we NORTH AND MIDDLE AMERICAN spams as THE ucuen GENERA ‘gomsvuospouw AND _LOPADIUNI WITH ANTHRAQUINONE PIGMENTS Thesis Iar the Degree of M. S. MICHIGAN STATE UNIVERSITY Judith McKiIIen Burgess 1963 ImmmumummquumulmuW L 3 1293 00108 LIBRARY Michigan State University MSU LIBRARIES .— ‘— RETURNING MATERIALS: PIace in book drop to remove this checkout from your record. FINES wiII be charged if book is returned after the date stamped below. a L ”Teri-1M} u; f '. AEBTRACF THE NORTH AND MIDDLE Al-‘EMCAN SPECIES OF THE LICIIEN GE‘IERA EMBHIOSPORA AIID IDPADIUTI WITH AIII'HRAQUINONE PIGI-IEIITS by Judith McKillen Burgess The lichen genus Ebmbyliospora is composed of two Species groups. The North and Middle American members of the group producing anthra- quinone pigments were the subject of this study. These species have long been supposed to be closely related to the Lopadia with anthra- quinone pigments. I have found that the genus, as presently constituted, is a The unifiying character of its unique spore is more im- natural group. The Spore should be called multi- portant than the pigments produced. locular rather than multicellular; the locules are formed when the endospore thickens and constricts the sporoplast, as in Caloplaca. The endospore sculpturing may be annular, in which case the original pri- mary locule is divided into several, lentiform secondary locules, or it may be spiroform, in which case the primany locule is not separated into parts, but is only forced to take on a spiralled shape. The fact that similarly variable endospore thickening was found in Caloplaca atroalba, during the course of this study, makes it even more certain that Bomgzliospora belongs in the Caloplacaceae. The section Spiro- blastia has been erected to include the anthraquinone-producing'ggm- Qzliosporae, among which spiroform locules are very common. Lopadium vulpinum has been transferred to Bombyliosporg because Judith McKillen Burgess it has the appropriate spore type. The other North and Middle American Iopadia with anthraquinone pigments cannot be transferred, since their spores are of another type. THE NORTH AND MIDDLE AIERICMI SPECIES OF THE LI CHEN GEI'JERA RDMBYLIOSPORA AN D IDPADIUI-I WITH ANTHRAQUII'JONE PIGIVIEI‘ITS by Judith IVIcKillen Burgess A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of I'LASTER OF SCIENCE Department of Botany and Plant Pathology 1963 7.5:: 7. l l . I‘ . '1‘ I ,, ‘ ,-2‘\ '. ’\ ‘. I/(’ f ‘ 1 CONTENTS List of plates..............................................i Introduction................................................1 Morphology Methods..............................................3 Thallus..............................................4 Apothecia............................................5 Spores: ontogeny....................................6 Spore wall sculpture................................IO Pycnidia............................................I7 Chemistry Hethods.............................................18 Results.............................................22 Taxonomy Families............................................25 Genera..............................................26 Key to species...............................29 Bombyliospora................................31 Subgenus Spiroblastia.................31 '5. ggggglg.....................32 o domngfinSi‘gooo0.00000000000033 kn In: 0 flaVidflaOOOOOOOOOO0.0.0.0.038 I WlEzina-OOOOOOOOOOOOOOOOO0.0Cl"O kn Notes on other possible Spiroblastiae..................43 Lopadium.....................................49 The Brigantiaea group.................50 L. cfr. ggdggi.................50 1;. fuscoluteum.................51 ‘L. lgucoxanthum................52 literature CitedOO0.0.00.0...OOOOOOOOOOOOOOOOOOOO0.0.0.0...56 iii LIST OF PLATES Chart Page 1. AnthraQUinoneSooooooooooocoo.oone.000.00.000.0000000021 Plate 1- WWe 2. Bomm§0§mra flavidula..u..........................64 30 BOMliosgora mafia...00.000.009.000.00000000000000066 no WWoooooooooooooooooooooooooooogoggéa 5. Apothecial Types......................................70 6. Spores of Bombzliosmra dondngeng’s..................72 7. Other Spore Types....................................74 8. Spore development in @mbzliospora domggensis.......76 9. Spore variation in Bombzliospora domingensis.........78 10. Plate10...0.0.0...CO...0....OOCOOOOOIOOOOOO0.0.0.00080 11 . Spore variation in W sp, Imshaug 1 5028. . .82 12. Lines of evolution in Bomgliospoga. ... u... ..... ....81+ 1 3. Distribution Map of Bombznomra domingensis. . . . . . . .86 11+. Metribution Map of Other Spinoblastiae. . . . . . . . . . . . . . 88 15° Distflbltion Map Of Lopadium leucoxanthumo o o o o o o o o o O .90 iv INTRODUCTION The lichen genus Bombzliospgra is composed of two Species groups Ihich are superficially dissimilar. The group whose North and Middle American species were the subject of this study is traditionally char. acterized by its yellow or orange apothecia and thallus, and small spam: with lenticular cells. Bombzliospora aureola, _B_. domingensis, and _B_. flavidula belong to this group. The other group, including the type species, 53. pachycarpg Mass.. is traditionally recognized by means of its lack of anthraquinone pigments and its large spores with cubical cells. I The genus lopadium has apparently analogous groups also occurring in the perscribed area. and its species with anthraquinone pigments have often been supposed to be closely related to the Bomgglio§porae with anthraquinones or congeneric with them. The Wooncerned are L. cfr. Magi, _L_. fuscoluteum, L. leucoxanthum, and L. vulpinum. The purpose of this investigation was to re-evaluate the position of the Bommliogpgrae with anthrequinones within the genus, to charac- terize their unique spam, and to determine their natural relationship with the anthraquinone-producing Lopadi . I have found that L. vulgnum must be transferred to Bommlio- m, so to avoid confusion, I will refer to it throughout the paper as Bomgxliospora Min . Specimens studied were obtained from these herbaria: University of Michigan (MICK), Missouri Botanical Garden (M0), Michigan State University (MSG). and the Smithsonian Institution (US). The photographs were taken by Mr. Philip Coleman. I am grateful for the guidance and encouragement of my advisor, Dr» Henry A. ImShaug. and I wish to thank Dr. Flordeliz R. Uyenco for her fresh collections of Bomgyliospora from Florida. Thanks are due to Mr. Dennis Jackson for the camera lucida tracings of fig. 10 and 11 of plate 9. MORPHOIDGY flethods. Freehand sections of the thallus and apothecia of every speci- men were made with a razor blade and crushed by gentle pressure on the cover slip. Apothecia of representative specimens of each species were sectioned at 25 p with a freezing microtome using Wetmore's (1960) method and were pemanently mounted in Diaphane (Will 0011)., Roches- ter, New York). Some were stained with 0.1 per cent acid fuchsin in 95 per cent ethanol, and others were mounted unstained in order to demonstrate the natural color of the exciple. Tissues were diagrammed with the aid of a Zeiss drawing apparatus, then stylized diagrams were made with these diagrams for guidance. Spores were drawn with the aid of a Zeiss drawing apparatus, and were measured, in distilled water. In order to photograph the spores, freehand apothecial sections were mounted unstained in Diaphane and squashed immediately, or were mounted and squashed in Turtox Mount- ing Medium to which a few crystals of acid fuchsin had been added, or were mounted, unstained, in water. The second medium reacted with the lichen tissues and bubbles were formed after a few days. Change in spore appearance with addition of 20 per cent aqueous ROI-I was noted. The location of cytoplasm in spores wasvverified by mounting in Amann's Medium to which acid fuchsin had been added. 3 Thallg The most conspicuous characteristic of the thallus is its color. Due to the presence of anthraquinones, herbarium specimens are usually orange or gold colored, and K+ rose (K+ means there is a color change with addition of XOR: K- indicates lack of color change). Some are at least partly glaucous, and a few are ochraceous. Bomgzlio- 5221;; aureola has a yellow thallus. Bombzliospora flavidula often has reddish maculi. Tuckerman must have been referring to these when he wrote (in Nylander, 1863a) "thallo interdum hinc inde intus miniato— tincto (erythrinoso)." Acharius (1811»), Muller (1887a), Tuckerman (1888), Shirley (1889), Bailey (1891), and Fink (1935) have all pointed out the great variation of thallus color occurring even within one specimen. Very little fresh material was seen, but without exception it was pale green. The fresh material seen consisted of p. domingensis and g. vulpina, species which have glaucous thalli much less often than does _B_. flavidula. This material, collected by Uyenco in Florida in February, 1963, had begun to turn yellowish by the end of April, 1963. It was air dried. It is suggested that the change in color is due partly to a slight loss of color by the algae, but mostly to the con- version of the anthraquinone in the thallus from a colorless sugar solution to a colored crystalline state before death of the thallus (see Thomson, 1955). In some herbarium specimens, birefringent gold or red particles can be seen in the medulla and algal layer. The thallus is thin (up to 150 p), heteromerous, and has no H311 defined cortex. The smooth upper surface is composed of indis- tinct periclinal hyphae. I have seen no rhizinae on the lower surface; in fact, the thallus is sometimes hypophloedal with some hyphae and a __._.__. _ q. , - 77' few algal cells among the bark cells. The thallus may be areolate, in which case the wide, thin, white hypothallus is prominent, or it may be rugose or smooth, becoming areo- late. This variation seems to lack significance. The algal cells are 7-16/u in diameter and have lobed chloro- plasts. They are thought to be Trebouxia. Large red-sheathed colonies of Gloeocapsa were seen rather fre- quently inside the thallus and apothecia of several specimens of,§. domiggensis and.§. flavidula. Apparently it is a common epiphyte which can he accidentally included in the lichen tissue. ggmggliogpora aureola has been described as having a rimose thal- lus, but the single example I saw was rimose merely because the bark on which it grew was cracked. Leprose and coralloid varieties of g. dondggensis and p. vulpina have been described, but not from this area, and I have not seen any. Apothecia The apothecia always contain an abundance of anthraquinone pig- ments. 'Relative colors of the disks and margins are of some help in determining Species but are not reliable. A few specimens of g, ngipg BET-5’33 and g. v’ulpina have yellow pruina which can be scraped off to reveal dark disks. Bomgyliospgra aureola disks are heavily yellow pruinose, but disks of p. flavidula are never pruinose. Disks which are not pruinose may appear fuzzy due to the anthraquinone crust on the tips of the paraphyses; this condition Should not be confused with pruinosity. The entire, persistent margin may be yellow, orange, or ochra. ceous. It is thick, except in _B_. flavidula, whose mature apothecia have thinner margins than those of the other species (plate 2). If the disk is pruinose, the pruina extends onto the inner edge of the margin, which is usually elevated. There is no signigicant variation in maximum size of apothecia. Young apothecia are flat and mature ones may remain plane or become slightly (rarely greatly) convex. The outline of mature apothecia is very often extremely gyrose (undulate, flexuous), and columns of sterile tissue the color of the margin rise through the hymenium to appear as raised pads of that color on the disk (plate 1). The apothecia often appear crowded, and many small ones sometimes arise on what seems to be the disk of a larger one. Fee (1824 and 1837) pictured some of the apothecial configurations. Iviicroscopically, there is a good deal of uniformity in apothecial structure. In general, the exciple is more or less intensely rust colored, although it may be colorless, and a distinct, colorless, nar- row subhymenium is usually visible. Malme's collection of _13. domineen- gig in Lriphenes Austroamericani, no. 31 g and p is illustrative; g has a colorless exciple and _s an intensely colored one. A stylized diagram of the tissues is presented in plate 5. The terms are used as defined by Imshaug (1957). The apothecial tissues of Ionadium cfr. dodrei and L. leucoxanthmn are also of this type, but those of L. fuscolutewn are different (fig. 2). Mass m The name Bombyliospora means bulging spore. De Notaris (181%)) chose the name, and he used it only in connection I—ith g. pachycama. However, the literature and my own observations indicate that the spore wall of this species is smooth, not bulged (see fig. of _B_. incana,= 13.. pachycgpa, in Zahlbruckner, 1905 and 1926). Perhaps the name applies to