953 (MSC); ridge between East Pk. and Sugar Loaf Mt., St. Thomas, Imshaug 14890, 1953 (MSC); between High Pk. and Main Ridge Gap, St. Thomas, Imshaquliggg, 1953 (MSC); summit of Sugar Loaf Mt., Imshauq415259,- 1953 (MSC); Knife Edge, near Sugar Loaf Mt., Imphgpg_l§§fll, 1953 (MSC). Detailed Description: Thallus: wide. The thallus is foliose and the lobes are 2-8 mm The upper surface is pale green and the lower surface is whitish. Soredia can be seen on the margins. Tomentum is seen on the upper surface but some areas are without any. The the the are lobes are divided into lobules and the ultimate size of lobules is between 2—4 mm. The thallus is attached to substrate by dense, pennicillate, black rhizinae. which limited to the margins. Only a few can be seen in the central part of the lower surface. 67 The upper cortex (Plate 5) is paraplectenchymatous and is 15—25,u or 2—3 cells wide. The hyphae run parallel to the surface of the thallus. Simple, one-celled, color— less, almost vertical hyphae make up a tomentum which can be seen on the upper surface of this layer. The algal layer is of short separate chains of Nostoc. The chains are vertical or oblique. The layer is 30‘s wide and the Nostoc cells are mostly 5 x 5‘s. There is no difficulty in identifying the phycobiont in this particular species. The medulla is formed by loose, nearly parallel hyphae. They are thin-walled, apparently non-septate. Special kinds of cells, called spheroidal cells by Asahina (1952) and moniliform cells by Culberson (1961), are seen in this layer along with the loose hyphae. The function of these cells is not known. The medullary layer is about 50,» wide and is exposed to the outside as the rhizinae are mostly marginal and there is no lower cortex. The rhizinae are 5-7,u.thick where they are widest. .Apothecia: They are mostly marginal, peltate, and stipi- tate. The disk is dark brown to black in color and the margins are lighter. The apothecia are 1-2 mm in diameter. 68 The upper surface of the apothecia is glabrous but the sides are covered with fine tomentum. The apothecia (Plates 5 and 7) are mycolecanorine. The outermost layer is the amphithecium and it is paraplec- tenchymatous. The hyphae run almost parallel to the surface of the apothecium in the center, turning upwards near the top and downward near the bottom of the apothecium. The outer layer of hyphae which form the one or two-celled to- mentum are thin-walled, colorless, and 10-12.u in length. The amphithecium is about 601p wide. The exciple is only a few Cells wide in the sides but it is 50-60,p.wide in the center. This is formed of irregular euthyplectenchyma. Some of the hyphae are thick- walled and pigmented. The tissue below the exciple and within the amphi- thecium is the stipe. This is formed of loose, irregular, thin—walled, and branched hyphae. The hymenium is 70.” wide and has oblong asci. They are 7-10 )1 wide and 25—30); long. There are eight spores in each ascus and are monostichous or distichous. The spores are ovide to subspherical in shape and measure 4—54p.x S‘p. Generally one—celled spores are found in 69 this species but occasionally two-celled spores can also be seen (Plate 7). Both kinds are not mixed in the same ascus. The spores are hyaline and mfiunxmy roughened with- in with no thick epispore (Plate 7). The paraphyses are filiform. 1.5 - 2‘p wide near the tips, and the tips are thick—walled. The paraphyses form the epithecium. Eriodepma polycarpa Fée Essai Cryptog. Ecorc. Officin., 146. 1824. The type specimen was collected in Cuba, Island of Bour- bone by Aubert Petite—Thouars in 1824. Material Studied: CUBA, Island of Bourbone: Aubert Petite-Thouars in 1824. Conservatoire Botinique. Geneva, Switzerland MEXICO. State of Pueblo. Pringle 12344.719OB-(Mich.); Near Honey Station, Cuyamalaya, Pringle 15578: 1908 (Mich.). Depailed Description: Thallus: The thallus is foliose, greenish-yellow above and pale yellow below, with lobes somewhat pleated. The upper surface is tomentose. The lower surface of the thallus is 7O highly veined. The veins anastomose and are yellowish in color (brighter than the lower surface). The rhizinae are black and abundant and are on the lower surface except the margins. The upper cortex is 20 - 301u thick and is composed of loose paraplectenchymatous tissue which is parallel to the surface of the thallus. There is tomentum on the upper surface as in E. physioides. The hairs are unicellular. straight or curved, colorless, and are 10-20‘u long. There are filaments of Scyponema in the algal layer. These are short, straight or coiled, composed of 3-6 cells. The filaments are not close together. The layer is 30-40;u wide and the Scytonema cells are 5-10‘p in diameter; they are almost round. The medulla is of loose, parallel, thin—walled un— branched hyphae. The layer is exposed to the outside ex- cept where there are rhizinae. The medulla is 50—80‘u wide. There is no lower cortex. The rhizinae are abundant and do not occur on the undersides of the margins. The thallus anatomy is the same as in E. physioides except that here the upper cortex cells are roundish and in g, physipiges they are isodiametric. and here (Q. polycarpa) the spheroidal cells are absent from the medulla. 71 Apothepia: There are more apothecia on the margins than on the rest of the upper surface. They are round and small (about 0.5 mm in diameter). The disk is dark brown with black margins. The stipe is much smaller than in the other species. The upper surface of the apothecia is glabrous. but tomentum is present on the sides. The apothecia are mycolecanorine. The amphithecium is paraplectenchymatous and is highly tomentose on the out- side (Plate 8). The orientation of the hyphae follows the growth pattern. The cells are roundish and loosely arranged. This particular layer is reminiscent of the same layer in ‘g. physioides. The amphithecium here is 50 - 80,u wide. The hairs are 10 - 30‘p.1ong and simple, straight or curved and are colorless. The exciple is not too distinct in the lateral arms but can be recognized easily in the center where it is 10 - 30‘p wide. It is formed of loose, thin—walled, irregularly arranged euthyplectenchyma. The subhymenial tissue is not distinguishable from the hymenium above and exciple below. The hymenium is 30 — 40 11 thick and is composed of ellipsoidal asci and club-shaped paraphyses. The mature 72 asci are 10 u wide and 45,p.long. There are eight subsper— ical, monostichous or distichous spores in each ascus. They are one-celled, hyaline. and without a thick epispore. The tips of the paraphyses are thick—walled and form the pithecium. The paraphyses are 3—8‘n wide near the tips. Erioderma.Wrightii Tuck. Amer. Jour. Sc., Ser. 2, 25:423. 1858. Type collected in Cuba on the tip of Loma del Gato by C. Wright. Material Studied: U. S. A., Michigan: Epippewa County, Imshaug 3341, gggp, gglg, 1949 (MSC); Mackingg,Counpy. Imshaug 3294, 1949 (MSC). HAITI: Massif de la Hotte. Dept. du Sud, Imshaug ggggp, 1958 (MSC); Imshaug 23169, 1958 (MSC); Imshaug _;;g;g, 1958 (MSC). WEST INDIES. Jamaica: Trail of Silver Hill, gripp, 1932 (Mich.); Spring Trial, Chinchoma, Plitt J-45, 1919 (Mich.); Chestervale. Blip; 1-45—13, 1932 (Mich.); Silver Hill Gap, St.Andrew, Imshaug 14085.1953 (MSC); 73 between Clydesdale and St. Helena's Gap, St. Andrew; Imshaug 15106, 1953 (msc). CUBA. Las Villas. Imshguq 24634, 1953 (MSC). COSTA RICA. Guana-Caste, Dodge and Thomas 6983, 1930 (Mich.). Detailed Description: Thallus: The thallus is foliose. pale green in color above and yellowish below (where there are no rhizinae). The lobes are 1.0 - 1.5 cm broad; they are larger than the lobes in the other species studied. There are thick. black rhizinae on the lower surface except on the margins. The upper cortex is of loose irregularly arranged paraplectenchyma(see Plate 9 for thallus anatomy). Tomen— tum is developed from the uppermost layer of the hyphae; they are unicellular and colorless. The upper cortex is 25-40 p.thick. The algal layer is formed of short filaments of Scytonema, irregularly arranged and not so dense as in the other species. The layer is 20-40 h thick and the Scytonema cells are irregular in shape. ranging between 2 x 3 u.— 5 x 6‘u in size. These are typical scytoneme— iform alga as described by Dahl (1950). 74 The medulla is made up of quite loosely woven. thin-walled. irregularly arranged hyphae. The layer is 50—150‘p wide. The lower margins are arachnoid tomen- tose, with groups of tomentum covering the lower margins which are otherwise free of rhizinae. Rhizinae arise in groups from the lower surface of the medulla as there is no lower cortex. The rhizinae are 35-50,p long and 5—7,n thick. Apothecia: The apothecia are marginal, peltate and stipi— tate. The disk is brown or black with lighter margins. The mature apothecia are about 1.5 mm in diameter. The apothecia are mycolecanorine. The amphithe- cium is paraplecuamhwmatous and the hyphae run parallel to the surface in the sides and vertical to the surface below; they are at an angle as they go upward. The de— velopment of the hyphae can be very easily traced back to the exciple (Plate 10). Some of the superficial cells are pigmented. The outer surface is tomentose and the tomentum can be seen in groups forming a sort of network. The amphithecium is 20-70‘p in the sides and 100—200)u below. 