spores in an intermediate stage of development, where the endospore annnli are just beginning to constrict the sporoplast (plate 10, fig. 4). KBrber (1855) and Frank (1886) described the spores of‘g. pachycagpa as caterpillarbshaped. Bomgyliospgra has always been described as having multiseptate spores, in contrast to the muriform spores of Lopadium. These two terms are no longer adequate in View of present knowledge of spore development and morphology; The appearance of mature spores cannot be accurately described without corfleration of how they got that appearance. This is particularly true with muriform spores. Hue (1912) pointed out that polarilocular and multilocular spores of Iicheni blasteniospori Mass. have no true septa and are therefore unicellular: "Les spores des lichens blasténiosporés sont simples, elles naissent simples at demeurent tounours telles, mais en meme temps elles sont polocoelees, c'est-a-dire que, par suite de l'épaississement equatorial de leur tegument, elles présentent aux deux phles des cavities hemiSphérique...." By the same line of reasoning, the spores of BoMbyliospora also are unicellular. At first, the spore consists of a single, continuous, cylindrical sporoplast surrounded by a uniformly thin wall of epispore and endospore. Very early in development, the endospore begins to thicken, causing the sporoplast to lose its cylindrical shape and be- come a tube having protrusions in a characteristic pattern. The pro- trusions represent locules of the mature spore. I use locule here as Hue used it, to mean a cavity containing a sporoplast within the spore wall, as Opposed to cell, which is a unit delimited by a wall laid down on a middle lamella. The original cylindrical locule of Mg- gggrg will be called the primary locule. In the mature spore it may remain primary, but take on a spiroform shape,.or it may be separated by endospore into several lentiform secondary locules. The number of secondary locules depends on the species. In blasteniosporous lichens, according to Hue, additional concen- tric layers of wall material are deposited inside the original epispore and endospore, constricting the cylindrical (primary) locule into two polar areas or into several (secondary);i.hocules. The polar areas are usually still connected by a tube at maturity, but sometimes not. Hue based his concept of the concentric layering on 'petites fentes appar- aissant ca et 15 sur la surface des spores" and considered the idea con- firmed when he observed a spore cut transversely. He saw three concen- tric circles: the outer, of many layers, he interpreted as the outermost spore wall; the middle he called the endospore, which was "de formation plus recente, [and therefore] moins apparentes;" the innermost circle he called the tube. As no species is Imam to have such a thick epi- spore, I believe Hue did not see the epispore at all. He probably saw a section showing the tube, a locule above or below it (his endo- spore), and the endospore. The "petites fentes" were probably folds in the walls of nonturgid spores. There is no indication of concentric layering of the spore walls in Bombyliospora, but in some way the endospore thickens and constricts the sporoplast, forcing it to assume the shape of the remaining space. The sequence of diagramsi’n plate8 represents the changes in appearance of one spore type of B. dondngensis as it develops; the process is continuous, therefore the diagrams do not represent distinct stages. Photographs of various stages of both spore types of g. domingensis appearinplate 6, andof§.m. inplate 7. Wall thickenings grating from opposite sides of the spore even- tually meet and obliterate the central tube, fanning secondary locules. Thus, although the final appearance is of a septate spore. it is no more septste than is a W spore: it is mnceuular, and multi- locular. Spores of all North and fiddle American species of W lack the central tube at maturity. The secondary locules nearly touch each other, and endospore wall material completely fills the very small area between then. The epispore is thin, and endospore alone is responsible for the formation of secondary locules or distortion of the primary locule into a spiral. Primary and secondary locules are each surrounded by a membrane. . Actually, they seem to be enclosed by a wall as thick as the epispore. There is em reasonforthinldngthisnaytrulybeawallandnot merely a membrane surrounding a plasnolyzed sporoplast: I expect study of the species uthout mthraqflmnes to clarify this point. The spore wall is indented opposite the secondary locules or opposite the spirofore locule, and bulged between. but this is not always apparent. The pattern is so synetrical that secondary locules look as though they are formed by overlapping circles (plate 6, fig. 3). This is accentuated by XOR. men the spore wall is mtured in KOH. locules often say be aqueesed out individually (plate 7, fig. 5. ani plate 10, fig. 3), each wrapped in its own membrane. Spore Me, number. and size varies with the species. 10 9.312811% The manner of thickening of the endospore is striking and nest simficanb. The cum thin endospore has the potential to develop in either of two we. Annular thickening results in the for- ution of more or less lens-shaped secondary locules. Thickening in the for: of a spiral causes the primary locule to take on a spiralled shapebutdoesnot separateitintosecondarylocules. Bothtypes occur in nearly equal numbers in g. M. The spdroforn type is much less canon in p. m, mile 3. m never has the entire endospore spiralled. W m never has an por- tion of the endospore spiralled, but there is indication of that tendency. Itissignflicantthatinatleast onespeciesofma, too, the endospore may be thickened in either way. I have seen one specimen “WM (mm, from South Dakota, rec), in which sons spores have the usual single annular thickening, but may have a Spiralled thickeMng instead. The spores have been described (Fink, 1935) as obscure]: one-septate. Shape and orientation of lentiforn locules in W varies between species and the shape also somevhat witldn species. In a. M, with an ovate spore,* locule shape depends on location in the Spore. mloculesarenearlyroundemeptthe oneatthetipofthe narrow end of the spore: that one is usually elongated (plate 10, fig. 1). The locules inthe smallerhalf ofthe spore are smallerthanthosein the other half. 'Tems referring to spore shape (except short fusiforn) are taken from mm 1: Descriptive Terminology, ‘w the systematics Assoc" 1962; the terms describe optical sections . 11 Secondaryloculesofg.Wand§.Waretms- versely lentifon; that is. each locule looks like a thick, double- convexlens those principalplaneis perpendicularto thelong ads or the spore. Those of g. M almost always form an acute angle at the edge or the spore. but these or a. m never do (plate 10, fig. 2). B. m has secondary locules oriented longitudinally: they look like portions of a thick double-convex lens those principal plane is parallel to the long axis of the spore. Secondary locales or _B_. m are always subdivided transversely (to the locule) into ter- tiary locales ty extensions of the locule membrane. and are sometimes further divided longitudinally by such extensions. In _B_. W. one transverse septum rarely divides one or more secondary locules; this condition, when frequent. is the basis for Malme's (1923) South Anrioan variety W and Asahina's (193M variety W from the Benin Islands. Tertiary locules can be easily seen and counted then locules are squeezed out or the spore wall (plate 10. fig. 3). The number is not constant, but in 3. W only one ring of tertiary locules forms, an in g. m, two concentric rings are present at maturity. The spores of m are superficially sindJar to those of E. m, but are actually very different. In L. W, material in the aseus shows transverse and longitudinal lines or separation eden before a spore wall is laid down. These lines. vhich because the septations or the spore, are entirely irregular at an age. All walls are uniformly thin; these is no thickening of the endospore and the spore is completely filled with nnute cells (plate 7, fig, 7):, In old. necroue spores vbose sporoplast has disintegrated. a three 6.1.33. sional reticulum of irregular septa can be seen throughout the spore. 12 The spores often break down within asci into oval conidia about 7 1' 2m. Apparently healthy spores at all stages of development are often totally brain. and 1+ arose. Spores of W never break down into caddie. and althoughthelocules of old sporesarerarelybromor red. and K-. the spore wall is never colored. Spores of L. W and L... cd'r. m are similar to each other in that regular transverse septa fem before further division proceeds. but at maturity. L. W spores look like those of L. W; as the spore grows in size. the cells become progressively further divided. and the divisions become irregular. The spores are often brown. . In lg. ctr. m, all septa originate regular and remain regu. lar. forming right angles (plate 7. fig 6). The mature cells are larger than those of the other two w. and the spores are never brown. although they are often 18- rose. Further. when pressure is applied to spores of the former two species, they split irregularly through the center. lon- ntuddnally. and do not come apart. In L. ctr. 93432;, however. the spores break up in a manner strongly remeniscent of g. m. They sometimes come apart along the seven or nine original transverse septa (the area Wen is divided into three rows of cells at maturity). or they may be crushed more completely to yield a flat sheet of locules finish remain connected or become separated. In additiOn. occasionally a wide, colorless. K- ring surrounds the spores I have not detemined the nature of this ring. The say these spores come apart then crushed implies to me that pmmp. ere cells lackalintingneu. brane, vhile L. cfr. 99.93%, like W. possesses a halting manomnneaohumt orthe spore. ' 13 Since Lopadium spore patterns are not due to endospore thicken- ing, the terms primary and secondary locule cannot be applied. No spiralled pattern has been seen in Lopadium. Spiroform endospore thickening is less common than annular thickening in Bombyliospora, although it is very frequent and easily recognized inig. domigganSts. From drawings and notes made by Tuck- erman and others on old specimen cards, it is certain that they inter- preted spores with spiroform endospores as submuriform. Malme (1923) and Asahina (1934), however, used subnmrifom to indicate septate secondary locules. A spiroform locule must be called a primary locule because it is never separated into units by endospore thickening. Spiral thick- ening of the endospore merely forces the sporoplast to assume a spiral shape. As the single, continuous ridge of endospore develops, it pushes further into the interior of the spore. Until it reaches the midpoint, the sporoplast looks like an auger; it has a cylindrical core with a horizontally flattened ridge running its length spirally. When the endospore reaches and passes the midpoint, the central core is obliter- ated, leaving only the spiralled part, which is no longer flat but is more or less cylindrical. Invaginations of the locule membrane often form, making the locule transversely septate, but it is still a primary locule. By transverse, I mean the septa are perpendicular to the sides of the locule, no matter what their angle in relation to the spore wall. The spiroform locule of‘g. vulpina is always very much septate, often appearing beady. The spiral ridge of endospore is narrower than in g. domiggensis, therefore more of the volume of the mature spore is occupied by sporoplast. These facts make the spiroform locule of'g. vulpina more difficult to recognize. I have found that slightly oblique 114 ”secondary locales“ are a reliable indication of spirals (plate 7, fig. 3). With lnck, one can confirm this by squashing the spore to release the locale. Sprinter. locales are usually broken at a septum in the pro- cess, but retain their form except for unidnding slightly. Lentiforn locules are unconnected disks . ‘ Mamas. mew have usuanxeight- locular spores (sometimes six.- or ten-locular), and accordingly. their spiroforn locules have eight turns (gyros). It is not at all unusual to see more than one spirofom ridge of endosporeon one spore wall, especially in 9,. W (plate 6, fig. 7, 8, 9). In the majonty of cases, the two ridges arise on opposite sides of the spore and turn in the same direction: clockwise. I saw one spore with spirals going in opposite directions; the endospore was not thicker at the intersection point. Marv spores have one complete ridge the length of the spore plus portions of one or more others. In other words, ridges of spiralled endospore do not always extend from one end of the spore to the other, but may arise and end at any point, 301111118 another ridge if they end in the middleoi‘ the spore .‘ Practically infinite variations and conunations or the two basic spore types-mular and spiroform endospore- have been seen, most of then in p. m, probably because they are easier to recognize in that species. All of them occur with or without septa dividing the locules. Usually either completely annular or completely spiroform endospore predominates in a given specimen. W m does not have completely spirofom eulo- SPOI‘O. but often that would berths two anmli at either eni of the 8901‘ Join to form a short spiral. In l. m, the end. nest annulus often does not reach the spore mdpoint, therefore the 15 two end locules appear joined (plate 10, fig. 1). I had thought the number of spiroforn locules per ascus might represent the number of nuclei present, but this now seems unlikely; the locale number is a result of the number of ridges of endospore, and that number is inconstant. I The spiroform locule is truly spiralled, not helical. Using Thompson's (1961) definitions, a spiral is "a curve which, starting from a point of origin, continually dimmshes in curvature as it recedes from that point; or in other words, whose we 2; mture continually increases.” A helix has the same radius of curvature throughout. Since the spore diameter decreases towazd an end, the wall thickening must be spiral. Further, the spiroform locule is an equable spiral, or spiral of Archimedes; its shape is that of a coiled cylinder, not a coiled cone. It should be pointed out that similarly variable wall thickenings are not unknown in plant cells. The secondary wall of tracheary ele- ments of may woody plants is deposited as rings or helices, with the type depending on age of the plant. (Age is not influential in _B_gn- me) However, Esau (1961) states "...the different types of wall structure intergrade. The annular thickenings may be intercon- nected here and there; annular and helical...thickenings may be com- l'lmed in the same cell...‘' This seems remarkably similar to the situ- ation in Bomliospora. Prelimnary study of 102 specimens of eight species of m. m which lack anthraquinone pigments has convinced me that their spores are in every way similar to those of the species with anthra- quinones. The sporoplast remains cylindrical. until the spore is rather large, then three to eleven (rarely more) rings or endospore begin to 16 pinch the sporoplast into secondary locules. One never gets the im- pression that they are connected by a central tube, though. As I have attempted to show in plate 10, fig. 4, the reason is purely due to the angles of curvature. Each locale is surrounded by a limiting structure ditch we become very wide (plate 11. fig. 1+). and seems to be a wall rather than a membrane. Secondary locules are longitudinally lentifom. Host investigators have called them cubical, but Zahlbruckner (1905 and 1926) and Fink (1935) called them lenticular. In a single specimen (W 159;, frm Jamaica, 1186), several spores hd completely or partially sporoform endospore, lending strong evidence that the genus is a natural group. Most of the rest of its locales were distorted like those pictured by Zahlbruckner (1905 and 1926) for _B_. m. The spore variations in this specimen are shown in plate 11. Many other specimens have locales distorted like these. An evolutionary sequence of spore types incorporating some species lacking anthraquinones is presentedin plate 12. Not all types of spores known in W are represented because I have not seen them: the types involved are those of .B_. wemaceae, E. W, _B_. W, _B_. egansa, and p. themengi . Toms used to describe these spores were chosen so they could be used in other genera also. Thus, one may say of most Caloplacae: primary locule reduced to two polar areas connected by a central tube, 1. e., polarilocular; of most M: primary locule mischoblastio- norphic, with or without a median transverse septum, i. e., mischolo- cular; of some W: primary locule reduced to two spheres con- nected by a narrow tube, with or without a median transverse septum, and if connecting tube is lost, one spherical secondary locule perk sists in each cell. i. e., pachylocular. 17 We The only species seen with pycnidia was _B_. domggensis, and even then only a very few specimens had pycnidia. The pycnidia are minute, flask shaped, and have carbonaceous walls. The contents are heavily conglutimted, meldng it very difficult to determine the attachment of the conidia. The conidia are oval, 7 x Z/u. Almost none of the literature reports mention pyonidia or coni- (119.. For his _B_. dowensis var. flavocrocea, from Africa, Nylander (1869) describes the pyonidia as convex lutescent, with conidia 5 x 2.5/11, aaclfixa mnutissima oblongo-cylimlrica" on jointed sterigmata 2.5m wide. Zahlbruclcner (1905) put Bombyhomra in the Lecideaceae, vhich he stated are seldom endobasidial, then in 1926, he transferred it to the Caloplacaceae. possibly on account of the corridial attachmsnt. He names only g. dMensis among K+ rose species, and does not describe the pyonidia or conidia. CHEMISTRY moss A 20 per cent aqueous solution of XOR was applied to the upper surface and exposed medulla of the thallus by means of drawn capillary tubes, and the color change was noted. This KOH solution was also added to apothecial sections in distilled water on slides and the color change of each tissue noted . Specific color reactions may be helpful in identifying the anthraquinone, according to Asahina (1954), but were not here. In an attempt to identify the arrthraquinone( s), extractions were made in hot acetone in small test tubes, then a few drops of 0.5 per cent methyl alcoholic magnesium acetate (MgAc) were added. For each specimen, one extraction was made from a few pieces of thallus and another from a few apothecia. According to Asahina, colors gotten in this way may be diagnostic. I found that too much depended on concen- tration of the substance, and the colors of several standards (species named by Asahina as sources from which a specific anthraquinone has been purified) seemed identical. I then tried paper chromatography of acetone extracts, using standard techniques. The method of Shibata, Takiko and Tanaka (1950), using petroleum ether saturated 16th 97 Per cent metmrl alcohol as solvent and 0.5 per cent methyl alcoholic MgAc to bring out the color of the spots, was not effective. The substances did not dissolve well and there was much streaking. 18 19 Three solvents mentioned by Hale (1961) were tried, ascending in tubes, also with little success. They were n-butanol saturated with marlin, n-butanol-acstone-water, and n-butanol-alcwol-wat or; for ratios and phases, see Hale, 1961 . The best solvent tried was ethyl acetate-pyridine-water (2:1 :1). It dissolved the substances completely and left distinct spots with little streaking. It was tried with ascending technique, in tubes, and descending, on sheets in a tank. The two methods gave comparable results, so I used the descending, since in that way I could run more samples at a time with the equipment available. What-m No. 1 chromatography paper was used, and the solvent was allowed to run about 11+ inches. One set of specimens was run 19 inches in the hope of separating a double spot, but without success. After drying, the sheets were sustained with long wave (3660 A) UV light and the position and color of fluorescent spots were marked. Some sheets were then sprayed with methyl alcoholic MgAc, dry-ed at 90°C. checked for color reaction, and re-exann.ned for color of fluorescnece. Because the colors could not be differentiated, other chromatograms were not Sprayed. Drawn capillary tubes containing 20 per cent aqueous KOH, 100 per cent ethyl alcoholic pherwlenediamine (PD), or MgAc were used to test each spot. The standards for each anthraquinone named by Asmna were treated in the same way. In an effort to determine the similarity of anthraqmnones in W, W, and other Caloplacaceae. the pigments of several species not otheMse studied were chromatogramed. Results are compiled in table 1. Because the epithecia and margins are not the same color, and sometimes turn slightly different colors in KOH, epithecia and mrgins 20 Chart 1 . Anthraquinones (1) Includes these species: (2) Includes these species: Bombyliospora domingensis Caloplaca cinnamomea B. flavidula C. tetraspora B. aureola C. arizonica B. vulpina Blastenia leucoraea Lopadium fuscolut eum L. leucoxanthum L. leucoxanthoides L. cfr. dodgei Caloplaca ulmorum C. stillicicdiorum C. feracissima C. flavovirescens C. aurantiaca C. carphinea C. pyracea C. cerina C. elegans C. lobulata 21 Chart 1. Anthraquinones standard Mange a? Xanthoria parietina parietin .98 Getraria endocrocea endocrocin .70 Solorina crocea solorinic acid .98 Xanthoria fallax fallacin .97 Teloschistes flavicans teloschistin .97 Parmelia formosana lichenanthone .80 unknown I .95 unknown II .95 unknown III .95 unknown IV’ 5 .76 unknown V .90 unknown VI .60 color before treatment yellow uv-orange bright yellow- orange - uv-deep orange yellow uv-orange yellow uv-erange yellow uv-orange none ‘11?- bright yellow uv-bright yellow-orange yellow uv-orange yellow-orange uv-yellow-orange yellow-orange uv-pal‘e ‘ orange pale yellow uv-pale orange pale yellow uv-pale orange color after kAc pink uv-pink-orange rose uv-deep rose pink uv-pink-orange pink uv-pink-orange pink uv-pink-orange ' blue uv-s pale orange-pink uv-pale pink pale rose uv-rose deep magenta uv-deep rose pale rose uv-rose pale rose uv-rose pale rose uv-pale rose color after KOH pink deep violet pink pink pink yellow uv-white pink uv-pink pale violet uv-violet deep violet uv-deep purple pale violet uv-violet pale violet uv-violet species Protoblastenia rupestrio (1 ) (2) (2) 8. cinnamomea 22 were chromatogrmed separately for one specimen of each species of W am W studied. Apothecia were dissected under magnification to insure that only the appropriate tissue was taken, then the pieces were placed directly on the chromatogram sheet and acetone was applied. Good spots were obtained, showing that enough material was present, but there was no difference in spots from margin and epithecium. All specimens were inspected macroscopically under UV for fluorescence of thallus surface, medulla, and apothecia. may: then KOH is applied to the thallus surface, orange, gold, or tan thalli tum some shade of red, indicating presence of anthraquinone. The thallus of p. agola turns purplish. Glaucous and fresh (green) thalli are K-. The medulla of g. was; is K+ yellow. In enmining thalli with endothallic W Moroscopically, one must take care not to confuse the color of the algal sheath with K reactions. The red sheath does not change color in KOH. When KOH is applied to apothecial sections, the epithecium turns rose or magenta, or sometimes pale violet. The cortex of the amphi- thecium tums intensely rose, and immeasurably small purple particles diffuse out. After standing a few mmtes, these particles group in M11 aggregates, but never form large crystals or gain any definite shape, even after standing for several days. Rudolph (1955) has stated that parietin particles form large crystals after standing one half hour, so perhaps .the Bomgliomsa anthraqminone is not parietin. The rust colored emciple is 1(- except in p. aureola, where it is not actually rust colored, but more nearly the color of the amphi- thecial cortex. This emiple becomes slightly reddish in K021. In 23 addition, the thallus of this species is more purple than the margin in KGB. The pyridine chromatograms eliminated endocrocin as a possible new for the W and Iopadimn anthraquinone. Still, parie- tin. fallacin, and teloschistin could not be differentiated, and the W and Lemur: spots have the same R7: and colors with every reagent as these three. However, it is not necessary to know the name of the substance; it is enough to know that all K+ rose Bommliogmrae and w produce the same pigment, and that the same pigment is the most col-son one among the Caloplacaceae . It is interesting that W may be separated from M by its anthraquinone in addition to spore type. Rudolph (1955) stated that parietin is the only anthraquinone in the Caloplacaceae, but he relied on literature reports which were rather old, published before the days of chromtograplv. I have found 811 anthraquionoes among Caloplacaceae with orange or cinnamon apothe- cia. The cinnamon group is most interesting, having at least two (sometimes four) unique pigments. It seems likely that further study of these will prove them to be a taxon worthy of autonomy. Spots at Rf .95 often appear to be double because of the pres- ence of atranorine at the same Rf: only one anthraquinone is there. Atranorine was proven to be present in the following species by re. czystanizing it in G.A.o'1'. (formula in Asahina, 1951+): W W, p. w, W cfr. 992%! L. fuscoluteum, Tel - m worms, Xanthoria W. and g. gag. The medulla of g. .v_ul_pi_.n_§ contains a substance, not an anthra- qmnone, that is diagnostic for the species. The