75 The exciple is 20—50,u thick in the center and only 7—10‘p thick in the sides. It is made up of loose. thin—walled euthyplectenchymatous hyphal tissue, which is somewhat parallel to the surface of the apothecium. The subhymenium is seen between the hymenium above and the exciple below. The hyphae are thin-walled and loosely arranged. The layer is only about a couple of cells wide. The hymenium is 100-140‘p thick and has oblong asci with rounded ends. They are 4—5 [.1 wide and 30—35 )1 long with eight spores in each. The spores are mostly monostichous, few are distichous and are ellipsoidal. one or two—celled, and with a thick epispore (Plate 10). They are between 4—5 x 3—4‘u. Discussion of Species of Erioderma Studied: Dodge (1933) separated p. physioides and made it the type species for a new genus, Malmella (M. physioides [Vainio] Dodge n. comb). up. physioides should not be re- Inoved from its present position as a species of Erioderma. It is evident from the Plates 4-10, the anatomical simi- larities are so profound within this genus that I cannot 76 help but disagree with Dodge. The similarities and dif- ferences are discussed in detail in Section VIII. Hydrothvria Russ. In Proceed. Essex. Instit. 1:188. 1853. Type Species: .H, venosa.5p§§. Collected in U. S. A., Vermont, in a brook on a rocky sterile soil on slopes of a hill of granite called Bald Mt., near Dummerston by J. L. Russell and C. C. Frost in 1851. ; Spepigg Studied: Hydrothvria venoga, l.c. Material Studied: U. S. A., New York: Ulster County. seepage area along West Neversink Creek, wooded foot of a trail up Slide Mt., 2410 ft. e1ev., §pith S. J. and Milipp A. G. 3371 (NYS). Detailed Description of Hydrothvria Venosa: Thallus: This is the only aquatic genera in the family. The thallus is basically foliose, the younger lobes be- coming almost fruticose as they start growing away from the substratum. It is gray, becoming blackish sometimes. Both surfaces are of the same color, but the lower surface 77 78 is veined. The veins are a little lighter in color and are very prominent. The lobes are subimbricate and pli- cate with round margins. They are 6—9 mm” wide. The rhizinae arise from under the surface of the thallus from the veins, mostly in the middle part of the thallus, form- ing thick tufts. The upper cortex is paraplectenchymatous, the cells are isodiametric and parallel to the surfaces. The layer is about 5 celled and 20-30’u thick. There is a thin cuticle on the upper surface. The algal component is Nostoc (scytonemeiform Nostoc, Dahl, 1950) but the filaments are short, either coiled or straight. Along with many vertical filaments there are occasional horizontal filaments also (as shown in Plate 11). The layer is 60-100‘p,thick and the sizes of the algal filaments and cells vary quite a bit. The filaments are 2—many celled long and the cells are any where from 5x5 )1 - 10x10 )1. i The medulla is of loose, periclinical and thin- walled hyphae, 25-55,u.in.thickness. The lower hyphae in the medulla are septate and are quite distinct from the top portion of the medulla 79 and can be called pseudocortex (Plate 11). This layer is 10-20,p thick and is composed of isodiametric. thin— walled and conglutinate hyphae which run parallel to the surface of the thallus. They are quite different from the upper cortex and hence do not form a true lower cortex. Thick tufts of black rhizinae arise from the cen- tral parts of the lower surface of the thallus; they are several mm long and 8-10‘p thick. Apophecig: The apothecia are marginal, occuring fre- quently in the notches or sinuses of the thallus. They are subsessile. tomentose on the sides and orange to brownish in color. The margins are always darker in color than the disks. The diameter of mature apOthecia range between 1.5 and 1.8‘p. The apothecia are mycolecanorine. The amphithe— cium is 50 p.thick in the sides and 130‘p.in the center and is paraplectenchymatous and tomentose on the sides. The tissue is regular and the hyphae are almost parallel to the surface. The hairs are simple or branched and are straight with pointed tips and 15—25 p long. 80 The exciple is not distinct from the amphithe— cium in the lateral margins but is quite prominent in the center. It is formed by loose and parallel euthy- plectenchyma (Plate 11). The layer is 25-35,» thick. The subhymenium is not distinguishable from the hymenium above and the exciple below. The hymenium is 70-90‘p thick and is formed of linear-oblong asci and filiform paraphyses. A mature ascus is 90,p long and 10,u wide and has eight ellipti— cal spores. The spores are one—celled. hyaline, and without a thick epispore; they are 9-10 x 5,n. The tips of paraphyses are 3-6,n thick, are thick-walled, and form an epithecium Which is about 10 )4 thick . 81 Massalonqia Korb. Syst. Lich. German. 109. 1855. TypeSpecies: Massalonqia carnopg (Dicks.) Korb. l.c. Lichen parposus Dicks. Orig. Coll.: Great Britain, Scot- land. 1790. p. 21. Species Studied: Massaignqia carnosa. Material Studied: U. S. A., Montana: Glacier National Park, over rocks. Virginia Falls, 5000 ft. elev., Imshaug 6456, 1950 (MSC). Detailed Desgription of Massalonqia carnosa: Thallus: The thallus is foliose with imbricate lobes; the lobules are 2-5 mm wide. It is brownish and has globose or cylindrical, marginal isidia. The lower sur- face is lighter in color and brown rhizinae arise from the middle part leaving the lower margins free. The upper cortex is almost uniformly 2-celled and 15-20‘p thick (Plate 13). It is covered with a thin cuticle—like layer. The cells are paraplectenchymatous 82 and pigmented and are vertical to the surface of the thallus. The phycobiont is Nostoc (Dahl, 1950) and the filaments are distinctly visible enclosed in gelatinous sheaths. The algal layer is 25-30‘p thick and the algal cells are 3—5 p.in diameter. The medulla is formed by loosely woven. thin- walled. irregular hyphae and 2—3‘p wide hyphae. The layer is 60-70 ,4 thick. There is a lower cortex present in this species. It is formed of vertical pigmented, paraplectenchymatous cells. This layer is a little thicker than the upper cortex (25-30‘p) but is mostly one-celled in thickness. The lowermost hyphae start growing downwards to form the rhizinae. The rhizinae are not very conspicuous as in the other species of the family. They are septate and brown in color. Apothecia: The apothecia are scattered, peltate. stipi- tate, and concolorous to the thallus with darker margins.- They are about 2 mm in diameter. Tomentum is seen on the sides of the apothecia. 83 The amphithecium is paraplchsxmymatous which is not very regular. Few algal cells are seen in the inner part of the amphithecium making it pseudo-mycolecanorine. The amphithecium is lSO‘p.thick. The tomentum is very sparse, the hairs are 20-30 p long and 5 )1 thick. The exciple is of loose, thin-walled, and parallel euthyplectenchyma. It is very narrow in the lateral margins. about 5 us but in the center it is about 20,u.thick. The subhymenial tissue is not distinct from the hy- menium above and exciple below. The hymenium is 30-60,p thick with clavate to club- shaped asci and club-shaped paraphyses. The asci are 10-12 p.wide and 40-60,p long. There are eight ellipsoidal two- celled spores (three-celled spores are seen occasionally) in each ascus. They are 11~27 x 3-4.5,u and are without a thick epispore. The paraphyses are about 10‘p thick near the tips and are pigmented in this area forming the epithecium. Pannaria Del. in Bory Dictionn. Class. Hist. Nat. 13:20. 1828. Type Species: Pannaria rubiqinosa (Thunberg in Ach.) Del. in Bory. Species Studied: .E~ leucosticta, P. lurida, p. pezizoides. .p. rubiqinosa. Detailed Description of the Genus Ba§ed on the Species Studied: Thallus: The thallus is squamose, subfoliose or foliose and is olive—green to gray on the upper surface and has a lighter lower surface. The rhizinae are abundant and well developed sometimes forming ablack cushion—like structure. The upper cortex is paraplectarhymatous, 4-5 celled thick with irregular or vertical cells. The phycobiont is true nostociform Nostoc and the algal layer as such is quite prominent. There are short chains or coils or Nostoc and the algal cells are mostly spherical. 84 85 The medulla is composed of loose, irregular, thin— walled. narrow hyphae sometimes forming a loose network (arachnoid). The hyphae in the lower part of the medulla are more densely woven than in the upper. The lower cortex is absent and the rhizinae arise from the lower surface of the medulla. They are brown, black, or even pale, and are very dense and prominent. Apothecia: The apothecia are usually abundant, scattered, sessile but constricted below. peltate, orange—brown to dark brown with thalloid margins. The apothecia are lecanorine with differentiation of a cortex and a medulla (the algae are toward the center of the amphithecium). The amphithecium is paraplectenchy- matous and sometimes has a tomentum developing from the outermost hyphae° The amphithecium completely encloses the disk on the sides. There are vertical or parallel filaments of Nostoc which are not so dense as in the thallus. The exciple is paraplectenchymatous or enthyplecten- chymatous. It is colorless, narrow in the sides and wide in the center and quite distinct. 86 There is no hypothecium and the subhymenium is not distinct from the hymenium above and exciple below. The hymenium is composed of mostly clavate asci and filiform or club-shaped paraphyses. The surface of the hymenium is flat with the amphithecium coming little above the thecium on the sides. The asci are lO-ll/u wide and 55-75‘p.long with eight spores in each of them. The spores are elongate to ellipsoidal or subspherical. hya— line. one-celled and without an epispore, they can be monostichous or distichous. The tips of the paraphyses sometimes form and sometimes do not form an epithecium. Even when the epithecium is developed it is not as con- spicuous as in some of the other genera. Pannaria leucosticta Tuck. in Nyl. Annal. Scienc. Nat., Bot., Ser. IV, 12:294. 1859. Material Studied: U. S. A., North Carolina: Durham County; in Duke Forest near Durham. On Quercus rubgg. Culberson 4806. 1958 (MSC). 87 Detailed Description: Thallus: The thallus is subfoliose, subimbricate, olive- green and marginally sorediate. The lobules are 0.5—1.0 mm wide. There are black rhizinae on the entire lower surface. forming a network which looks like a hypothallus. The upper cortex is paraplectenchymatous and is about 32‘p thick. The cells are almost vertical to the surface. The algal layer is composed Of definite coils of Nostoc and is about 65‘p,thiCk. The medullary hyphae are arachnoid and the layer is 80‘p thick. The hyphae are thin-walled, colorless, and are loosely arranged. The rhizinae arise from the lower surface of the medulla and form a conspicuous network. Apothecia: The apothecia are scattered, orange-brown with light colored (concolorous to the thallus) pruinose margin. They are peltate, and constricted below and are 0.3—0.9 mm in diameter. The amphithecium is 160‘n thick and is paraplec- tenchymatous and has the algae dispersed almost in the 88 entire thickness of the amphithecium. It exceeds the sur- face of the hymenium and is seen like a rim on the sides. There are dense coils of the phycobiont as in the thallus. The exciple is composed of conglutinate, parallel euthyplectenchyma and is about 32‘s thick in the sides as well as in the center; it is light brown. The hymenium is about 95 ;u thick and is almost flat surfaced. There are clavate asci with eight ellipsoidal, hyalline, thin-walled. one-celled spores in each ascus. The spores are mostly monostichous but distichous spores can be seen too. They are 13.8 x 9.0,» in size. The para- physes are filiform with thickened tips forming an epithe- cium. Pannaria iurida (Mont.) Nyl. Memoir. Soc. Sc. Nat. Cherbourg 5:109. 1857. Collemg luridum Mont. Ann. Sc. Nat. Bot. II, 18:266. 1842. Mapgpigl Studied: U. S. A., North Carolina: gipyen COunty; near Havlock, about 15 ft. up the trunk of a large Quercus gai- 93p; felled in the woods during hurricane, Culber- son 10373, 1960 (MSC). 89 Qetailed Description: Thailus: The thallus is foliose, gray and pruinose with no soredia or isidia. The lower surface is whitish. The lobes are imbricate and the lobules are 2-4 mm wide. The rhizinae are pale to brown and do not develop on the lower margins. They are about 1 mm long and do not form a net— work. The upper cortex is paraplecflxmhymatous, 4—5 celled or about 23,p thick. The cells are uniform and parallel to the surface of the thallus (Plate 15). The algal layer is 65,n thick and is composed of mostly vertical filaments of the phycobiont (Nostoc). the cells of which are 3—7 x 3—6,p. The filaments are not so dense as in g. leucosticta. The medullary hyphae are thin-walled, colorless. loose and form a network. The layer is 90,M thick and the hyphae are 1.5-2.5 p wide. The rhizinae are abundant and pale or brown arising in tufts. They do not form a network or cushion—like struc- ture as they do in g. leucosticta. 9O Apophecia: The apothecia are scattered, orange—brown, pel— tate and constricted below. The margins are pruinose and concolorous with the thallus. Many apothecia become irreg— ular in shape through crowding. The amphithecium is paraplectenchymatous and is differentiated into a cortical tissue and an inner layer with the algae, the latter come almost in the cortical tissue near the upper tip of the amphithecium. The cells are regular and uniform in size and Shape and the whole layer is about 80 51 thick (Plate 16) . The exciple is paraplectenchymatous and is uniform in thickness on the sides and in the center, it is 20‘p thick. The subhymenial layer is not distinguishable. The hymenium is 120,p thick and has clavate asci which are about 55 u long and 10 In wide. There are eight ellipsoidal. one—celled, thin-walled, hyalline, monosti- chous spores in each ascus; they are 8-10 x 3—5,u in size. The paraphyses are filiform and have slightly thickened tips forming a very poorly developed epithecium about 10,u thick. 91 Pannaria pezizoides (Wever) Trevis. Lichenotheca Veneta, 98. 1869. Liphpn pezizoides Weberi. Spic. F1. Goet. 200. 1778. Matppiai Studied: CANADA. Britisp Columbia: Yoho National Park. Twin Falls. on a mossy soil bank at west of trail is spruce- fir forest, pamb 6266, 1951 (MSC). Dgtgiigd Description: The thallus is squamose. The squamules are minute (0.1—0.2 mm wide). densely imbricate and brown with a smooth surface. The upper cortex is paraplectenchymatous and is 20‘» (3-4 celled) thick and is light brown. The algal layer. almost 65‘p.thick, is composed of dense coils of Nogtoc, which are made up of almost spherical cells 2.0 x 2.0 A1 in size. The medulla is composed of thin—walled, irregular. loose hyphae. and the whole layer is about 35.M thick. The lower cortex is absent. The rhizinae are black and shiny and form a narrow network appearing like a hypothallus, but not as well de- veloped as in Anzia or PannOparmelia. 92 A o ia: The apothecia, scattered on the upper surface. are usually numerous and brown with the margin concolorous with the thallus. They are peltate and sometimes become irregular in shape through crowding, but a normal spherical apothecium is about 1.2-1.5‘p.in diameter. The amphithecium is paraplectenchymatous and has (the algae in the central area where the amphithecium is differentiated into a cortical and medullary tissue (as in g. lprida). The amphithecial cells are regular and uniform in size and the layer is about 105.fl thick. The exciple is paraplectenchymatous, 10 )4 thick in the sides, 70‘p thick in the center. and it is light brown. The hymenium is composed of clavate asci which are about 75.0x3.0.u in size, with eight one-celled. ellipsoid (15—25 x 8—13,p) hyalline spores which are monostichous or distichous and without a thick epispore. The paraphyses are club-shaped with pigmented and thick—walled tips forming a very well—developed epithecium 20,u thick. It is best developed of all the species of ngnapia studied here. 93 Pannaria rubigippsa (Thunb.) Del. In Bory Dictionn. Class. Hist. Nat. 13:20. 1828. Lichen rupiginpsus Thunb. in Ach., Lich. Suec. Prod. 99, 1798. Mgpepial Stgdigd: U. S. A., North Carolina: Near Havlock, Graven County. Culberson 1023;, 1960 (MSC). Depailed Description: Thaiius: The thallus is foliose, olive green and minutely granulose in certain areas. The lobes are imbricate. waved and have crenate margins. they are 0.8 - 1.5 mm.Wide. The lower surface is light or dark and is covered with pale or black rhizinae on the entire lower surface. The rhizinae are 0.1 — 0.2 mm long and do not form a network or cushion- 1ike structure. The upper cortex is paraplectenCMatous, 13-25 M (4—5 celled) thick and the cells are vertical to the sur- face of the thallus. The algal layer is about 103.5‘p.thick and is com- posed of dense coils of Nostoc cells which are 4.8 x 3.2‘p in size. 94 The medulla is composed of loose, thin-walled al- most parallel hyphae which make the layer about 701u thick. The rhizinae arise from the lower surface of the medulla, the lower cortex being absent. The rhizinae are about S‘p thick and are either pale or brown. Apophecig: The apothecia are orange—brown to reddish brown. 0.6 w 1.5‘p.in diameter and are scattered. The margins of the apothecia are concolorous with the thallus and are some- what crenate. The amphithecium is paraplecusmhymatous and is dif- ferentiated into a cortical layer and a medulla, the algal filaments and coils are in the inner area (medulla). The outside of the amphithecium is tomentose. the latter being colorless and dense. The exciple is of irregular paraplectenflyma and is about 13 41 thick in the sides and 62 A thick in the center. It is brownish and quite distinct. The hymenium is about 131,u thick and is composed of clavate asci which are 62,u long and l3.8.u.thick (in the center). There are eight one-celled spores in each ascus. they are ellipsoidal, 14—26 x 6-11,u, hyalline. and are arranged in one or two rows. 95 The paraphyses are filiform and their tips form a brownish epithecium which is 20.5.p thick but is not so conspicuous as in p. pezizoides. Parmeliella Mfill. Arg. Memoir Soc. Phys. Hist. Nat. Geneve 36:376. 1862. Typg Species: Parmeliella tripppphylla (Ach.) Mflll. Arg. Lg. corallinoides (Hoffm.) Zahlbruckner]. Speciep Studied: ‘g. lepidiopa, g. micpophylla, g. pannoga. .3. plumbea. Detailed Description of the Genus based on the Species Studied: Thallus: The thallus is squamose to foliose. the tips be- coming almost foliose even if the lobes are subfoliose or squamose. The squamules or lobes are quite minute and are brownish-gray. There are abundant black hyphae on the un- dersurface of the thallus forming a thick mat and extending beyond the thallus as a black margin. This probably is a prothallus; Dodge (1933) called it a hypothallus. The upper cortex is paraplectenchymatous and is 2- many celled or 10—60|p thick. The hyphae are vertical or almost vertical to the surface of the thallus. 96 97 The phycobiont is Nostoc and the thickness of the algal layer differs considerably in the different species. The Npptoc filaments are densely or sparsely arranged and are mostly vertical to the surface of the thallus. Some- times there are regular coils of Nostoc filaments as in.