medulla fluoresces mite, and the substance forms a flocculent white precipitate with 24 HgAc in a test tube. Its Rf the spot is pale yellow: it is more yellow with MgAc. and fluoresces in pyridine in .95. Before treatment, yellow-vhite. KOH does not change the color in ordinary light, but changes the fluorescence to purple. PD has no effect. The substance does not crystallize in 6.3. or G.A.oT. (both formulas in Asahina, 195a). Apothecia of L. W yield a gray, non-fluorescing spot at R4? .07. It is MgAc-, Ko, and PD-. Apothecia of L. leucoxanthoides, whose tissues are K+ blue, yield a streak beginning about Rf .93. It is gray before MgAc and faintly blue after. It absorbs UV, is PD-, and is made colorless by KOH. It is not known whether this substance is responsible for the blue color. TAXONOMI muss Massalongo (1852) described Bompllioggra in the Lecideaceae. Zahlbruckner considered it to belong to that family in 1905, but in 1926 he transferred it to the Caloplacaceae on account of its endo- basidial conidia and thick spore nails. All members of the Lacideaceae except Megalogpgra, according to Zahlbruckner, have oxobasidial conidia. In addition, members of the Lecideaceae seldom have Spores with thick- ened walls. Lopgggum has always been placed in the lecideaceae even though its spore walls are often much thickened. The conidia, according to Zahlbruckner (1905 and 1926), are exobasidial. I have not seen direct descriptions of pycnidia or conidia of the species included in this study, but Santesson (1952) stated that the foliicolous species, at least, have exobasidial conidia. Asahina (1931+) evidently suspects that the corticolous species may be endobasidial. he proposes that, if they are, Lopflum and Megalosmra should be transferred to the Caloplacaceae. Then, according to him, the three genera would repre- sent one evolutionary series within the family. Spore septation would therefore be given less importance than it is now. Des Abbayes (1951) stated that the position of Bomgzliomg in relation to the Caloplacaceae was uncertain because of its "pluri- cellulaire" spores. ‘ Santesson (1952) pointed out that Brigantiaea Travis. (1853, not seen) was used before Lew KBrber (1855). Trevisan intended 25 26 W to include all m with anthramlinone pigments. £9,13- 2).-19.22% m was not known until 1863, but Santesson includes it in what he calls the Brigantiaea group. He comments also that 3.3.. d931,- m2 is probably a member of the group. Rudolph (1955) stated that Bomgznoggerg cannot be included in the Caloplacaceae because of its spore type. I have shown this to be untrue. In 1v opinion, Bomgzgospgra is closely related to Caloplaca, and belongs in the Caloplacaceae, as shown by the spore development and by the anthraquinones produced. I believe Iopgdium should be left in the Lecideaceae, since it has exobasidial conidia, and its spores do not have thickened endospores. gems Ban e rs. Massalongo (1852) credits De Notaris with first recogrn.tion of the genus W, but De Notaris never published its description. De Notaris wrote only that g. m, ILDNtrs. hex-bu" differed from Biatora icmadopgla in the structure of the apo- thecia and the size and septation of its spores. Massalongo designated 1;. w the type species, and described 13. versicolor at the same time. He thought then that Lecidea (=Bomflomra) versicolar F59 and _L_e_cidea (=Bogmgospgra) tgberculosa Fée were synonymous, but in 1860 he recognized g. tgbercglosa Fee as a separate species. Bomgzuosmra W is pictured by Massalongo as having two Spore types in the same apotheciuln: some large spores, with thin walled cubical locales, and some smaller spores, with thick walled transversely lentiform locules. Obviously Massalongo was mistaken in picturing both types from 27 one apothecium. I think the source of the error is discernable. He states that he is not sure his concept of the genus agrees with that of De Notaris, and in the legend for the drawing of _B_. pachzgaga, he states "Quests figure vennero cavate da un exemplars favoritonti dal prof. De-Notaris...." Thus it is probablya composite drawing; he observed one spore type and erroneously thought De Notaris' illustra- tion of the other was from the same species. Maesalongo (1852) claims that the spores of _B. versicolor are 91 .5—122.0 x 214.11-30.5/u and one per ascus, but he pictures them with transversely lentiform locules. Massalongo was the only person ever to report seeing such large spores with that type of locale. He wrote that he took his drawing of _B_. versicolor from Fe'e's favorite example of lecanom versicolog, but F66 (1837) made it clear that his 9. 1.23;. 9313}; had two-celled spores, and this is now the basiormn for Megalogora W. Massalongo thought his :3. versicolor was the same as Fee's L. tuberculosa, but Fé'e drew cubical (not longitudinally lentifonn) loculed spores for that species. Zahlbruckner (1931) recognized that Maesalcngo's _B_. veflcolor was not the same as Fee's Lecidea versicolor and Zahlbruckner therefore lists a Bombfliogmra versicolor Mass. in his Catalogus. However, Massalongo actually made a new combination of Fee's basionym, so Massalongo's Bommylogpgra is without a name. There is reason to believe, however, that no such lichen endsts and that Masealongo's description and illustration are the result of compound errors. A nomenclatural note concerning g. m is in order here. Zahlbmckner (1931) thought g. W should be called _B_. m because its basionym was was incanus Smith. Actually Smith (in Smith and Sowerby, 1807) stated that his species was based on meg 28 1.5m Ash” inch is the besionvl of m m. Therefore m cannot be used for aw W. The correct basiom is W W Del. in baby (1830). The only K-I- rose tam now in 1the germs that were not placed there by Zahlbruckner in his Catalogue (1931) are _s. M, which was added as g. m (Spreng.) Mass., and its variety Qyocroce . iiich was added as g. flavocggcea (”1.) Mass., both in 1860. Several other species now in the genus were known before that date, including the K+ rose W described in 1811, mtgg var. W, 1833, and W, 1829. There is no explanation as to why Massa- longo did not include them. His inclusion of s. W (as m) and its variety flavocrocea was the only indication until lahlbruckner's Iona-ental work that am K+ rose species belonged to M. I think, Massalongo would not have thought them members of this genus ex— cept for his confused concept of the locule shape. I see no reason why anyone would have thought the K+ rose and K- species groups related before the spore ontogeny was studied. Although one might think the difference in pigments produced would indicate that two genera are involved here, I believe the strongly unifying characteristics of spore development and a tendency to spiro- form endospore thickening is more important than pigments. Recognition of two subgenera is, however, warranted. I have chosen to call these tan subgenera rather than sections because the differences between then are too basic to give them only sectional rank. My. Until now, g. m was thought to be one of a natural group of K+ rose Lopedia in North and Middle America. I have found that the other members of the group cannot belong to W 29 since they lack the unique spore type of W. Also, the thallus in w is thicker and granular or sorediose. Santeseon (1952) stated that medium fonns no good taxonomic unit. Three foliicolous gerera have already been segregated from it and Santesson saw at least four groups of mainly corticolous species resuming. The type species of Loggia, L. W, is different from all. the other species except .I_._. W and probably 1,. gaggl- m in having paraphyses with dark, swollen calyptra. Until such time that further study proves that additional gerera should be segregated from M, Santesson proposed Low as a m conseMum against W. Brigantiag is the older name, and must be reserved for the K+ rose species. BoMliospgra (1852), however, is older than both, so it it should be decided that the K+ rose m must be united with the K+ rose Bomgzliomra, the latter name will take pre- cedence over gigantiaea. In view of the absence of a monographic treatment of the poly. norphic "genus” W, and in the absence of any-suitable section name, the species of w discussed here are merely referred to as the Brigantiaea group. KEY TO SPECIES This key includes all the North and Middle American species of Bommliosmra and Lopadium with anthraquinone pigments. 1. Spores muriform, 45-129 1: 1940a, one per ascus; thal- lus white or ashy, granular or sorediose.....................2 1. Spores mdcellular, multiloculate (may give false impres- sion of being muriform; see text), 12-‘47 x 6-21/u, two 3. 5. 5. 30 or more per ascus: thallus glaucous, orange, or ochra- ceous, smooth............................................".4 2. Apothecia stalked, with algae in margin; exciple colorless, with highly refractive shite crys- tals: musoicolous.............................._L_. fuscolutgg 2. Apothecia not stalked, no algae in margin: ex- ciple rust colored, without crystals; corticolous....3 Apothecia up to 0.7 m across: disks dark ochraceous: margins black; spores (ll-$59475 x (18)21-2641, septa regular at maturity...................................L. cfr. gage; Apothecia up to 1.6 mm across; disks heavily yellow pminose or nodular ochraceous; margins orange; spares Sit-115029) x 19J+O/u, wepta irregular at maturity....L. leucoxanthum it. Spores ellipticus, more than 25m long...............6 lb. Spores snguste-ovatus or short fusiform, less than 25/u long............................................5 Spares anguste-ovatus, 18-20(2l+) x 5m, secordary lacules nearly round: disks yellow pminose..........§. aureola Spores short fusiform, 12-18 x 6.7/11, secondary locules transversely lentiform; disks light red to fulvous..............................................g. flavidga 6. Spores (MM-8: nae, secondary locules trans- versely lentifom, rarely septste, spiroform locules prevalent; medulla UV-................§. d ensis 6. Spores 2 : nae, secondary locules longitudinal- ly lentifom, always much septate; medulla nv+ mites m 31 EMOSPOEA Mass. Ricerch. Anton. Lich. 1111. 1852. Psomthegg sect. Bombyliospora (Hum) Stizenb. Bericht. Thfitigk. St. Gallischen nat. Gas. 160. 1862. W sect. Bombyliospora Mass.) Tuck. Nsy. Lich. 2:55. 1888. Egg-cg subgen. Bombyliospora (143.88.) Wainio, Acta Soc. Faun. Fl. Fenn. 7:30. 1890. W sect. Bombyliospora Mass.) Mtll. Arg. Bull. Herb. Boissier 2(appendix1):65. 1891+. TYPE SPECIES: 5. W an.) De Not. m sect. Platylecania Mall. Arg. Flora 65:327. 1882. TYPE SPECIES: Earns (=Bomflogmra) fllificans (Nyl.) Mfill. Arg. W Stein, Schles. Ges. vaterl. Cultur 60:231. 1883. TYPE SPECIES: We (=Bon1pzliomra) m Stein. W. Thallus crustose, heteromerous, epiphloedal, scar- ticate: primary phycobiont trebousdoid. Ascocarp an apothecium of the molecanorine type, adnate. Paraphyses simple, nonseptate, not capi- tate, conglutinated. Spores one to eight per ascus, colorless (locules occasionally red or mscescent in age), short fusiform, anguste-ovatus, or ellipticus: primary locule cylindrical in earliest stages, becoming scalarifom or spirofonn: secondary locules it» to 12 (rarely 1h), trans- versely or longitudinally lentiform, persistently nonseptate or becom- ing transversely septate, sometimes with a few longitudinal septa: locules occasionally partly lentiform and partly spiroform in the same spore. Conidia endobasidial, according to Zahlbruckner (1905 and 1926). Subgenus SPIROBIASTIA Burgess, subgen. nov. awosi . Thallus smooth, rugose, or areolate, usually con- tairdng an anthraquinone . Hypothallus wide, thin, mite . Apothecium up to 1.8 m across, often in aggregates, often gyrose, with sterile 32 columns: margins yellow, orange, or ochraceous: epithecia and mnphi- theoial cortices K+ rose or violet, rarely blue. Spores 2 to 8 per ascus, 12495 x 7-25/u. TYPE SPECIES: _B_. W (Pers.) Zahlbr. somosrom mason (Nyl.) Zahlbr. Cat. Lich. Univ. 7:119. 1931. m m1; Nyl. Ann. Sci. Nat. Bot. IV. 19:353. 1866. Heterothecium aureolum (Ny1.) Tuck. Syn. Lich. 2:159. 1888. Lecflra aureola (Nyl.) Nyl. in Hue, Nouv. Arch. Mus. Iii. 3:53. 1891. ms: collected by Wright, 3. 3., in Cuba: hb. Tuckerman. W. Thallus yellow, smooth: medulla UV-. Apothecium up to 1.2 mm across, flat, somewhat imbedded in thallus; disk yellow pruinose; mamn nearly concolorous with disk, becoming crenulate. Emiple orange-rust colored, K+ orange-red: bark carried up into apo- thecium. Spores 6-7-8:nae, anguste-ovatus, 18-20(211) x 5/11, with 5-6 nearly round secondary locules, end two locales often Joined but not twisted: endospore never spirofom. gscusgon. Stirton (1898) intended this species to be in his germs Miltidea but he failed to make the combination. This species is rare. It has been reported from Cuba (Tuck- erman in Nylander, 1863, and Tucker-man, 1866) and from southern Africa (Stirton, 1898). I doubt that Stirton's material was actually _B_. m: the spores were 5-9 locular. There has been no mention of substrate in the literature. Mateggl seen. WET INIIL'E mam-o RICO: Aibonito, in a wood, Mm, 3 Jan. 1916 (HIGH). Mapped in fig. 111». 