§. plpppea (Plate 19). The medullary hyphae are thin-walled, loosely ar- ranged, and irregular or parallel to the surface of the thallus. There are conglutinate, thin- or thick—walled hyphae below the medulla in some species. This tissue is called the "prothallus." (also see pp. 18-21). The pro- thallus is 5-30‘p thick in different species. The rhizinae arise from the lower hyphae of the prothallus and are seen on the entire lower surface of the thallus. They are black. thickvwalled and very conspicuous. Apophecia: The apothecia are scattered, peltate, sessile or constricted below, light to dark brown in color with margins concolorous with the thallus. They are quite abun- dant and conspicuous. their diameter ranging between 0.5- 1.2 mm. The apothecia are_mycolecanorine and the amphithe- cium is para or prosoplectenchymatous. The sizes of the 98 hyphae and the thickness of the layer differs in differ- ent species. The exciple is distinct and varies in thickness in different species. In 3. pannosa the hyphae below the exciple are deeply staining suggesting the presence of hy— pothecium. The excipular tissue is loose euthyplectenchyma becoming parallel in the lateral margins. The subhymenium is not distinct from the hymenium above and the exciple below. The hymenium has linear to oblong asci and paraphy- ses with clubeshaped tips. There are eight ovoid to ellip- tical, one-celled, mono- or distichous spores in each ascus. The spores are hyalline and are with or without a thick epispore. The tips of the paraphyses may or may not be thick— walled and pigmented, hence the epithecium is developed or undeveloped. Parmeliella lepidiota Wainio Termeszetr. Fuzetek, 22:308. 1888. Maperial Studied: CANADA, Albepta: On ridge opposite Angel Glacier on Mt. 99 Edith Cavell, Jasper National Park, 6500 ft. elev., Imshaug 6954, 1950 (MSC). Detailed Description: Thallus: The thallus is squamose. the squamules being 0.2—0.3 mm wide and black. The external morphology of the thallus is very reminiscent of Placynthium nigrum. There are black rhizinae on the entire lower surface of the squamules. The upper cortex is paraplectenchymatous and is 60‘p thick. The hyphal tiSsue is almost parallel to the surface of the thallus. The algal layer is 90‘p.thick and is composed of regular ensheathed coiled filaments of Nostoc. The medulla is formed of thin-walled, irregular hyphae, the layer being 160‘p thick. The hyphae are 3—5‘p thick. There is no prothallus in this species and the rhizinae arise from the lower hyphae of the medulla. The rhizinae are black and about 5,u.thick. Apothecia: The apothecia are scattered on the upper surface. They are dark brown or black with the same colored margin 100 and are sessile and the disk is convex. The diameter of the apothecia varies between 0.5-0.7 mm. The anatomical studies show that the apothecia are mycolecanorine, with a very well-developed epithecium (more than in any other species studied here). The amphithecium is prosoplectenchymatous and is about 85‘p.thick. The exciple is of loose, irregular euthyplectenchyma and is 58,p.thick. The hymenium is 60‘p thick and is formed by clavate asci and filiform paraphyses. There are eight one-celled. thick-Walled, hyalline spores arranged monostichously or distichously in each ascus. The asci are 60-80,u long and 10-15 )1 wide. The spores are 8—10 x 5—8 p and ovoid. The tips of the paraphyses are thick—walled and pigmented as to form a very well-developed epithecium which is about 10,u thick. Parmeliella microphylla (Sw.) Mfill. Arg. Flora 72:507. 1889. Lichen microphyllus Sw. Vetensk. Akad. Handl. 301. 1791. 101 Material Studied: U. S. A., Michigan: Keweenap_County: Isle Royal National Park, along southwest shore of West Caribou Island, Wetmore 4416. 1959 (MSC). Depailed Description: Thallus: The thallus is squamose, the squamules 0.1—0.2 mm wide, sorediate, and gray. There are black rhizinae on the lower surface. The thallus morphology, like 3. lepidiota. reminds one of Placynthium nigrum. The upper cortex is paraplectenchymatous and 3-4 celled or lo‘p thick. The cells are pigmented and parallel to the surface of the thallus. This layer in g. micrOphylla (Plate 18) is narrowest of all the species of Papppliella studied here. The algal layer is 80‘p thick and is composed of vertical filaments of Nostoc, which are very compactly ar- ranged in almost the whole thickness of the thallus, thus reducing the size of the medulla to nil. This is similar in Placynthium nigrum (Plate 21), and different from other species of Parmeliella studied here. The Nostoc cells are 102 There is a paraplectenchymatous hyphal tissue be— low the algal layer which I suggest should be called the pseudocortex. It is 2-3 celled or 8—9,u thick. The rhizinae are very inconspicuous and arise from the lower hyphae of the pseudocortex mostly from the lower margins of the squamules. Apothecia: The apothecia are scattered, black with margins concolorous with the thallus and are sessile. The disk is 1.0-1.5 mm in diameter and is peltate. The amphithecium is prosoplectenchymatous with the outer cells pigmented. The layer is about 70,».thick. The exciple is formed of irregular euthyplecten- chyma. The tiSsue is very prominent in the central area but is not so much developed in the lateral margins; it is 60 n thick. The subhymenial tissue is not distinguishable from the hymenium above and exciple below. The hymenium is 80‘» thick including the epithecium. There are linear to oblong asci which are 35-45,» long and 5;: wide. There are eight one—celled ovoid monostichous or distichous spores in each ascus. They are hyalline and with- out a thick epispore (unlike 2. lepidiota) and are 5-6 x 41h. 103 The paraphyses are unbranched and linear with club-shaped tips. The tips of the paraphyses are pig— mented and thick-walled forming an 8‘p thick epithecium. Parmeliella pannosa (Sw.) Mfill. Arg. Flora 64:86. 1881. Lichen pannosus Sw., Nov. Gen. Sp. Pl. Prodr. 146. 1788. Matppial Studied: U. S. A., Hawaii: island of 0ahu in Nfiflanu, on Han tree. Heller A. A. 2107, (1895 (MSC). Depailed Description: Thalius: The thallus is subfoliose, the lobes being 0.5-1.0 mm wide. and imbricate. It is brownish and the dark prothallus extends about l-l.5 mm beyond the thallus forming a black margin. There are black, conspicuous, well develOped rhizinae on the entire lower surface of the thallus. The upper cortex is paraplectenchymatous. The cortical cells are not uniform in size and are vertical to the surface of the thallus. The cortex is several- celled and 32—35 p thick. 104 The algal layer is very thin as compared to the algal layer in other species of iParmeliella studied here, being only lz‘p.thick and containing short filaments of Nostoc mostly seen as coils which are not densely packed. The algal cells are 5x3-4p in size. There are thin-walled, loose, almost parallel hy- phae in the medulla making it about ZO‘prthick. There are more conglutinate, parallel and black hyphae below the medulla which form the prothallus. It is 15/u thick below the medulla. The rhizinae arise from the lower part of the pro- thallus. They arise in groups on the entire lower surface of the thallus and are black and quite conspicuous. The rhizinae are 3—5 11 thick. Apothecia: The apothecia are scattered on the upper sur— face. They are mostly brown or infrequently black with margins which are concolorous with the thallus. The apo- thecia are sessile, peltate, and 0.5—1.0 mm in diameter. The amphithecium is of irregular paraplectenchyma. It is 70‘u thick and the orientation of the hyphae clearly shows its development from the exciple. 105 The exciple is ls‘p.thick and is composed of euthy- plactenchyma. It cannot be distinguished very well from the amphithecium in the lateral margins. There is a layer about 18‘p thick of pigmented, thick-walled cells above the exciple and below the hymen- ium which probably is the hypothecium. This tissue is absent in all the other species of the family Pannaria- ceae that are studied here. The hymenium is 45‘p.thick and has clavate asci which are about SS‘p.long and 10.u wide. There are eight oblong, 12—15 x 4-5‘p.spores in each ascus. They are thin— walled, one-celled, and hyalline. The paraphyses have club-shaped tips, 2.5-3.0‘n wide which form the 3,n thick epithecium. The paraphyses reach the surface of the apothecium, and even extend a little above it. Parmeliella plumbea (Lightf.) Mull. Arg. Bullet. Herbier Boissier. 2 (appendix I): 44. 1894. Lichen plumbeus Lightf., Flora Scotica 2: 826. 1777. Material Studied : SCOTLAND, Argyll: shores of Lake Leven near Glen Coe, 106 Weber W. A. angvgrundwell A. C. S; 14300, 1958 (MSC). Detailed Description: Thallus: The thallus is foliose, grayish, and has scattered cylindrical isidia and orangish soredia. The thallus lobes are imbricate, 2-4 mm wide and pruinose. The prothallus ex- tends about 1 mm beyond the thallus lobes as a black margin. sometimes appearing like a cushion of hyphae. The rhizinae are abundant, black, and 2.5 mm long. Sometimes they are not seen under the slightly involute margins of the lobules. The uppercortex is paraplecflnchymatous (Plate 20), 20,p.thick and the cells are almost vertical to the surface of the thallus. The algal layer is SO‘p thick containing Nostoc filaments which are either coiled, vertical, or horizontal. The Nostoc cells are 4-5 x 6—8,u. The medulla is 80,u thick and is composed of thin- walled, almost parallel hyphae. There are conglutinate and black hyphae below the medulla which form the prothallus which is 20,u thick and form a mat of hyphae which appears almost cushion-like in the marginal areas at the tips of some lobes. 