33 EMBYIIOSPORA mMIINGENSIS (Pers.) Zahlbr. in Waura at Back, Itin. Princip. Coburg. 2:153. 1888. Patellarig WPers. Ann. Wetterauisch. Gas. 2:12. 1811. Leeanora W (Pers.) Ach. Syn. Irish. 336. 18111. Pamelia 3223.928. var. W Eschw. in Martius, Fl. Brasil. 1:189. 1833. W W (Pers.) Nyl. Acta Soc. Sci. Fenn. 7:862. 1863. rm m sect. Lopadium dflggense (Pers.) Tuck. Proc. Am. Acad. 7:229. 1868. utideg domiggefls Stirt. Trans. Free. New Zealand Inst. 30: 386. 1898. Placodium doMense (Pers.) Wain. Dansk. Bot. Ark. #(11): 11. 1926. m dowense (Pers.) Fink, Lich. Fl. United States 235. 1935. ms: collected entrees in Santo Domingo (Hispaniola) by Persoon. Legidea m Spreng. K. Svenska Vet. Akad. Handl. #7. 1820. m mes (Spreng.) Mont. in Sagra, Hist. 1'Ile Cuba, Bot. 212. 1842. Heterothecium menu: (Spreng.) Flat. Bot. Zeit. 8: 551+. Esa- EEO—5213 m (Spreng.) Mass. Atti I. R. Istit. Veneto III. 5: 262. 1860. TYPE: on bark of 232295113 mosmillomora in Guadalupe: collector not stated. Lecanog acemata Raddi, Mem. Soc. Ital. Sci. Modena 20:51, 1:. 3. f. 2. 1827. Type not stated. m W 1‘. W Wain. Ann. Acad. Sci. Fenn. A. 6(7):121+. 1915. Bomflliogra «Mange var. glaucotmpa Wain.) Wain. Ann. Acad. Sci. Form. A. 15(6):103. 1921. TYPE: Mm, on bark of W ingisus in Basso-Terra, Guadalupe. WWfo WW. Ark. Bot- A. 18(12):5. 1923. TYPE: yam 1.6.2.: Rio de Janeimv and M .‘téi s and g, Pot-to Alegro. 31+ M We var. m Steiner. Bull- Herb- Bois. II. 7:616. 1907. TYPE: Sanat. sub. no. 995. Cape Province, Africa, gm, 1900. W. Thallus glaucous, yellow-orange, golden, or rarely 0611ch then dried, smooth, rugose, or areolate: medulla UV-. Apo- thecimn up to 1.8 mm across, flat, slightly convex, or rarely strongly convent: disk dark red to fuscous, rarely sparsely yellow pminose. kciple usually rust colored, sometimes colorless, always K-: bark not carried into apothecium: algae sometimes present in stipe. Spores usually 6-81nae, sometimes 2J+:nae and than larger, 28-145 1: 11-14(15- 17) In: endospore thickening annular or spirofrom or a combination of the two, locales nonseptate, or occasionally with a few transverse septa especially in spiroform locules: lentifom locules forming acute mgles at edge of spore (rarely not acute). W. This is the most common of the species studied. It has been reported on Cappazis enemallophoza (the type, Sprengel, 1820), Xanthoglum caribaeum (foo, 1824), and Artocms Ecisus (Wain- io, 1915), and from lable data, on Oxamdra, Hyperbena, lig_ui_damb_a£, m, Mates, Poinciana 5% Age; m, and M. If one were to separate those specimens with mostly spiroform endospore from those with mostly amular endospore, an interesting distribution becomes evident. The former group would be found in large numbers in southeastern United States, but not at all in the West Indies or Middle America. The latter group is found mainly in the West Indies and Middle America, with only a few (5 out of 31) specimens from southeastern United States. I have examined material from Brazil, Paraguay, Hawaii and the Benin Islands and find that it is the same as that from North and 35 Middle America. One specialen (M 565 E, from Rio Grande do Sul, Porto Alegro, Brazil, 27 Sept. 1892, in M0), which was listed in Malme's type description of his f. tmggressa, looks like normal _8. W with locales occasionally septate. The single specimen I have seen from the Benin Islands (m, g. g" 1853-56, in US) has many spores vdth spirofom endospores, so perhaps it is not Asahina's (1939) var. bordnensis. From the descriptions, I can see no difference between W and Mamie other than locality. Steiner's var. i_13_sp_grsa is based on the yellow pruinose disk, with a darker color showing when the pruina is scraped off. I have seen this variation and consider it insignificant since it varies greatly within a single specimen. Wainio's var. glaucotropa is probably not a variety since a single thallus may be partly glaucous and partly orange. I have seen a specimen (Dopgland 8&8, from the Philippines, April 190"», in US) which was designated and published as f. M- W by Wainio, but which seems to me to fit the description of E. W var. intermedia. The spores are like those of 13. W except for their size-20-30 x, 9-13/u, according to Maller (1891+). Those of Copeland's specimen were consistently six locular, and the size was 19-21 x 9-12/11. No material like it was seen from the area covered in this study. No specimens fitting descriptions of other published varieties (of this species were seen. Bommliogora domingensis has been reported in the literature from Europe (southern France, Zahlbruckner, 1905 and 1926), Africa (not-thorn Africa, Zahlbruckner, 1905 and 1926; Port Natal, Nylander 1896b, Hue, 1891, Stizenberger 1892: Usambara, Mdller 1891+; Ivory Coast, 36 dos Abbayes 1955: Ben Spei, Nylander 1859 and 1869b, Hue 1891), south- eastern United States (South Carolina and Louisiana, Tuckerman 1872: Florida and Texas, Tuckerman 1888: South Carolina, Louisiana Florida, Texas and Alabama, Rink 1935). Central America (Mexico, Hedrick 1935), West Indies (Guadalupe, Hue 1891: Santa Domingo, Persoon 1811; Cuba, Homtagne is Segre 181m enhance, Riddle1920), South America (Brazil, Eschweiler in Martius 1833, Miller, 1886, Maine 1923: Argentina, Krem- pelhuber 1878: Colombia, Nylander 1863b: Argentina and Colombia, Nylan- der 1869b, Hue 1891: Argentina and Paraguay, Muller 1891:), Hawaiian Islands (Oahu, Tuckerman 1868), Asia (Benin Islands and Taiwan, Asahina 1931+), Philippine Islands (Comiran and Samal, Wainio 1921), Australia (Stimberger 1895: New Caledonia, Nylander 1869b: Queensland, Shirley 1889: Brisbane , Miller 1891: Toowoomba and Queensland, Miller 1882: Queensland , Bdley 1891), am New Zealand (Hooker 1867, kbington 1855 [pot sooty; north island, Hellbom 1896, Muller 1891+). A nomenclatural note is in order here. Zahlbruckner (1931) was in error in listing Nylander (1869a) as the author of B. lgprolytg. He and Hue (1891) called it a subspecies of _B_. domingensi . Muller was first to use the name at the specific level, in 1882. Zahlbruckner mitted the homonym Patellaria dgggensis var. leproMa Shirley (1889). The species is mentioned by Bailey (1891) but he misspelled it legg- m. dutch may account for Zahlbruckner's classion of that reference. M §_e_e_r_1,. Exsiccati. Malme, Lich. Austroam. 1+1 A 8: B. 11mm: arms , A Fromm: county unlmom, m _1_9_, 1889 (use 119979 and 119981.); Honorsit, Callcins 35; (mm); 6 Mile Base, W (no); 6 Mile Creek, m 161013; mmedtdth p. m): Caloosa River, m, 1878 (as ex hb. Willey): southwest m" m a mg. 1878 (vs). 37 001mm 00mm. Lake city, 1835322151., July 11-19. 1895 (MSC, us). BABE (DUNTI, Homestead, Owaissa Bauer Hammock, m 12.2, 9 Feb. 1963 (160). 310mm mUNTY, Highland Hamocks State Park, Sebring, m 22, 11 Feb. 1963 (use; mixed with _s. _v_ul_2i_r_1_a and p. loucoxanthum). POLK 00mm. Scott Lake near Lakeland, Mgh'haxmnock, Mozarlin 39. 4/6/31 (Inca). summon: 00m. Black Bottom Hammock, 3 mi E of Oveido, mac 2:, 9 Feb. 1963 (use; mixed with a. 221.22%). TAIIDR 001mm. Aucilla River on Route 98, W 52_2_8_, 11 Sept. 1951+ (MSG). IDUISIANA: parish unknown, M (HIGH): Coteau, hamlets _r, 2 Aug. 1892 (US); chhamps, Mg, 17 Sept. 1894 (US); John Durands', w, 2 Jan. 1891» (us): Pninte aux Loups, Lansing. 30 Sept. 1895 (vs); St. Leo, M, 21 Mar. 1895 (HIGH 6: hb. Fink 1986). PLAQUMNBB PARISH, Points a la Hache, M393, 20 Nov. 1885 (US). SAINT LANDR!’ PARISH, Grand Coteau, M )3, 6 June 1894 (US), Opelousas, Mg, 22 June 1885 (as). SOUTH CAROLINA: county unknown, m (MICE! ex hb. Fink 1511-11, 0 mixed with g. m. MO 341%; as; and as ex hb. *mley), Santee Canal, m, 1849 (mm, Reliquiae Tuckemarde 104). TEAS: county unknown, gel; _6_, 1872 (us ex hb. Willey). 05mm AMMCA MTISH HONDURAS: locality unlmown, M__a_.i_.n_;_s_ 3122, 3 July 1936 (mm): E1 Cayo District, along Belze River, Egan's; 3535, 17 June 1936 (Midi): Valenten, M 35;, 23 June 1936 (HIGH). moo: locality unknown, ”Tamasopo, m, 10 June 1890 (use £15823): San Luis Potosi, Las Palmas, Lrigglg L63, 10 June 1890 (vs). Yucatan Peninsula, Chichen Itze, in San Yui Cenote, its-em 3.7.9.1.: 29 June 1932 (men); Lake Cbicbanoanab, Quintana R00, 8335: 2361, 28-19 July 1932 (mm); San Miguel, Cozumel Island. Quintana Roo, m 38 3957. 6-8 Aug. 1932 (HIGH). WET INDIE CUBA: Province unknown, m (US); M g; 192 (US); map all (mm 82: hb. Bennett): m £31015). MATANZAS PROVINCE, Canar- ioca, m a \«al___go_n 3312;, 7 Feb. 1916 (NO). OKIENTE PROVINCE, Gran Piedra, south slope about halfway up, wooded recky ravine with small waterfalls, Sierra Maestra, W 35916, 25951, 25312, 16 Aug. 1959 (166). HAITI: DEPAMI‘ DU NORD, hill west of Cap Haitian, mm 366;, 10 July 1958 (rec). JAMAICA: HANOVER PARISH, Dolphin Head, 1750 ft, m 1.5.6.3.. 23 Apr. 1953. ST. ANN PARISH, Albion, 2400 ft, pm 1593;, 7 May 1953. ST. CATHERINE PARISH, Montpelier Sumnfit, 2300 it, M M, 28 Jan. 1953: Hollymount, Mt. Diablo, 2754 ft, gem 1129.8: 28 Jan. 1953. all in BBC. PUERI'O RICO: locality unknown, Earle, summer 1903 (MICE! ex hb. Fink 1111-198). Aibonito, in a wood, M 1182 (HIGH ex hb. Fink 8336, mixed with g. rlayidula) and Link 17;. 1/3/1916 (Inch ex hb. Fink 8102): Candelaria, near Bayamon, m. Egg 8: m 2883. 11 April 1913 (HIGH): near Martin Pena Station, 30 ft, giggle; 3331, 19 Jan. 1900 (US. MD): and He or g. 3., 19 Jan. 1900 (men exhb. M14909); Pele de la Piedra, Elmira 8: m 159, 16 Feb. 1923 (MIKE): Rio Piedras, in a wood, _Fi_n_k§1_6_§, 18 Jan. 1916 (MICE, US). The localities are mapped in fig. 13. HDMBTIIOSPORA FLAvIUULA (Tuck.) Zahlbr. Cat. Lich. Univ. 7:53. 1931. 133132 MTuck. in Nyl. Ann. Sci. Nat. Bot. 11?. 19:253. 1863. w W * flavidgg 39 (Tuck.) Nyl. in Hue, Nouv. Arch. Mus. 1:11. 3:53. 1891. TYPE: col- lected in Cuba by Wright, 93.: in hb. Tuckerman. m. Thallus grey-green or glaucous and K-, or occasionally tan and K-l- rose, smooth, entire: medulla UU-g. Apothecium to up 1.7 mm across, sometimes gyrose, often with sterile colunms: disk flat, light red to fulvous, never pruinose: margin rather narrow at maturity; false thalline margin often present, K-. Spores uni- or biseriate, 6-8znae, short fusiform, 12-18 x 6-8 (rarely 5) u: secondary locules 11.-6, trans- versely lentiform, not forming an acute angle at edge of spore, never septate: end two locales sometimes joined and twisted sprially in 6- locular spores. mgcussion. This is a rare species and was previously known «fly from the original collection. The Puerto Rican specimens were con- sistently h—locular except for the one in US, which had a few spores with six locales, the end two Joined. The Haitian material was consistently 6-1ocular except for Wetmore 11215;, which had a very few 5- (nct 311:) locular spores. Since the other characteristics are identical, I be- lieve that this variation may be interpreted as incipient speciation, but not yet worthy of taxonomic recognition. Populations isolated by water have come to differ by one annular thickening per spore. I have not seen authentic Colombian material of g. doMgensis var. ineggcata (Nyl.) Zahlbr., but the description by Nylanier (1863a and 1863b) fits _B_. flavidga. Mate _s_e_9_:_1_. WEST INDIES HAITI: DEPARTMENT DE L'OUEST, ravine on east slope above road NW of For'et des Pins, Imshagg m, 16 July 1958: elfin woodland for- est, northof Forert dos Pins (Shade Station), near Dominican border, 1&0 5800 ft. mm. mast!» July 1958: elfin woodland, ridge above (east of) Foret dos Pins, 5800-5900 ft, W 3_1_§1_, 3123, 1125, 13- July 1958. all in MSG. PUERTO RICO: Aibonito, in a wood, _F.‘_i_nk_ 1182 (US: men ex hb. Fink 8062: and MECH ex hb. Fink 8336, mixed with _B_. W): Ha.- cicada Pulgillas, Coamo, M £21. 25 Aug. 1920 (men). The localities are mapped in fig. 11:. EDWIOSPORA VULPINA (Tuck.) Birgess, comb. nov. m 3.11M Tuck. in Nyl. Ann. Sci. Nat. Bot. IV. 19:354. 1863. Heterothegm sect. Lopadium m (Tuck.) Tuck. Syn. Lich. 2:57. 1888. mtidea m (Tuck.) Stirt. Trans. Proc. New Zealand Inst. 30:386. 1898. W BEE-EM (Tuck.) Wain. Ann. Acad. Sci. Fenn. A. 15(6):88. 1921. TYPE: collected in Cuba by Tuckennan, g. 3.: hb. Tuckeman. Loflium vulpigum var. personatum (Tuck.) Malme, Ark. Bot. A. 29(6):25. 1937. TYPE: M 2&1... at the edge are woods, Santa Anna da Chapada, Matto Grosso, Brazil, 28 Feb. 1891+. Iocidea subvulpina Nyl. Bull. Soc. Linn. Normandie II. 2:89. 1868. m m var. $11me (Nyl.) Zahlbr. Cat. Lich. Univ. 4:317. 1927. TYPE: collected by Tuckeman, g. g. Diagnosi . Thanus golden , occasionally glaucous: smooth, ru- gose, or areolate: medulla UV+ white. Apothecium flat to slightly convex: disk fulvous, occasionally sparsely yellow pruinose. Spores Zmae (rarely1 or innae), unisefiate, ellipticus, (ZN-27)28-leo(l+2-lr7) I (9)12-19(21) u: spiroform or secondary locules much septate trans- versely, sometimes longitudinally septate: secondary locules longi- tudinally lentiform. 