107 The rhizinae arise from the lower part of the pro- thallus and are black and cover the entire lower surface except some marginal tips. Apothecia: The apothecia are scattered on the upper surface. They are dark brown with the same colored margins and are 0.8-1.0 mm in diameter. The disks are peltate and glabrous and are constricted below. The apothecia are mycolecanorine, the amphithecium being prOSOplectenchymatous and about 60 )4 thick. The exciple is 20‘p thick in the lateral margins and is 804p thick in the center. It is composed of euthy- plectenchyma and is colorless. The subhymenium is not distinct from the exciple below and the hymenium above. The hymenium is 70—100,u thick and has oblong asci which are 35 u long and 8,u wide. There are eight ovoid, one-celled, 4—5 x 4‘p, hyalline, each with a thick epispore. They are monostichous or distichous, often within the same apothecium (Plate 19). The paraphyses are either fusiform or with club- shaped tips. Their tips are not pigmented or thick~walled in most cases, hence an epithecium is not differentiated. Placynthium Gray A Natural Arrangement of British Plants, 1:395. 1821. Type §pecie§: Placynthium nigrum Gray, l.c. Specigs Studied: Placynthium niqum. Material Studied: U. S. A., Michigan: Leelanau Counpy: along the shores of Lake Michigan at Lighthouse Point, Imshaug 19481, 1958 (MSC). Depailed Description of Placypthium nigrum: Thallps: The thallus is crustose but broken into angular areoles. Each areole is composed of several granules. The granules are 0.5~1.0 mm in diameter. The thallus is dark olive-brown, appearing black. It is attached to the sub- strate by conspicuous rhizinae Which are black. The thallus anatomy is discussed in Section III in detail but some of the important things are repeated here. A thallus section shows that the granules are rather inter- mediate between being homoiomerous or heteromerous. There is a one-celled paraplectenchymatous cortical layer seen 108 109 as the outermost layer of the granule (Plate 21). The thickness of the granule is occupied by the phycobiont and fungal hyphae. The medulla is not seen as a definite layer, since the filaments of Nostoc are dispersed irregu- larly in the entire central region. The Nostoc filaments are 2-6 cells long and are sometimes in coils. The algal cells are 2-3 x 3.5 - 4%». The filaments are mostly ver— tical. The rhizinae arise from the lower surface of the areole and are black and conspicuous. Apophecia: Apothecia are scattered on the upper surface. They are concolorous with the thallus, 0.2-0.3 mm in dia- meter and with promenent black margins. The apothecia are mycolecanorine, constricted be— low and with straight surface. The amphithecium is para— plectenchymatous, and the orientation of the hyphae clearly shows that the ampthithecium has developed from the exciple (Plate 22). The tissue of amphithecium is about 15‘p thick laterally and about 90,p.toward the base. The exciple is composed of thin-walled, loose and almost parallel euthyplectenchyma. It is a very thick llO tissue (about 100‘p) in the center of the apothecium and very narrow (20—40‘p) in the lateral margins. Among the species of the Pannariaceae that have been studied here, this species seem to have a unique exciple. The hymenium is composed of linear aSCi, pointed at both ends, and filiform paraphyses. The mature asci are 32-35‘p long and.4—5‘p wide. There are eight spores in each ascus. The spores are one—celled or two-celled with both kinds equally common in the same apothecium. They are elliptical, monostichous (arranged side to side slanting in one direction), thin-walled, and hyalline. They are 5-7 x 3-4fin size. The paraphyses are without thickened tips hence an epithecium is not differentiated. The tips of the paraphyses are 2-5 )1 wide. Psoroma Nyl. Mem. Soc. Sci. Nat. Cherb. 3:175. 1855. Type Species: Psoroma Hyphoppm (vahl). S. F. Gray Species Studied: Psoroma hyphorum, and Psoroma pallidum. Detailed Description of the Genus Based on the Species Studied: Thallus: The thallus is squamose, with minute, glabrous, grayish squamules, having scattered white soredia. The rhizinae are pale and absent from the lower margins, they are few and inconspicuous. The upper cortex is paraplectenchymatous which is usually several cells thick and the cells are someWhat ir— regularly placed. The algal layer is usually narrower than the upper cortex and the medulla. The algal filaments are not so densely packed as in the other genera of the family. The alga is Doctylococcus or more probably Coccomyxa (a green alga). lll 112 The medulla is the widest layer and is composed of thin—walled loose hyphae which form a network or are irregular. The lower part of the medulla is composed of hy- phae which form a kind of tissue which is different from the medullary tissue and is not like upper cortex either. This, then, is the pseudocortex. The rhizinae arise from the pseudocortex and as numerous tufts. They are thin-walled and usually non- septate. Apothecia: The apothecia are scattered on the upper sur- face. The disk is concave and the margins are crenate and concolorous with the thallus. They are adnate to sessile. The apothecia are mycolecanorine, the amphithecium is or is not differentiated into a cortex and a medulla. The exciple is of loose and parallel euthyplace— tenchyma and colorless. The asci are linear to oblong and have eight hya- 1line, thin—walled, monostichous or distichous spores which are usually ellipsoid to almost spherical. The 113 paraphyses are filiform or with club—shaped ends, they do or do not form an epithecium. Psoroma hypnorum (Vahl.) Gray. A Natural Arrangement of British Plants, 1:445. 1821. Lichen hypnorum vahl., Icon. P1. Dan. fasc. 956, 1787. Parmelia hypnorum (vahl.) Ach. Pannaria Hypnorum (vah1.) Koerb. Material Studied: U. S. A., Colorado: Jackson County: 3 miles north of Willoe Creek Park, Rabbit Ears Range, Shushan S—840l, 1956 (MSC). Detailed Description: Thallus: The thallus is grayish—green, squamose, the squamules are minute and imbricate with rounded margins and are smooth with occasional globose soredia. The rhi- zinae are pale and few and are not very conspicuous. The upper cortex is paraplectenchymatous, 4-5 cells (50,u) thick with irregular cells. The algae filaments are quite conspicuous (more than in the thallus of g. pallidum). They are 2—4 cells long and are placed irregularly, the algal cells being 2-4 x 2-5‘p in 114 size. The algal layer is 404p thick (a little less than in g. pallidum). The medulla is composed of thin—walled, loose ara- chnoid hyphae, 60 p.thick. There is no pseudocortex below the medulla as in g. pallidum and the rhizinae arise from the lower part of the medulla. The rhizinae are not pres— ent on the lower margins and when present are not very conspicuous. However, below the apothecia they are some— times arachnoid. Apothecia: The apothecia are reddish brown, 1.5-2 mm in diameter with a concave disk. The margins are crenate and concolorous with the thallus. The margins are involulte so as to cover the disk to a great extent. The apothecia are lecanorine and the amphithecium is not differentiated into a cortex and a medulla. The algae are dispersed through the entire thickness of the amphithecium, which is 225,u.thick. The amphithecium is paraplectenchymatous and short algal chains are found along with the fungal tissue. The hyphae run parallel to the surface of the apothecium. On the sides the amphithe- cium is tomentose (Plate 24). The tomentum is dense com- posed of hairs which are short, one-celled, and colorless. 115 The exciple is euthyplectenchymatous. The tissue is loose and composed of parallel hyphae which are thin— walled and hyalline. It is of equal thickness laterally as well as below the hymenium and is 20‘» thick. The hymenium is composed of clavate asci. There are eight ellipsoidal one-celled, monostichous or distich- ous hyalline spores in each ascus. The spores are 15-22 x 7—10‘p in size and are without a thick epispore. The paraphyses are filiform. Their tips are not pigmented or thick-walled, hence an epithecium is not de- veloped here (Plate 24) as it is in g. pallidum. Psoroma pallidum Nyl. In Annal. Sc. Nat., Bot., Ser. 4, 12:294. 1859. Material Studied: Misplaced after making the slides. Collection data: not available. Detailed Description: Thallus: The thallus is squamose, the squamules are 0.1- 0.3 mm wide, they are almost corolloid and grayish on the upper surface and pale on the lower. The thallus is gla— brous and has occasional scattered, globose, white soredia. 116 The rhizinae arise from the lower surface leaving the mar- gins free; they are pale and inconspicuous. The upper cortex is paraplchanhymatous but the cells are irregularly arranged. It is almost 10 cells (about 804p) high. The algal layer is about 35,p.thick and is composed of spheroidal cells of probably Cocgpmyxa. The algal cells are surrounded by a gelatinous sheath and are usually united in colonies of indefinite extent, which are not very densely packed. The algal cells are 5-10 x 5-81p.in Size. The medulla is composed of thin-walled, arachnoid hyphae, which are loosely arranged. The layer is 120‘p thick and is the thickest layer in the thallus. The lower part of the medulla is cellular and 30.“ thick; it appears to be a pseudocortex. I The rhizinae are not very conspicuous, and arise from the pseudocortex. Apothecia: The apothecia are scattered on the upper sur- face, and are minute as compared with those of P. hypnorum. The disk is brownish, flat, and the margins are concolorous with the thallus and crenate. The apothecia are adnate. 117 The amphithecium is differentiated into a cortex and a medulla and the algae are found in the middle areas. The amphithecium is paraplectenchymatous and 160‘p thick. The sides are not tomentose as in §.'hypnorum. The exciple is hyalline and is formed of loose. parallel, thin—walled euthyplactenchyma, 10 p thick in the sides, and 40‘p thick below the hymenium. The hymenium is composed of linear asci which are 70—90 x 12-18,u. They contain eight one-celled, ovoid to spherical, hyalline spores. 