41 Discussion. This is the second most frequently collected spe- cies, but is found far less often than is _s. dgggensi . Although the type locality is Cuba, Imshaug did not find it in extensive collecting there. It has been reported in the literature from Florida (Tuckeman 1888), the West Indies (Cuba, Tuckerman in Nylander 1863b, Tuckerman 1888, Miller 1881a, Hue 1891, Iynge 1939: Hispaniola, Wainio 1915), Cen- tral America (Hoodoo, Davis 1936), Asia (Ceylon and East Irdies, MfiJler 1881a), Africa (Mauritius and Guyana, Tuckerman in Nylander 1863a: Ivory Coast, des Abbayes 1955), and Australia (Shirley 1889, Stirton 1898: New Caledonia, Nylander 1868 and 1869: New Caledonia and Noumea, Miller 1893a: Brisbane and Queensland, miller 1891). The only substrates identified on the lables were Magnolia, gag- mug, 91-3292: Quercus vigiflana, and palm. The only specimen which had some spores singly in asci was m 51: only the specimen Lash, Fla., had some spores innae. Mdllcr (1881a) stated that Heterothecium Hfleanm Mass., 11. gristg Hampe and E. fuscenscens Hampe are synonyms of _E. village. These are all BM 1.3292: not mentioned by Zahlbruckner (1931 ). Malme (1937) mistakenly thought a K+ violet blue reaction of the epithecimn and amphithecium was normal for _8. vulgina, and on that baSis he described var. Emonatum for K+ rose specimens. Tuckeman did not mention the K reaction in the original description, but I have seen Malme's exsiccatum Lichenes Austroamericani no. 195 (MSG) mich he called m, and the K reaction of this specimen is distinctly different from that of the isotype. Therefore, Malme's var. permtum is iden- tical to var. vulgina and must be discarded. Specimens with a K+ violet blue epithecium and amphithecium represent Lecidea subvulm Nyl. , designated as "epithecium ert con. ceptaculum hydrate kalico coerulee tinguntur' by Nylanier (1868). l+2 There is some question, however, as to whether or not the name sum as validly published. Nylander!s (1868) wording, "Disjuncta scilicet when subvulpina Nyl., quae disnoscitur tanquam varietas [of mm)" leaves one wondering if he meant to call it a species or a variety. This would fall under Article 1} of the Rules (1956); the name was not validly published since the author failed to recognize the taxon at arm certain level. This taxon is for the present treated as a synonym of _B_. E12293 since the significance of the color reaction is not hilly understood. It seems to be analogous to the situation with Lopadium leucoxanthum (K+ rose) and L. leucoxarrthoides (K+ blue). Many authors have comented on the similarity between g. do en- gig and _B_. Wilma and their varieties in the past (Tuckeman, 1866, 1868, 1872, 1888: Tuckerman in Nylander, 1863b; Nylander, 1869a: Miller, 1882 and 1891; Shirley, 1889: Stirton, 1898; Malme, 1923; Asahina, 1934: Santesson, 1952), but if they were made members of the same genus, it was g. domimensis that was transferred, because of its "submxifom" spores. It is interesting that the two species have similar fame and varieties reported in the literature, namely glaucescent, ochraceous. and coralloid. None of these were described from North or Middle Amer- ican material but I have seen both glaucescent and ochraceous forms. Material m. UNITED STATES FLORIDA: county unknown, Austin 8: _S_n_fi._t_t_1_, Mar. 1878 (US ex hb. Smith): Caloosa River, m, 1878 (us ex hb. Willey); Calldns 8, 1889 (MSG); m 8, no date (MSG); Mg 1Q, 1889 (M80): 6 Mile Creek, M 151 (US, mixed with g. dogggensis). ALACHUA COUNTY, Gains- Ville, Ravenel, 1877 (US ex hb. Willey). COLUMBIA COUNTY, lake City, @2111. July 11-19, 1895 (no, US). HIGHIANDS COUNTY, Highland 43 Hammocks State Park, Behring, m _82, 11 Feb. 1963 (NSC); Lime ber- ry trail, Highland Hammocks State Park, Sebring, ERIE—9. 81, 11 Feb. 1963 (MSG); Wm. lieder Trail, Highland Hammocks State Park, Sebring, m 22, 11 Feb. 1963 (mac, mixed with _a. domiggensis): LAKE comm EusthS, M 1252, May 28-June 15, 1895 (MO, US). POLK COUNTY, Peace Creek, M, Mar. 1880 (US ex hb. Smith). SMNOLE WUNTY, Black Bottom Hammock, 3 mi E of Oveido, m 51 and fl, 9 Feb. 1963 (MSC, it mixed with g. domingensis); Oviedo, Heed (no ex hb. Moore). SOUTH CAROLINA: county unknown, Ravenel, on trees (MICH ex hb. Fink 15411). CENTRAL MEGA BRITISH HONDURAS: El Cayo District, Cohune Ridge, EELS. 3182i, 10 July 1936 (HIGH). MEDECO: Yucatan peninsula, in forest, Champoton, Campeche, £3213 L8_2_6_, 7-15 July 1932 (PITCH); Yocatan peninsula, San Miguel, Cozumel Island, Quintana Roo, in forest, Steers. 3221, 6-8 Aug.t1932 (HIGH). WEST INDIEB CUBA: locality unknown, M; (US ex hb. Willey ex Miss Wilson; no spores seen by me); M II 110a (US); 1113113 233 (US, ISOTYPE). Localities are mapped in fig. 11». Notes 9_r_1_ other possible W. In my study of the liter- ature, I found numerous references to certain species of Bommliospora and Lopadium commenting on their similarity to one or more of the spe. else just diagnosed. Because they have not been found in North or Mid- dle America, they were not included in this study, and no material was seen. However, since spores with spiral patterns have been mentioned M in the literature only in connection with the species below, I feel this discussion is pertinent. LOPAUIUM EEFERUM (Nyl.) Zahlbr. Cat. Lich. Univ. M300. 1927. Lecidea m Nyl. Ann. Sci. Nat. Bot. IV. 15:47. 1861. Heterothecium m (Nyl.) Mill. Arg. Flora 6+: 101%. 1881. TYPE: collected by Pancher in Mauritius and Guyana. Iodidea leucoxantha var. bigora Nyl. Mem. Soc. Sci. Nat. Cher- bourg 5:123. 1856. (nomen nudum) Disggsion. Nylander's original desceiption (1861) stated that the spores are "transversim pluries divisis vel fere murali-loculosis ..." and two per ascus. He later (1863a) described them as "quasi intus oblique vel spiraliter murali-divisis (seriebus circiter 10)" in comparing the species to B. vulpina. In 1868, he wrote "Sporis binis intus spiraliter (nonnihil oblique) murali divisis mox distinguenda Species." His description of the thallus varies from "Similis externe [_Iepadiunz] leucoxanthae" (1861) to cinerascent (1863a) to 'glaucescens tennis vel glauco-virescens vel macula glauco-flavida indicatur" (1868). The epithecium is K+ purpurascnns. The species was originally described from Mauritius and Guyana, as was )2. vulgna, and was later reported from New Caledonia and Lifu (Nylander, 1868), Brisbane (Miller, 1891), and Noumea (Miller, 1893a). Stirton (1898)stated that _L_. m and L. graham probably belong to the group (muriform-spored initidea) of which _B_. vulpina is the type. LOPADIUM EIFERUM var. PLUEEFERUM (Nyl.) Zahlbr. Cat. Lich. Univ. 11:300. 1927. legidea more var. plurifeg Nyl. Bull. Soc. Linn. Normandie II. 2:89. 1868. TYPE: collected by Nylander in New Caledonia and Lifu. 45 Discussion. This lichen was described by Nylander (1868) as a variety which has spores 2-6 per ascus. and epithecium K-I- "fuscescente '(obscuriorem sumente colorem, raro purpurascente)." LOPADIUM PARAELUM (Nyl.) Zahlbr. Cat. Lich. Univ. M309. 1927. Lecidea parabola Nyl. Bull. Soc, Linn. Noraandie II. 2:90. 1868. Heterothecium parabolum (Nyl.) Mdll. Arg. Rev. Mycol. 9:80. 1887. TYPE: collected by Nylander in New Caledonia and Life. Discussion. This species was described as similar to _L_. £21313 but with thallus dilute glauco-flavid, spores "vulgo 8 spiraliter con- timzis, parum divisis,“ 30-34 x 13-152u, and epithecium K+ purpuras- cens (Nylander, 1868). Miller recorded its occurrence in Australia (1887a) and Noumea (1887b). maximum PARAHJLUM var. SUBWLPINUM (Mini. Arg.) Zahlbr. Cat. Lich. Univ. 4:309. 1927. Heterothecium parabolum var. _s_u_b;vu_l_..- 91-23% Mall. Arg. Flora 70:338. 1887. 14993925 W var. §_u_1_3_1n_i_]_.- 2% (Mill. Arg.) Shirley, Proc. Roy. Soc. Queensland 6:184. 1889. TYPE: collected by Sayer at Trinity Bay, Australia. Discusgon. The thallus of this vanety is “aurantiaco flavus wt in Ii. Mina Tuck.';' (Mfiller, 1887a), but the spores are like those of normal L. parabolum. or L. biferum except "intus simplicius spiral. iter divisae." Shirley's (1889) _I_.._. parabolarhad a golden yellow thal- lus but he commented on the color variability with single specimens. Maller (1887a) stated that thallus color is "in tota eerie Specierum t affinium, _H_. leucoxarrthi, H. m, a. M at H. Lug-m evidenter inconstans ert saepe in eodem specimine e flavescenti plus minusve 1:6 glaucescent: v. albicans." Zahlbruckner (1927) erroneously listed this as var. ghalflum: actually, Killer and Shirley called it var. subvulpigg. EDHBYIIOSPORA COCCODES (Be1.-) Mass. Atti I. a. Instit. Veneto III. 5:262. 1860. M coccodes 861. Voy. hides-Orient. 2:125. t. 13. r. 2. 1846. magi-a coccodes (361.) soot. Ann. Sci. Nat. Bot. III. 10:126. 181:8. Lecidea coccodes (1361.) Nyl. B111. Soc. Linn. Nomandie II. 1:89. 1868. TYPE: collected by Delanger on Reunion. W. This species is more difficult to interpret because of the lack of clear descriptions of its spores. The thallus surface is different from that of the species Just discussed: according to filamger (18%), the surface is 'subtartarefi, granulatoé-iconglomerat’a, albe-psllescente,‘ but the granules become "stalactiformes, et verda‘tres dans an age plus avanc‘é.‘ He did not describe the spores. Montagne (1848) discussed the species briefly in comparison with three other species, but placed emphasis on the characteristics of the thalli, writing only that the spores of _B_. coccodes have '1: i 6 nuclEnsJ‘ The Species it was com pared vdth are one-septate, multilocular, and muri- form, respectively. Nylander (1868) evidently felt B. cgccodes might be confused with I. m var. plurifem, since he pointed out that the yellow material in the margin of the latter forms a purple liquid with KOH, tile in g. goocodes, the yellow pigment of the margin is "statim violacee purpurascit.‘ In his (1868) description of g... m. Nylander noted that its spores are almost the same as those of g. m. If one assumed 117 that the specific name was chosen with regard to the spore shape, then incorporated hints from the literature, one would expect the spores to be cocciforn and to have a spirofonn endospore. This invites curiosity as to the sinlarity between 2. coccodes and L. 111m 1’. corallinum (um. Lrg.) Zahlbr., which has a yellowish thallus. Tuckerman (1866 and 1888) described the spores of g. m as cocciforn to elongate- ellipsoid. I have seen one specimen from the Philippines (McGregor, 19111, in 08), those spores must be called cocciform except for occasion- a1 ones thich are alone in the ascus. and then ellipsoid; its yellow- orange thallus in snooth rather than coralloid. It seems possible that g. coccodes could have either white and granular, or yellow and smooth thalli; the thallus of L. leucoxanthum has greatly variable surface tex- ture, from w observations, and its color ranges from white to sulfur yellow if one accepts Lecidea bicolor Spreng. as a synonym of L. M- man- Trevisan'a 1853 report on _B_. coccodes in Spighe e Paglia was not available. ' Many facts about each of these species and varietiesnthe Spiralled appearance of the spore interior, spore size, K reactions, the sometimes yellowish thallus, and the distribution-{Indicate that they are probably members of Bombzliomra sect. Spiroblastia. LECIDEA mmmmcuwus Nyl. Ann. Sci. Nat. Bot. Iv. 11:228. 1859. W guadrilgggaLis (Nyl.) Mass. Atti. I. a. Istit. Veneto III. 5:262. 1860. Lecanoga W (Nyl.) Nyl. Ann. Sci. Nat. Bot. Iv. 16:378. 1861. W sect. Tetrophthalnidium tetranerum (Nyl.) Man. Arg. Bull. Soc- Bot. B9181(1‘10‘328135o 1893. WWW. 1+8 Ark. Eat. A 20(9):9. 1926. Blastenia W (Ny.) Zahlbr. Cat. 11101). uni-Vs 731‘8e 19310 Cfloglaca W (Nyle) Eahlbre Cat. 11011. M's 7:202e 19310 W seats momma W, (Nyl.) Gyel. Acta Faun. Fl. Univ. II. 1(5—6):8. 1933. TYPE: collected by Wedd on Cinchona 93.33% in Tipusn‘i, Bolivia. W. In my conception of the genus Bomgggogmra, this species (would belong-to it only if the spores develop in the character- istic manner. I have not been able to see any _I_.. W but the literature infomation does not exclude it from Bommgogara, where it was placed by the author of that genus. Nylander (1859) described the species as having spores usolidae h-loculares loculis contractis sphaeroideis invidem saepius tubule Junctis." Malme (1926) left out that important word 'saepa,‘ writing 'h-loculares, loculis tube contrali conjunctis, intermediis lenticu- 18.111118, membrane in apicibus tenui." Hue (1912) stated that Nylander was wrong to call the spores h—locular; he thought Nylander saw, but didn't understand the significance of, the tube. ‘ The spores are eight per ascus. ellipsoid, 211-30 2: 9-11(12) /u; the thallus is ashy and rugose, the apothecial disk fuscous, the mar- gin pale. The original description includes no mention of algae, but Helms (1926), vho saw the type, stated that algae are numerous in the medulla of the margin. Zahlbruckner (1931) took Oglopisma W, from Brazil , to be the same as L. ggadz‘iIOMS, although MBJler (1893b) thought it was not. It has spores 22-25 x 12-15/u, middle locules distinctly larger, thallus white and granular, and there are algae in the apo- thecial margin. Thus there is a possibility that LeQdeg W Nyl. belongs to W, but I camot tell at this time. #9 It should be pointed out that Zahlbmckner (1931) erroneously based all the synonyms except the Bomgzliogggra and Lecanora Quad- 343533535. Nylsnder (1861) clearly stated, however, that his Lecanora W was based on his Legdea q_uadgilocula§_is. IOPAIIIUM Kdrb. Syst. Lich. Germaniae 10. 