'The spores are mostly monosti- chous and are without a thick epispore. They are 5-8‘u in diameter. The paraphyses are linear, unbranched, and non- septate. They have club-shaped tips, forming an epithe- cium 10,p thick. VIII DISCUSSION Lichens like any other organism can be classified in many different ways. Ever since these organisms at- tracted the attention of naturalists, they have been placed in different systems of classification, based on external characters as well as the anatomy of the thallus and apothecium and even the phycobionts. Most of the time the classification was quite artificial (classification based upon characters of convenience without relation to phylogenetic significance or classification based on a single arbitrarily chosen criterion, instead of an evalu— ation of the totality of characters). Zahlbruckner (1912) was the first to propose a satisfactory natural classification. Our present knowledge of the lichen ascocarp makes the development of the lichen thallus of secondary taxonomic importance (Santesson, 1954). The micro characters of the fungus in the apothecia, e.g., 118 119 spore septations, nature of the exciple, nature of amphi— thecial tissue, are very important and must be considered if one has to come to a meaningful phylogenaic grouping of the species. The micro characters of the thallus, e.g., distribution of the algae, presence or absence of a lower cortex, and nature of medullary hyphae, etc., can be of assistance along with the micro-characters of the apothe— cium. The characters of the phycobionts are more important in identification than in classification. Culberson (1961) in discussing lichen nomenclature proposed, "The algal and fungal components of a lichen shall not be considered dis- cordant elements." Thus, emphasizing the importance of considering both algal and fungal components together, and implying that a lichen thallus is, biologically, something more than just two plants living together. Santesson (1950) emphasized that we have to deal with the lichenized fungi alone and not with the lichen consortium when it comes to taxonomy of lichens. He said that "lichen systematics" based on algal characters is unnatural. My own studies suggest that phylogenetic relationships and development throughout the family Pannariaceae are shown.lwflthe development of mycobiont tissue. I have not been able to 120 see any characters of the phycobiont which can be considered primitive or as as highly advanced. Naturalness of the familnyannariaceae: The family Pannariaceae except for the genus Placyn— thium, seems to be a natural group since phylogenetic rela— tionships can be seen among the genera and a gradual evolu- tion from a less developed to a more developed structure or tissue is quite noticeable. The family probably is of poly— phyletic origin and the genera have taken different lines of development. The following characters in the remaining genera show some probable phylogenetic relationships (less highly advanced character given first): 1. The spores range between one to two and three- celled. The exciple is from loose to conglutinate euthy- plectenchyma. The amphithecium is prosoplectenchymatous to paraplectenchymatous. The apothecia are lecanorine to mycolecanorine. The thallus is squamose, subfoliose to foliose. The thallus ranges between a less developed het- eromerous (Parmeliella microphylla) to a highly 121 developed heteromerous (Massalongia carnosa) type. 7. The presence of prothallus in some species and its absence in the others. Placynthium should be removed from the Pannariaceae, because it has many characters (as seen in the type species _2. nigrum) which are different from the other genera studied here. They are as follows: 1. The exciple is cup-shaped as compared with a saucer-shaped exciple in the other species. 2. The cortical layers are one-celled thick rather than 2—many celled in the other genera. 3. The medulla is absent, with the whole thickness of the areole being occupied by algal filaments. 4. The thallus is squamose-areolate, rather than squamose, subfoliose, or foliose. Dahl (1950) also suggested that Placynthium should be removed from the Pannariaceae and be placed in a family of its own-—P1acynthiaceae. He included other genera in the Placynthiaceae (see p. 10). He separated the two families, Pannariaceae and Placynthiaceae, on the basis of two dif- ferences. 122 Dahl considered the Pannariaceae to have only a nostociform phycobiont and the Placynthiaceae to have only a scytonemeiform phycobiont. He did not consider Psoroma at all which has a green phycobiont. In Coccocarpia the phycobiont is chtonema (according to Bornet's 1909 fig- ures). Dahl suggested that Coccocarpia be removed from Pannariaceae, though he did not assign it a definite fam- ily. If keyed out according to his key it comes under Placynthiaceae. My studies of the anatomy of the apothe— Acium of Coccocarpia show that the apothecial tissues are similar to the other genera, hence Cpcgocarpig should not be removed from .Pannariaceae simply on the basis of its having scytonemeiform alga. It has not been determined whether the phycobiont of Erioderma is Scytonema or Nostoc or both. Dahl (1950) suggested that Erioderma has scytonemeiform alga but since its morphology and anatomy are different from the members of the Placynthiaceae, it should be put into a family of its own. The characters of the apothecial tissues of Epig- Q§£E2 are closer to the other members of the Pannariaceae and therefore I suggest that Erioderma be left in its present position. 123 The second criterion that Dahl gave for the separa- tion of Pannariaceae and Placynthiaceae was the orientation of the algal filaments. I found that there are both verti- cal and horizontal filaments of the phycobiont in the thal- lus of Placynthium nigrum and other species. The horizontal filaments are quite prominent in Qgggggarpi§,but then they are equally so in Hydrothvrig, which was placed in the Pan- nariaceae. Choisy (1960) recognized the Placynthiaceae as a separate family with Placynthium as the type genus. Fur- ther, he considered the Pannariaceae to contain just one genus, Pannaria, with the other genera formerly included here as sections and subsections of it. I consider the morphological characteristics of these genera support their being kept at the generic level. Naturalness of the genera: One of the main characters by which genera of Pan- nariaceae are identified is the presence of a green or blue-green phycobiont. Only two genera, Psoroma and Egg;- omaria, possess a green phycobiont, but their mycological characteristics bind them together with the other genera in the Pannariaceae. 124 The type genus, Pannaria, has the following char- acters: l. Thallus squamose to foliose. 2. Thallus almost heteromerous (lower cortex absent). 3. Upper cortex paraplectenchymatous. 4. Phycobiont is blue-green. 5. Medulla of loose, arachnoid hyphae. 6. The outermost layer of the apothecium is the amphithecium.. 7. The exciple is distinct. The characters mentioned above are seen with some variations in the other genera studied here. Some of the distinct differences are mentioned below. Hydrgthvria is an aquatic genus hence one would expect some morphological specializations of the thallus such as, shield-like thallus lobes, prominent veins, thal- lus becoming frutiose as it grows away from the substrate. etc. However, the anatomy of the apothecium including a parapleCUthymatous amphithecium, a euthyplactenchymatous exciple, etc., follow the same pattern as the other genera in the Pannariaceae. 125 Dodge (1933) removed Erioderma physioides from the genus Erioderma and made it the type species of a new genus, Malmella, with three new species and another new combination. The way he differentiated between Malmella and Erioderma is that Malmella has less developed tomentum than Erioderma, whereas he has overlooked the fact that .g. limbatum which he has included in Erioderma does not have any tomentum at all (although he says it is tomentose). According to him the phycobiont in Malmella is Nostoc and probably chponema, though his new combination M. physio- .iggg'has Nostoc. The other differences that he gave between Malmella and Epioderma is that the apothecia are scattered in the first, and marginal in the latter. This is not a very dis— tinct difference, because scattered apothecia are seen in ‘g. microcarpa and marginal in p. physioides. Further, he said that amphithecium is absent in Erioderma and present in Malmglla. Maybe this is due to differences in termin- ologies of the tissues of the apothecium, according to the terminologies used here amphithecium is present in all the species of Erioderma that I have examined includ- ing E. physioides (Plate 5). 126 The only difference that is seen between‘g. phygio- i§§§_and other species is the presence of a special kind of cells in the medulla of g, physioides. These cells which have been called moniliform or spheroidal cells, have not been mentioned by Dodge. Asahina (1952) saw them in Pappelia sublaeviggpa and Culberson (1961) noticed them in Parmeliea galbina. The significance of the cells is still unknown; probably they are of no taxonomic value. Relagionshipg of the Genera.based on the Species Studied: A phylogenetic arrangement of the genera of the Pannariaceae was suggested by Reinke (1895), who traced possible relationships of the genera within the family. and to other genera and families (see Section II). zahlbruckner (1926) also arranged the genera of the Pannariaceae into a possible phylogenetic order (see Section II). This study has revealed some characters which I considered important in arranging the genera phylogenti— cally. If Placynthium is to be included in the Pannaria- ceae, then it is probably one of the most highly evolved 127 genera, as shown by its highly developed exciple and What appears to be a retrogressive evolution of the thallus. The latter is indicated by the presence of a cortical layer occupying the whole thickness of the areole. If Plggynthium should be removed from the Pannariaceae as suggested by Dahl (1950) and Choisy (1960), and I believe it should,then Massalongia is the most highly evolved genus. Henssen (1963) treated Massalongia, goerberia. Vesgeggreopgis and Placynthium as closely related genera belonging to the family Peltigeraceae. The reasons she gave for including Massalongia and lac th um in one family were: 1. Both have hemiangiocarpic development of the apothecia, and 2. Their spores are ellipsoid to fusiform and septate. My studies of Massalongia carnosa and Placynthipm piggpm did not show any relationship either between the thalli or the apothecia of these species (see Plates 13, 14, 21, and 22,, and the descriptions on pages 81 and 108). 128 The apothecial anatomy of Massalongia carnosa is quite similar to that of Hydrothyria venosa (Plates 11 and 14 respectively). However, the latter has one-celled spores. The thallus anatomy of Coccpcgrpia Gayana (Plate 1) and that of Hydrothyria venosa (Plate 11) show two very strong similarities: the first is the presence of a lower pseudocortex; and the second is the occasional horizontal orientation of algal filaments in the medulla. Coccocappia gaygpa isia more highly evolved species of Coccogarpia, showing the presence of two-celled spores and slightly stipitate apothecia. The apothecia of Coccocgppia and Erioderma are my- colecanorine (also of Hydpothyrig, Massalongia, Parmeliella, and Placynthium) but it is very hard to say whether the outermost margin of the apothecia of Coccogarpia is an amphithecium or exciple. The apothecia of Erioderma have a well-developed stipe which makes them more highly devel— oped than those of Coccocarpia. The genus Psoroma, though having a bright—green phycobiont as compared to blue-green in other genera, has apothecial characters quite similar to those of Pannaria. Both have lecanorine apothecia but in the amphithecium 129 of Pannppia lurida (Plate 16) a cortical layer is developed. In Psproma hypporum (Plate 23) there is no such differentia- tion. Two characters which suggest that Psoroma is more highly evolved than Panparia are larger spores and the presence of a lower pseudocortex (Plate 23). It seems that Parmeliella is less highly evolved genus than the other genera, because of presence of a pro— thallus, and prosoPlectenchymatous amphithecium. .§-.E12£Q’ phylla (Plate 18) has a lower pseudocortex, no medulla, and a very thick algal layer, making this species more highly evolved than the others in the genus. Probably Parmeliella, Cocgocgrpia, Hydrothyria. and flaggglongig follow the same line of evolution, Epip- {gpgpg being a divergence from somewhere near Coccocappia. The genus Panpapia falls between Papmeliella and the other more developed genera, and is related to Esopoma. At this point I would like to make some suggestions for further studies on the Pannariaceae. 1. Extensive research on the chemistry of the family. 2. More work on the isolation and identification of the phycobionts is needed in the Pannaria- ceae . 130 3. More intensive works on monographs of some genera. IX SUMMARY This study has been done on morphology and anatomy of some of the species of each of the following genera: Cpggocgppia, Eripdepma, Hydrothyria, Massalongia, ngppria. Plggypthium, Parmeliella, and Psopoma, of the family Pan— nariaceae. The following conclusions have been drawn after studying the different species. 1. The thallus in Pannariaceae is squamose to foliose, and is heteromerous or between a true homoiomerous and a true heteromerous type. Only Massalpngia garposa has a true lower cor— tex. Cogcocarpia, Hydroghyria, and Psopoma pallidum.have a lower pseudocortex and the others do not have a lower cortex at all. In Plagypthium nigrum and Parmeliella microphylla, the cortical tissue is present but the algae are not confined to a definite layer. 131 5. 132 The hyphal tissue which appears as a dark or pale line beyond the thallus lobes in PE;- meliella is a prothallus and not a hypothallus. The apothecia are either lecanorine (algae in the amphithecium), or mycolecanorine (no algae in the amphithecium). The outermost layer in the apothecium is an amphithecium (questionable in Coccocarpia). The genus Placynthipm represents many diversi- ties of morphological and anatomical characters from the others, hence should be removed from the Pannariaceae. Eripderma physioides does not present any dif- ferent characters from the other species of Erioderma, except for the presence of monili— form cells in the thallus, hence should not be made the type species of the new genus Malmella (Dodge, 1933), but should be left as a species of Erigderma. The orientation of the fungal hyphae and algal chains was noted but it is not of any taxonomic value, except that in Cpcgocarpia Gayana and 133 Hydppthypia venpsa occasional horizontal fila— ments are seen. In Section XI are plates of longitudinal and cross- sections of both the thalli and apothecia of Eggpgggppga Ggyap . Epiodepma physioidgs, Epiogerma‘Wpighti , figpyp- Lhypip vgnpsa, Massalongia carposa, Pannaria lurida, §2£' meliella plumbea, and Placynthipm nigrum, cross sections of the thalli of Parmeliella picrophylla and Psoroma pg;— ligpm, and longitudinal sections of apothecia of Cocgocap— sis selliss messy: W and Psoroma JB.______h norum- The last plate is of different kinds of tissues that make up the exciple in Collema (redrawn from Degelius, 1954, Fig. 11). LITERATURE CITED Asahina, Y. 1951. Lichenes Japoniae novae vel minus cog- nitae (6), Journ. Jap. Bot. 26:289—293. . 1952. Lichens of Japan. II, p. 98. Genus Parmelia. Research Inst. of Nat. Resources. Tokyo. and Shabita, S. 1954. Chemistry of Lichen Sub- stances. Japan Society of The Promotion of Science. Tokyo. ' Bornet, E. 1909. Rech. Gon. Lich. P. 28, Plate 11. figs. 1-60 Choisy, M. 1960. Classification et Nomenclature Des Parmeliales (suite) Neophysciales:2, Pannariaceae et Placynthiaceae. Bull. Soc. Mycol. France. 76:fasc. 2. ' Corner. J. M. 1929. Studies in the Morphology of Discomy— cetes. I. Trans. Brit. Mycol. Soc. 14:263-291. Culberson, W. L. 1961a. The Pappelia Qpercina group in North America. Amer. Journ. Bot. 48(2):l68~l74. 1961b. Proposed changes in the international code governing the nomenclature of lichens. Taxon lO(6):l6l—l65. E. 1950. Studies in macrolichen flora of South West Greenland. Medd. om Groenland, 150(2). Degelius, G. 1954. The lichen genus Collema in Europe. Symbolae Botanicae Upsalienses, 13(2). 134 135 Dodge, C. W. 1933. The foliose and fruiticose lichens of Costa Rica. I. Ann. Miss. Bot. Gard. 2: 373—467. Dughi. R. 1952. Un problems de lichenologie non résolu: l‘origine et la signification de l'apothécie 1e5— anorine. Ann. Faculté Sci. Marseille II, 21(3): 219-243. g. 1954. L excipulum proprium des apothéoies des Discolichens. Rev. Bryol. Lichénol. II. 23: 300-316. Fink, B. and Richard, C. A. 1915. The Ascomycetes of Ohio. II. The Collemaceae. Ohio State Uni. Bull. 19:28. Columbus. Ohio. Fink. B. 1935. The Lichen Flora of the United States, Uni. of Mich. Press. Ann Arbor. Forsell. K. B. 1885. Beitrage zur Kenntniss dur Anato~ mie und Systematik der Gloeolichenen. Nova Acta Reg. Soc. Sci. Upsal., ser. 3, p. 118. Stolck— holm. Fries, M. 1861. The Lichenes arctoi, Nova Acta Reg. Soc. Sci. Upsal., ser. 3, 72-80. Gaumann. E. A. 1952. The Fungi. (Trans. by F. L.‘Wynd.) 420 pp., 440 figs. Hefner Publishing Co., New York. Gyelnik. V. 1940. Lichenaceae and Pannariaceae. In Rab- enhorst's Kryptogamenflora. 9,(2). Leipzig. Hale, M. E. 1961. Lichen Handbook. Smithsonian Inst. Washington, D. C. Henssen, A. 1963. The North American species of Massalon- gia and generic relationships. Can. Journ. Bot. 41:1331-1346. Hesse, O. 1901. J. Prakt. Chem., 63(2):541. 136 Hue, A. M. 1906. Lichenes morphologice et anatomice dispoSiut. Nouv. Arch. Mus. Hist. Nat. Paris. ser. 4, VIII: 237"2720 . 1912. Lichenes morphologice et anatomice dis- posiut. Nouv. Arch. Mus. Hist. Nat. Paris, ser. 5, II. Imshaug, H. A. 1957. The lichen genus Pyxipe in North and Middle America. Trans. of the Amer. Micro. Soc. 76(3):246—269. Kbrber. G. W. 1855. Syst. Lich. Germ. Die Flechten Dent. Breslau. Lindau. G. 1899. Beitrage zur Kenntniss der Gattung Gy— r0phora. Festcher. Schwendener. Berlin (Hedwigia, 38:101. Beibl.). Massalongo, A. 1855. Syst. Lich. in Sched. Orit. ad Lich. exc. Ital. Mudd, W. 1861. A Manual of British Lichens. Darlington. Nylander, W. 1854-1855. Essai d'une nouvelle Classifica- tion des Lichens. I-II. Memoir. Soc. Sci. Nat. Cherbourg. 3:163-194. . 1861. Notiser un Sallskapets, Fauna Flora Fen- nica. 5:231. , Rasanan, V: 1943. Das System de Flechten. Acta. Bot. Fennica. 33:82. Helsinki. Reinke, J. 1894—1895. Abhandlungen fiber Flechten. I-II. Berlin. Runemark, H. 1956. Studies in Rhizocarpon. I. Taxonomy of the yellow species in Europe. Opera Botanica. 2(1):29. Lund. Santesson, R. 1944. Contribution to the lichen flora of South America. Arkiv. for Botanik. 31 A, no. 7. 137 . 1952. Foliicolous Lichens. I. A Revision of the Taxonomy of the Obligately Foliicolous Lichen— ized Fungi. Symbolae Botan. Upsal. 12(1). Upp— sala. . 1953. The new systematics of lichenized fungi. Proceed. 7th. International Bot. Cong.. pp. 809— 810. Schwendener, S. 1866. Ueber den angeblichen protothallus der krustenflechten. Flora. 49:401-412. . 