1855. fleterothecigm sect. Iopadium (K6rb.) Tuck. Syn. Lich. 2:57. 1888. LeQdeg subgen. Lopadium (K5rb.) Wain. Acta Soc. Faun. Fl. Fenn. 7:25. Diplotomma sect. Lapadimn (K5rb.) Jatta, Syll. Lich. Ital. 11.32. 1900. Lecidea subgen. Biatora E. Iopadium (K8rb.) Ham. et Clam. Guide Elem. Lich. 71. 1901+ (not seen) TYPE SPECIES: L. maizoidem (Ach.) KB'rb. We Travis. Spighe e Paglia 7. 1853 (not seen). TYPE SPECIE: (selected) Biatora tgcglog Mont. (=Iopadium leucoxanthum Spreng. Zahlbr. Lecidea subgen. 1.0pr I. Gymnothecium Wain. Acta Soc. Faun. Fl. Fenn. 7:25. 1890. TYPE SPECIES: Lec_idea leucoxarrtha Spreng. W (adapted from Santesson, 1952). Thallus crustose, heteromsrous, epiphloedal, ecortic'ate, K+ yellow (atranorine); medulla UV-; primary phycobiont trebouxioid. Ascocarp an apothecium of the molecanorine type. Spores usually 1znae, sometimes 2-8:nae, ellip- ticus, hyaline, fuscescent in age, murifom or su‘mmnform, endospores not thickened. Paraphyses almost simple, or richly branched. Conidia embasifial, ellipticus to flask shaped or dumbbell shaped; uppermost part of spores or entire spores sometimes brealdmg into conidia within asc'i. Oephdodia containing species of Scytonemataceae sometimes present . 50 BEIGANTIAEA GROUP mm. Thallus K+ yellow (atranorine), corticolous or muscle- olous. Apothecium adnate or stalked, up to 3.5 m wide, often gyrose am with sterile columns: margin yellow, orange, ochraceous, or black; disk gold, ochraceous. or nearly black: epithecia and amphithecial cortices K+ rose, rarely K+ blue. Spores 1:nae, rarely 2:nae, often fuscescent and K-I- rose. Cephalodia not present. LOPAIIUM cfr. DODGE]: Herre Bryologist 53:296. 1950. TYPE: collected by§_e_1_'_rg _s_.3_1_., on Qgercus M Pacific Grove, Calif.: in Herre hb., isotypes in Dodge hb. and US. mg. Thallus ashy, sorediose, corticolous. Apothecium up to 0.7 mm across, adnate; disk ochraceous pruinose, becoming black: margin black. Epithecium and ampithecium K+ rose; exciple rust colored; hymerrium with clusters of colorless needles in KOH: paraphyses branched. Spores 1:nae, rarely 2:nae, always hyaline, sometimes K+ rose, (115)59- 75 x (18)21-26/u, all septa regular in young and mature spores, making cells nearly round; spores sometimes with wide, colorless halo: spores not breaking into conidia. Discusgog. I am not sure the material I have seen is 3.4. m, but that is the only name available. The type description does not include K reactions or descriptions of apothecial tissues. The spore of this species is very interesting: its appearance and the way it breaks when crushed is very much like g. MEL—2%: I cannot Justify inclusion of L. cfr. gm in @mfiliomra, however, because the locules are not formed by endospore thickening. Material se 2- UIIITED STATES 51 PENTANA: Glacier National Park, Snyder Creek, 3400 ft, on cot- tonwood, M 1296, 25 July 1950. WASHINGTON: Mt. Ranier National Park, Nisqually River, 2700 ft, mm, 11 Aug. 1948; both in MSG. IDPADIUM Fusoommm (Dicks.) Mudd Manual Brit. Lich. 190, t. 3, f. 73. 1861. £19933 fuscolgteus Dicks. Fasc, Plant Crypt. Brit. 2:18, t. 6, f. 2. 1790. Patelléga fuscolutea (Dicks.) Hofm. Descr. Adumbr. Plant. Lich. 3:10, t. 65, f. 1. 1801 (not. seen). Pamela-3‘s Escolutea (Dicks.) Ach. Meth. Lich. 180. 1803. Lecidea fuscolutea (Dicks.) Ach. K. Svenska Vet. Akad. Handl. 266. 1808 (not seen). Lecidea aurantiaca var. fuscolutea (Dicks.) Schaer. Lich. Helvet. Spicil. 11.5:179. 1833(not seen). Biatora fuscolutea (Dicks.) Fries, Summ. Veget. Scandin. 1:112. 1846. Leeanora £931,313 var. fuscglutea (Dicks.) Linds. Trans. Bot. Soc. Edinb. 7:122. 1863. 1.1313..- ercthecimn fuseoluteumwicks.) Tuck. Gen. Lich. 176. 1872. Legidea 7T7” Bombyliospora fuscolutea (Dicks.) Shirley, Proc. R.) Soc. Queensland 6: 175. 1889. Smropodium fuscoluteum (Dicks.) Wain. Acta Soc. Faun. Fl. Fenn. 53:269. 1922. TYPE: collected in the Scottish alps by Ben Lawers. W. Thallus smooth, muscicolous. Apothecium stalked, up to 3.5 mm across, thalline margin with algae and with warts containing algae; only this write false thalline margin showing on young apothecia; amphithecium ochraceous. Exciple colorless, full of large white crys- tals usually visable only in polarized light, crystals sometimes faint- ly pink, always K-; paraphyses simple. Spores 1:nae, 82-106 3: 28-35/u, thin walled, septa regular in young spores, irregular in mature ones; spores often fuscescerrt and K+ rose, not breaking into conidia. Discussion. Stirton (1898) meant to include this species in Milt dea _ hit failed to make the comflnation. 52 The abundance of algae in the margin makes one wonder what is the true relationship of this species with Caloplac . The apothecium has a different structure from the other two Lopadia discussed; see ago So mammal _asee . UNITED STATE ALASKA: near terminus of Mendenhall glacier, elev. 150 ft, Juneau, M 28525, 12 July 1963; alpine smtmdt of Mt. Juneau, {Mg 28260,, 15 July 1963, both in M80. IDPADIUM IEUCOXANTHUM (Spreng.) Zahlbr. Sitzungsber. K. Akad. Wise. Wien, math.-naturw. Cl. 111(1):398. 1902. M lggcoxanthg Spreng. K. Svenska Vet. Akad. Handl. 1+6. 1820. Patenagia leucoxanthg (Spreng.) Spreng. Syst. Veget. 4(1):267. 1827, (not seen). gators leucoxantha (Spreng.) B51. Voyag. Indes-Orient. Crypt. 2:128. 1886. Heterothecium W (Spreng.) Mass. Misc. Lich. 39. 1856 (not seen). Xanthgcgzpia leucoxantha (Spreng.) C. M811. Bot. Zeit. 15:386. 8157. Spgropflum leucoxanthum (Spreng.) Wain. Ann. Acad. Sci. Fenn. A. 15(6):88. 1921. TYPE: on m obloggii‘olia Mutis, no locality given except I'trepicarum pugillus:" no collecter nnw. ‘ Lagdea bicolor Spreng. K. Svenska Vet. Akad. Handl. 1+8. 1820. Hetercthegum Qcolo; (Spreng.) Flot. Bot. Zeit. 8: 5511. 1850. TYPE: on rock in Guadalupe; collector not stated. Pamlia chgsocma Meyer in Spreng. Syst. Veg. 4(1): 329. 1827 (not seen). Legidea olivaceogfa Zenk. in Goebel et Kunze, Phat-mazeut. Waarenk. 1:152. t. 17, r. 6. 1829 (not seen). w W F33, Suppl. Essa Crypt. Ecorc. Offic. 53 111, t. L12, 1:. 35. 1837. WW (F'ée) Mass. Atti I. R. Istit. Veneto III. 5:262. 1860. TYPE? on W in Peru: col- lector not stated. m t_r_i_c_9_l_o_; Mont. Ann. Sci. Nat. Bot. II. 18:266. 1882. W W (Mont.) Travis. Spighe e Paglie 9. 1853 (not seen) TYPE: collected by Gaudichaud, on bark in the Sandwich Islands. Megalomm 2,1133 Flot. Linnaea 17:21. 1843. TYPE: no data. canora crocata Fee, Bull. Soc. Bot. France 20:315. 1873. TYPE: Egg gggg, from Brazil. legidea crocata var. W Fee, Bull. Soc. Bot. France 20: 315. 1873. TYPE: _Fée 121}, from Brazil. Spgrogdimn leucoxanthoides Wain. Ann. Acad. Sci. Fenn. A. 15(6): 87. 1921. Iopadium leucoxanthoides (Wain.) Zahlbr. Cat. Lich. Univ. 114305. 1927. Wu Thallus granular, often becoming coralloid, corti- colous. Apothecium adnate, up to 1.6 mm across: disk yellow pruinose or with ochraceous nodules. Exciple rust colored: ampithecium without algae: paraphyses simple. Spores 1:nae, 515115029) 3: 19Mm, thin walled, septa irregular even in young spores, spores often fuscescent and K+ rose, often breaking into conidia. Discussion. Stirton (1898) meant to include this species in his Miltidea but he failed to make the combination. I have seen one specimen of I... leucoxanthoides (W _2_1_§_§, Hawaii, in M50). Wainio described that species as differing from _I_.,. W only in the blue color reaction with KOH. The two "spa... Cies' have the same variation, as noted by Wainio, namely with ochra- ceous disks or with fuscescent disks ( the latter variety of L. £01132- m, var. epiphaeizae Wain., is apparently a nomen nudum), and they have the same distribution. is Wainio pointed out, the situation 58 seems to be analogous to that with _B_. vulgiga and its variety 811le- m, but incongruously, Zahlbruckner allowed L. W to be listed in his Catalogue as an autonomous species. This taxon is for the present treated as a variety of g. leucoxanthg since the signifi- cance of the color reaction is not fully understood. It is thought to be a matter of monoquinones vs. diquinones. Mategifl Egg}. Exsiccati. Malme, Lich. Austroam. 1+2 8: 393; Merr. II. 101. UNITED STATES FLORIDA: county unknown, Eckfeldt, 1886 (US): Faulkner‘s High Hamock, east of Kissinger Springs, McFarl__ig 191, 1L: July 1931 (men). ALACHUA COUNTY, San Felasco Hammock, 11% mi NW of Gainsville, m 18, 6 Feb. 1963 (MSG); scrub oak forest, 8 11:1. SE of Gainsville, 13122.02 53, 7 Feb. 1963 (MSG). HIGHLANDS COUNTY, Highland Hameoks State Park, Sebring, m 22, 11 Feb. 1963 (MSC mixed with g. vulpinaand g. domngenggs). LAKE COUNTY, Eustis, figs}; 1251, May 28-June 15, 1895 (MSG). NEW JERSEY: county unknown, _;_Lchn__e_l_:'_t_ L88 (US). NORTH CAROIINA: DUPLIN COUNTY, 1 mi E. of Beulaville, Culberson 10_2_5_z, June 1960 (MSG). WEST INDIES (118A: LA VILLAS PROVINCE, near Topes de Collantes (TB senator- ium), Trinidad Mts., Imshaug M, 2 Aug. 1959. OKEENTE PROVINCE, Loma del Gato, mania-cultivated area on slope of El Gato (towards sea), Sierra Maestro, Imshagg 23813, 12 Aug. 1959, both in MSG. HAITI: DEPT. DE L'OUEST, elfin woodland type thicket, ca. 5300 ft, road from Fomt des Pins to Same-Zombie, Wetmre 39_0_2_ pr. p. DEPT. DU SUD, Massif de la Hotte, near summit of Meme Macaya, 7000 ft. W 215, 27 July 1958, both in MSG. 55 JAMAICA: HANOVER PARISH, 131th Hill, 1809 ft, m 155529 and 11638, 211 Apr. 1953. ST. ANDREW PARISH, Bellvue to Mt. Rosanna, 3800 ft, Imshagg M, 1+ Feb. 1953; Bloxburgh Ridge, 3100 ft, Imshagg 15351. 3 Mar. 1953; mast of Coopers Hill, Red Hills, 2500 ft, Imshaug £113}, 27 Jan. 1953; Flora River, 2700 ft, Imshagg 1513322, 3 Feb. 1953; Halberstadt House to Bloxburgh Ridge, 2800 ft, Imshaug 1591. 3 Mar. 1953: John Crow Peak, 5750 ft, Imshaug 3.52264 27 Mar. 1953: New Haven Gap, 5750 ft, Imshagg 15115, ’4 May 1953. ST. ANN PARISH, Green Park to Claremorrt, 11100 ft, Imshagg 15873, 7 May 1953: Alexandria to Brown's Tom, 1800 ft, M 1.52%.: 7 May 1953. ST. CATHERINE PARISH, Holly. mount, Mt. Diablo, 27511 ft, Imshaug 115m, 28 Jan. 1953; Montpelier swat, 2300 ft, Lmshaug 11319;, 20 Jan. 1953: Mount Diablo, 2600 ft, Imshagg 1.22.2.2! 18 Dec. 1952. ST. THOMAS PARISH, north slope below Cuna Guna Gap, 2300-2500 ft, lmshagg m2; Fam Hill, 4000. ft. m M1, 21 Feb. 1953; south slope of Mossmans Peak, 5000 ft, Blue Mts., Inshagg M, 19 Feb. 1953, all in MSG. PUERTO FUZOO: near Barranquitas, Jelliffe, 22 Feb. 1927 (HIGH ex hb. Fink 1588). LITERATURE CITED des Abbayes, H. 1955. Lichenes lécolt’es en Cuiné'e Frangaise et en Gg'l'ée-d'IVO‘er. VIII. 3111. Instit. Fr. Afr. Noire 17 A3973 9 . . 1951. Trait’e de lichenologie. Paris. 1: + 21? p. Acharius, E. 1814. Synopsis methodica lichenum. Lundae. xiii + 392 p. Asahina, I. 1931+. Lichenologische notizen 5. J. Jap. Bot. 10:352- 357- . and S. Shibata. 1951+. Chemistry of lichen substances. (English edition) Tokyo. 171 + 2140 p. Bailey, F. M. 1891 . Contributions to the Queensland flora. Dept. of Agriculture, Brisbane, Bull. 9. Belanger, C. 18116. Voyage aux Indes-orientales, 2me part. Crypto- gamie. Paris. 192 p. Duby, J. E. 1830. Botanicon Gallicum seu synopsis plantarum in flora Gallica descriptarum. Pars secunda. Paris. p. 545-1868. Esau, K. 1961. Anatomy of seed plants. New York. xvi + 376 p. Eschweiler, F. G. 1833. Lichenes in Martins, Flora Brasiliensis seu enum- eratio plantarum in Brasilia etc. provenientium. p. 51 -293. Fae, A. L. A. 1820. Essa sur les Cryptogames des écorces exotiques officinales. Paris. 167 p. _______. 1837. Essai sur les Cryptogames des écorces exotiques officinales. Dememe partie. Supplément et révision. 178 p. Fink, B. 1935. The lichen flora of the United States. Completed for publication by J. Hedrick. Ann Arbor. x + 1126 p. Frank, A. B. 1886. Spezielle botanik, Kryptogamen, in Tennis, Synop- sis der pflanzenkunde, v. 3. Third edition. Hanover. xix + 675 p. Hale, M. E.. Jr. 1961. Lichen handbook. Washington, D. C. x + 178 P. 56 57 Hedrick, J. 1935. Lichens from the Yucatan peninsula, Q Botarv of the Maya area: Misc. papers VI. Carnegie Instit. Wash. Publ. 461. p. 107-114. Hellbom, P. J. 1896. Lichenaea Neo-Zeelandica seu lichenes Novae Zeelandiae. Bihang K. Svenska Vet. Acad. Handl. 21, Afd. 3 (13). 150 p- Hooker, W. J. 1820. Flora Scotica: or adescription of Scottish plants. Part II. Edinburgh. 29? p. Hue, A. M. 1892. Lichenes exotici a professors W. Nylander descripti vel recogrnti. Paris. 378 p. . 