1867. Ueber den Bau des Flechtenthallus. Verb. Schweiz. Naturf. Ges. Aarau. p. 88. Smith, A. L. 1921. Lichens. Cambridge Uni. Press. Cam- bridge. ' Tuckerman, E. 1872. Genera Lichenum: an arrangement of North American lichens. Lewiston, Maine. Tulasne, L. R. 1852. Mem. pour servir a l'Histoir Organo— graphique et Physiologique des Lichens. Ann. Sci. Nat., ser. 3, 18, p. 5-128 and 153-249. Wainio, E. A. 1921. Lichenes insularum phillippinarum. III. Ann. Acad. Sc. Fenn. Ser. A:15. Helsingfor— siae. ' Zahlbruckner, A. 1907. Lichenes (Flechten). B. Specieller Teil. Engler—Prantl. Die Natflrlichen Pflanzenfamil— ien. 1:1. Leipzig. . 1926. Lichenes (Flechetn). B. Specieller Teil. Engler-Prantl, Die Natfirlichen Pflanzenfamilien. Ed. 2. Bd. 8. Leipzig. Zukal. H. 1895. Morpho. bio. Untersuchungen Flechten. Sitzungsh. K. Akad. Wiss. Math. Natur. W. C1. Wein. civ. pp. 529-574 and 1303-1395. 138 Plate I Coccocarpia gayana.. Cross section of the thallus, illus- trated diagramatically and in detail. UC = upper cortex AL = algal layer M = medulla PC = pseudocortex R = Rhizinae Note: 1. Upper cortex of irregular paraplectenchyma. 2. Algal layer with occasional horizontal filaments. 3. Medulla of parallel hyphae. 4. Pseudocortex developed. 140 Plate II Coccocappia Gayapa. Longitudinal section of the apothe- cium illustrated diagramatically and in detail. HYM = hymenium "EXC"= "exciple" "AMP"= "amphithecium" Note: 1. Mycolecanorine apothecium. 2. No distinction between "amphithecium" and "exciple." 3. Septate paraphyses. 4. Spores two-celled with thick epispore. 141 PLATE I I HYM u « IXC \\\ ’ o In“ ‘ ~“~‘ \\ <2: QCE ‘0 ‘mgx \u \‘u‘ go “ \“ \2 \\ (has: \‘§ ‘ 142 Plate III Coccocarpia pellita. Longitudinal section of the apothe- cium illustrated diagramatically and in detail. APO = apothecium THA thallus EPI = epithecium E hymenium "EXC"= "exciple" "AMP": "amphithecium Note: 1. Adnate apothecium. 2. Well-developed epithecium. 3. No distinction between amphithecium and exciple. 4. Spores very minute. 143 PLATE III T r/ I "I -g/‘(". I / I I g///{ If,” 0 :’;.Q.'.":/“: (Z... .7. II/ ’I’ I . ' / J g. I O. "o.;".‘£ ‘ O ' 0%‘0 / .._°..'.::..::.’"~ 1:. o o M‘? H.453 ” o i O/ :06 'g’, \‘3.’ .}(0 _ Chaim nu \ 0 1 o ( \‘\\ \ ." ‘. ) / ’4...‘D:‘... S \ P- Q? ‘\\\\‘ e: o J-.. . .. c.) 'u-fi" “ / "' ~— 144 Plate IV Erioderma microcarpa. Longitudinal section of the apothe- cium, illustrated diagramatically. EPI = epithecium HYM = hymenium EXC = exciple AMP = amphithecium ST = stipe Note: 1. Mycolecanorine apothecium. 2. Presence of stipe. 3. Well—developed epithecium. 4. Exciple almost of uniform thickness. 145 PLATE IV fil'l ‘HYM IXC AMP 87 3%. . look 146 Plate V Erioderma physioides. Cross section of the thallus illus- trated diagramatically and in detail. UC = upper cortex AL = algal layer M = medulla SPH.C= spheroidal cells T = tomentum Note: 1. Tomentum developed. 2. Medullary hyphae almost parallel to the surface of the thallus. 3. Spheroidal cells in the medulla. 147 i PLATE V ”W Vb "‘ ’ ‘ -"' .95 ‘9 I ”s“ w :- / O l . 9 v 53/ mwkfl \ ’— 35??? 51% vi? 5': §= 148 Plate VI Egiodermayphysioides. Longitudinal section of the apothe— cium, illustrated diagramatically. EPI = epithecium HYM = hymenium EXC = exciple AMP = amphithecium ST = stipe Note: 1. Mycolecanorine apothecium. 2. Well—developed epithecium. 3. Stipe developed. 149 PLATE v: I f U ; I X .- r a “V , ,CS” // s \ / r “J I/ ; /’ Q~Nu : //i , '2“. K/ ' n/“""Q~w .1 _\ J‘\ ‘3 1 1 150 Plate VII Erioderma physioides. Longitudinal section of the apothe- Note: cium illustrated in detail. AMP = amphithecium EXC = exciple EPI = epithecium HYM = hymenium ST = stipe SP = spore Mycolecanorine apothecium. Well—developed epithecium. Exciple narrow at the sides and thicker in the center, of loose, irregular euthyplectenchyma. Stipe developed. Diagram of spore (without scale) to show minutely roughened inner wall. 151 PLATE VI I i»: Z. v.2: 1 UXT si< 152 Plate VIII Erioderma polycarpa. Longitudinal section of the apothe- cium illustrated in detail. AMP amphithecium EPI = epithecium HYM = hymenium EXC = exciple ST = stipe T = tomentum Note: 1. Exciple of irregular, loose euthyplectencyma. 2. Stipe developed. 3. Tomentum well-developed, hairs curved or straight, unicellular, and unbranched. AMP HYM 153 PLATE VIII [KC 8! loo 4. 154 Plate IX Erioderma Wrightii. Cross section of the thallus illus- trated diagramatically and in detail. UC = upper cortex AL = algal layer M = medulla R = rhizinae Note: 1. Upper cortex of irregular paraplectencyma. 2. Algal layer of sparse algal filaments. 3. Medulla of irregular hyphae. 155 ‘ PLATE IX 156 Plate X Erioderma Wriqhtii. Longitudinal section of the apothe- cium illustrated in detail. EPI = epithecium HYM = hymenium EXC = exciple S.HYM = subhymenium AMP = amphithecium T = tomentum SP = spore Note: 1. Mycolecanorine apothecium. 2. Well developed tomentum. 3. Subhymenium seen. 4. Spores 2-celled. 5. Diagram without scale of a 2-ce11ed spore. 157 PLATE X 2 >. _ 2 u I s >- >< . “I I III a .— 50A. 158 Plate XI Hydrothyria venosa. Cross section of the thallus illustrated Note: diagramatically and in detail. UC upper cortex AL = algal layer M = medulla R = rhizinae PC = pseudocortex Upper cortex of paraplectenchymatous hyphae, which are running parallel to the surface of the thallus. Algal filaments horizontal and vertical both. Medullary hyphae parallel to the surface of the thallus. Pseudocortex developed. 159 PLATE XI 160 Plate XII Hydrothyria venosa. Longitudinal section of apothecium Note: illustrated diagramatically and in detail. HYM = hymenium EXC = exciple AMP = amphithecium T = tomentum Apothecium mycolecanorine. The exciple hardly one-celled in the sides, euthyplectenchymatous. Amphithecium paraplectenchymatous. Tomentum well—developed on the sides. 161 PLATE XII HYM AMP \‘ f 1 Us A 162 Plate XIII Massalonqia carnosa. Cross section of thallus illustrated diagramatically and in detail. UC = upper cortex AL = algal layer M = medulla LC = lower cortex Note: 1. Lower cortex present, and is exactly like the upper cortex. 2. Medullary hyphae septate and running irregularly. 163 PLATE XIII 164 Plate XIV Massalonqia carnosa. Longitudinal section of the apothe- cium illustrated in detail. EPI = epithecium HYM = hymenium EXC = exciple AMP = amphithecium T = tomentum Note: 1. Apothecium mycolecanorine. 2. Exciple narrow. 3. Amphithecium paraplectenchymatous. 4. Spores two—celled, rarely three-celled. 166 Plate XV Pannaria lurida. Cross section of thallus illustrated diagramatically and in detail. UC = upper cortex AL = algal layer M = medulla R = rhizinae Note: 1. No lower cortex. 2. Upper cortex paraplectenchymatous hyphae which are parallel to the surface. 3. Medulla of irregular hyphae. Pannaria lurida. trated in AMP EPI = HUM EXC 168 Plate XVI Longitudinal section of apothecium illus- detail. amphithecium epithecium hymenium exciple Note: 1. Apothecium lecanorine. 2. A cortical tissue differentiated in the amphi- thecium. 3. Exciple paraplectenchymatous. 169 PLATE XVI AMP EPI HYM EXC 170 Plate XVII Pannaria pegezoides. Longitudinal section of apothecium illustrated diagramatically. EPI = epithecium HYM = hymenium EXC = exciple AMP = amphithecium Apothecium lecanorine. Apothecium sessile. Exciple wide in the center and almost indistinct in the sides. 171 ‘7 '0: Id! WAN .1. JWV 172 Plate XVIII Parmeliella microphylla. Cross section of thallus illus- trated diagramatically and in detail. UC = upper cortex AL = algal layer . M = medulla PC = pseudocortex Note: 1. Lower cortex absent. 2. Algal layer very wide, wider than the species, filaments vertical. 3. Medulla not distinct. 4. Pseudocortex developed. 173 PLATE XVI I I "C *Al PC 174 PLATE XIX Parmeliella plumbea. Cross section of the thallus illus- trated diagramatically and in detail. UC = upper cortex AL = algal layer M = medulla PT = prothallus Note: 1. Upper cortex paraplectenchymatous, hyphae almost vertical to the surface of the thallus. 2. Medulla of parallel hyphae. 3. Mat of hyphae forming the prothallus. V 175 PLATE XIX 176 Plate XX Parmeliella plumbea. Longintudinal section of apothecium Note: illustrated in detail. HYM = hymenium EXC = exciple AMP = amphithecium No epithecium. Mycolecanorine apothecium.) Spore monostichous and distichous, with thick epispore. Amphithecium prosoplechtenchymatous. 177 PLATE XX g‘c. uXu i»: 178 Plate XXI Placypphium nigrum. Cross section of an areole of the thallus illustrated in detail. F = fungal tissue A = algal filaments Note: 1. The cortical layer is one-celled. 2. Algal filaments are vertical as well as irregular. 3. Medulla absent. 179 PLATE XXI 180 Plate XXII Placynthium nigrum. Longitudinal section of the apothe- Note: cium illustrated diagrammatically. hymenium = exciple amphithecium areole Epithecium absent. Exciple very wide in the center. Spores two-celled‘without.thick epispore. loo A '— IV. fi‘l' 181 PLATE XXII 182 Plate XXIII Psoroma hypnorum. Longitudinal section of the apothecium Note: illustrated in detail. AMP = amphithecium EXC = exciple HYM = hymenium Apothecium lecanorine. The tips of the amphithecium curved inwards. disk concave. Epithecium absent. Exciple of parallel and loose euthyplectenchy- matous tissue. 183 PLATE XXI I I AMP IXC HYM 50." 184 Plate XXIV Psoroma pallidum. Cross section of the thallus illustrated Note: diagramatically and in detail. UC = upper cortex AL = algal layer M = medulla PC = pseudocortex. R = rhizinae Upper cortex of irregular paraplectenchyma. Algal layer narrow. Medulla of arachnoid hyphae. Lower pseudocortex developed. 185 PLATE XXIV 186 Plate XXV Collema. Types of excipular tissue (redrawn from-Degelius, 1954. Fig. 11, p. 83). a. Euthyplectenchymatous. b. Subparaplectenchymatous. c. Euparaplectenchymatous. 187 PLATE XXV lllllllllllllllllllillllllll‘ 77 4437 ll 1 m3 "0