1912. "Licheni blasteniospori" Mass. Bull. Soc. Bot. France 58:67-86. Imshaug, H. A. 1957. The lichen genus £2ng in North and Middle America. Trans. Am. Microsc. Soc. 7 :246-269. Harbor, G. W. 1855. Systems lichenum Gemaniae. Breslau. xxxiv + 1158 p. Krempelhuber, A. 1878. Lichenes collecti in republics Argentina a Dr. Lorentz et Hieronymus. Flora 61 :h33-b39, 461-0611, 1176- 480. 4924196. 516-523. Lanjouw, J. 1956. International code of botanical nomenclature. Regnum Vegetabdle 8:1-338. Lynge, B. 1939. Index collectionum "Lichenes Exsiccati.” Supple- merltm I. Nyte Mag. Naturv. 79:233-323. Malina, G. O. 1923. Die Flechten der ersten Regnellschen Expedition. Die Gattungen Bomgzlio¥ra, Megalomra, Catillaria and @26- cggpon. Ark. Bot. 18 A 12 x 1-7. . 1926. Lichenes blasteniospori Herbarii Regnelliani. . Bot. 20 A(9):1-51. Massalongo, A. B. '1860. Esame comparative di alcune generi di licheni. Atti R. Istit. Venet. Sci. Let. Art. III 5:247- 276. 313-337- . 1852. Ricerche sull' autonomia dei licheni crostosi. Verona. xiv + 207 p., 64 tab. Montagne, J. 1845. Criptogamia o plantas celulares, i3 Historia fisica, politics y natural de la isla de Cuba, Segunda parte: Historia natural. Tom. 9. Paris. 328 p. . 18118. Sixi‘e’me centurie de plantes cellulaires exotiques nouvelles. Cryptogamae Taitenses. Dec. I et II. Ann Sci. Nat. III. 10:106—136. 58 Miller. J. 1881. Lichenologische Beitrége XII. Flora 60:81-88, 100-112. . 1882. Lichenologische Beitrage XV. Flora 65:291-306, 316- 322. 325-337. 381-385. 397-402. . 1886. Lichenologische Beitrflge XXIV. Flora 69:252-258. 286-290, 307-318. . 1887a. Lichenologische Beitrage XXVI. Flora 70:268-273. 283-288, 316-322, 336—338, 386-1402, 423-1429. . 1887b. Enumération de quelques lichens de Noumea. Rev. Mycol. 9:77-82. . 1891. Lichenes Brisbanenses. Nuovo Giorn. Bot. Ital. 23: 385-1104. . 1893a. Lichenes Neo-Caledonici a c1. B. Balansa in Nova Caledonia lecti nex non alii nonnulli ab aliss ibidem observati, quos enumerat. J. de Bot. 7:51-55, 92-94, 106-111. . 1893b. Lichenes, in Th. Dirand et Pittier, Prinfitiae Florae Costaricensis. Second enumeration. Bull. Soc. Bot. Belg. 32(2): 122-173. . 1891+. Lichenes Usambarenses. Eng. Bot. Jahrb. 20:238- 272. p. 273-298, 1895. de Notaris, G. 1849. Nuovi caratteri di alcuni generi delle tribu delle Pameliacee, ed osservazioni sulla classificazione dei licheni. Mem. R. Accad. Sci. Torino II. 10:365-389. Nylander, W. 1857. Enume’ration g’engrale de lichens, avec l'indication sommaire de leur distribution g’eographique. Mem. Soc. ImpEr. Sci. Nat. Cherbourg 5:85-146. Supplément p. 322-339. . 1861. Expositio lichenum Novae Caledoniae. Ann. Sci. Nat. Bot. IV. 15:37-94. . 1863a. Lichenographia Novo-granatensis prodromus. Acta Soc. Sci. Fenn. 7:14-15-5011. ______. 1863b. Lichenes, in Triana and Planchon, Prodromus Florae Novo-granatensis. Ann. Sci. Nat. Bot. IV. 19:286-382. . 1868. Synopsis lichenum Novas Caledoniae. Bull. Soc. Linn. Nomandie II. 2:39-140. . 1869a. Lichenes Kurziani Bengalenses, coll. II. Flora 52:69-73- . 1869b. Note sur les lichens de Port Natal. Bull. Soc. Linn. Normandie ILI. 3:0—15. 59 Persoon, C. H. 1811. Novae lichenum species. Ann. Wetterauische Ges., Hanau 2:9-20. Riddle, L. W. 1920. Noteworthy lichens from Jamaica. Bull. Torrey Bot. Club. 1111:321-330. Rudolph, E. D. 1955. Revisionary studies in the lichen family Blasteniaceae in North America north of Mexico. Thesis, Washington Univ., St. Louis, Mo. Santesson, R. 1952. Foliicolous lichens I. A revision of the tax- onomy of the obligately foliicolous, lichenized fungi. Symb. Bot. Upsal. 12(1):1-590. ' Shibata, 8.. M. Takiko, and 0. Tanaka. 1950. Paper chromatography of anthraquinone pigments. J. Am. Phar. Assoc. 72:2789-2790. Shirley, J. 1889. The lichen flora of Queensland. Proc. R. Soc. Queensland, 1888, p. 80-110; 1889, p. 33-103. Smith, J. E., and J. Sowerby. 1807. English botany 21+. London. Sprengle, C. 1820. Plantarum cryptoganricarum tropicarum pugillus. K. Svenska Vet. Akad. Handl. p. 116-53. Steiner, J. 1907. Lichenes Austro-africani. Bull. Herbier Boissier II. 7:637-646. Stirton, J. 1898. On New Australian and New Zealand lichens. Trans. Proc. New Zealand Inst. 30:382-393. Stizenberger, E. 1890. Lichenaea Africans. Jahresb. d. St. Gallischen naturw. Ges. 1888-89. p. 105-149. . Lichenaea Africana II. Addenda et corrigenda ex annis 1893-94. Jahresb. St. Gallischen naturw. Ges. p. 215-264. Thompson, D. 1961. On growth and fem. Abridged edition, J. T. Bonner, ed. Cambridge. xiv _ 346 p. Thomson, R. H. 1957. Naturally occurring quinones. New York. vii + 302 p. Tuckerman, E. 1866. Observationes lichenologicae. Observations on North American and other lichenes. Proc. Am. Acad. 6:263-287. . 1868. Lichens of the Hawaiian Islands. Proc. Am. Acad. 7:223-235. ______. 1872. Genera lichenum: an arrangement of the North Amer- ican lichens. Amherst. 283p. . 1888. A synopsis of the North American lichens. Part II. New Bedford. 176 p. 6O Wainio, E. 1915. Additamenta ad lichenographiam Antillarum illus- trandam. Ann. Acad. Sci. Fenn. A. 6(7):1-226. Wetmore, C. M. 1960. The lichen genus Nephroma in North and Middle America. Publ. of the Mus., Mich. State Univ.: biol. ser. 1(11):371-’+52. Zahlbruckner, A. 1905. lichens in Engler and Prantl, Die natflrlichen Pflanzenfamilien. Edition 1. Teil 1, Abteil 1*. ' . 1926. Lichens in Engler and Prantl, Die natfirlichen Pflanzenfamilien. Edition 2. 8:61-270, 96 fig. . 1927. Catalogue lichenum universalis IV. Leipzig. 75*» p- . 1931. Catalogue lichenum universalis VII. Leipzig. 784 p. 61 Plate 1 Bpmbyliospora domingensis, showing rugose thallus, and gyrose apothecia with sterile columns. WW Phte1e 62 63 Plate 2 Bombxhgepom W, showing aggregated apothecia with rather thin margins. The apothecia are more regular in shape than those of other Spiroblaetiae. Plate 2- W ___vi_f1a dula 65 Plate 3 Bomggliospora vulpina, showing gyrose and aggregated apothecia, smaller than those of other Spiroblastiae, with fuzzy disks. 08 realm Plate 3. 67 Plate 4 Lopadium leucoxanthum, showing large apothecia nth fuzzy disks; pruina clings to the inner edge of the margins. The thallus is white and granular. Plate 1+. M leucoxanthum Q 1 ”1" J t e" 1 .‘\l ‘;1.1.~ _ \‘ 5 ) I ,3" Fig. 1. Fig. 2. 69 Plate 5. Apothecial Types epi--epithecium, hym--hymenium, sbh-subhymenium, exp-- exciple, alg--algae, med-medulla, cor-cortex, amp-- amphithecium, sub-substrate. Represents all K+ rose Bomgyliogporae and Lopadia except‘L. fuscoluteum. Represents‘L. fuscoluteum. Apothecial types Plate 5. ’1’“ 13‘ MED E116 ALP Fig. 1. AH IflD EXC Sm; “R EPIH'IM Fig. 2. 70 Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. 9. 71 Plate 6. Spores of Bombyliospora domiggensis Young spores in an ascus. The endospore has just begun to thicken; three lines of thickening are visible. Bulges on the inner wall are ridges of endospore. A.mature spore with annular endospore thickenings separating eight secondary loculee. An oblique view of an immature spore like the one in fig. 5. where the endospore thickenings are approaching each other from the sides of the spore, but have not yet met. An immature spore with eight protrusions connected by a central tube. (The above all in Turtomeedium) A mature spore with two annuli, and the rest of the endospore spiroform. The primary locule has become septate. Freshly mounted in Diaphane. A mature spore with a combination of annular and spiroform endospore. A week-old mount in Diaphane. A mature spore with two ridges of spiroform endospore. starting opposite each other. In Diaphane. A mature spore crushed in water: the outer wall has broken. allowing the two spiroform.locu1es to extend like released springs, and curve slightly to the right. The scale units represent 10 microns. Plate 6. Spores of Bombzliggmm dog agengl s 73 Plate 7. Other Spore Types Fig. 1-5 are Bomgyliospora vulpina Fig. 1. Fig. 20 Fig. 3. Fig. 4. Fig. 5. Fig. 6. Fig. 7. Fig. 8. Fig. 9. Young spore in ascus. All the endospore annuli have met in the center except the top one. In Diaphane. Two young Spores in an ascus. The secondary locules have a few'traneverse septa. (The below were all mounted in water.) Two mature spores in an ascus. Several locules of the lower spore are situated at an angle to the long axis of the spore. indicating that the endospore there is probably spiroforn. A.mature spore with the falsely-muriform appearance. The outer wall is ruptured, and individual secondary locules are released. Four are in the picture: one is lying nearly flat. L22§QLEE.Cfro‘QQQg2i. showing the minute, round, regularly- arranged spore cells, and the wide ”hale." W W, showing the minute, variously-shaped, lrregularly-arranged spore cells and the thin wall. A single secondary locule of‘B. domingeneis, lying flat: eight tertiary loculee are present, with one circle around a central locule. A single secondary locule of B. vulpina showing two concentric circles of tertiary loculee.. The scale units represent 10 microns. Pla te 7 . Other Snore . Type s 71+ 75 Plate 8. Spore development in Bomgyliospogg domiggensis Fig. 1. very young spore; the endospore has not begun to thicken. Fig. 2-5:the spiroform tendency Fig. 2. Fig. 3. Fig. A. Fig. 5. endospore has just begun to thicken primary locule has the appearance of an auger: the central tube is visible. central tube has been obliterated. primary locule has become septate Fig. 6-9:the lentiform tendency Fig. 6. Fig. 7. Fig. 8. Fig. 9. primary locule has just become ecalariform future secondary locules are becoming lentiform: the central tube is still present central tube has been obliterated; secondary locules are complete secondary locules have a few septa Plate 8. Spore development in Bomgyliogpgra domiggensis 76 77 Plate 9. Spore variation in Bomgzliospgra domingeggis Fig. 1. only one annulus is complete Fig. 2. half the spore has spiroform endospore, half, annular. Fig. 3. both ends have spiroform endospore, the middle, annular. Fig. 4. a young spore in KOH; the ends have annular endospore, the middle, spiroform. Fig. 5. ends spiroform, middle, annular. Fig. 6. upper end with annular endospore, lower with spiroform: one annulus incomplete. Fig. 7. most of spore with lentiform secondary loculee not forming acute angle at spore edge, and some septate; lower end with spiroform endospore. Fig. 8. half the spore has septate lentiform loculee forming acute angles at spore edge, half has septate spiroform locule. Fig. 9. septate loculee not forming acute angles at spore edge, all thickenings annular. Fig: 100 sparsely septate primary loculee; one complete spiroform locule, part of an additional one. Fig. 11. much septate, Single spiroform locule; all thickenings spiroform. WW Plate 9. Spore variation in Fig. Fig. Fig. Fig. 79 Plate 10 Mature spores of Bomgyliospogg aureola: the end two each have one incomplete annulus. Mature spores of‘B. flavidula. Septate secondary loculee of‘B. domingensis var. transgressa (Malme 1500 A3, MO) escaping from ruptured spore wall. Stylized diagram.of typical young spore of a member of section Bombyliospora; the endospore annuli are beginning to pinch the primary locule apart, but the spore does not appear to contain a tube. Plate 10 81 Plate 11. Spore variation in Bomgzliospgrg ep., Imshaug 15028 Fig. 1. Spore type found in most specimens, of least frequency in this specimen. Fig. 2. Spore type most frequent in this specimen, similar to that pictumd by Zahlbruckner in Engler and Prantl 1905 and 1926, for _B_. incan . Fig. 3. One end has spiroform locule, becoming septate; the remaining has annular thickenings. One spore was seen with entirely spiroform thickening. Fig. 11. The "membrane" surrounding each locule is greatly enlarged: it is a stmcture and not a cavity. The lines between are not cell walls, as shown by the fact that they do not extend into the endospore (compare Nephroma helveticum var. W in Wetmore, 1960). 83 Plate 12. Lines of evolution in Bomgzliosmra fig. 1. young spore, represents all species Fig. 2. B. domiggeneis, lentifom type Fig. 3. B. vulpina Fig. 14. B. flavidula Fig. 5. B. aureola Fig. 6. B. pachcheila Fig. 7. W sp., represents B. W, B. tuberculosa, B. incana, B. amplificans, and others. Plate 12. Lines of evolution in W 85 Plate 13. Distribution Map of Bomgliospora domiggensie Plate 13. Distribution Map of Wm W 87 Plate 11%. Distribution Map of Other Spiroblastiae solid circles = Bomgzliomra vulpina open circles = B. flavidula solid triangle = B. flavidula and B. aureola Plate 14. Distribution Map of Other Spiroblastiae C r" | ' 1 . ’J . \ I 1 I’ “‘ ,‘\ 1 ‘ - } ‘I 1 ' \ \. ~-_/Q..."’~ ~‘ A' I 5 1 1 1 I r . . -..- l.‘ . 1' I}. ‘ I 1 ’ O " f . 1 ’1 I ' I P — — v ‘- - -1 ' 1“ ‘0 9 o .) O I a f ? , I k <" , I) .l _ I 88 89 Plate 15. Distribution Map of Logadium leucoxanthum All maps are Good's Base Map No. 213. Plate 15. Distribution Map of LoEadium leucoxanthum $00M USE GMLY 'T’ ‘V ‘7. - r t... _, . , { -~: -‘ .l w -: f. r 1“ '. ' - ‘ I- ..~. ’.~-.'>~- ~' ado" t V l “X. I ‘ . ““ k“: .- I , . k -...c q..- P w . 5." . "WEI“EIHE'IJ Mimi?!" 1W '1'“