ANNALS OF THE MISSOURI BOTANICAL GARDEN ERRATA May cover, last line of Contents—186 should read 184. Page 65, line 29—var. nov. should read Sp. nov. Page 113, line 34, and on following pages of Gage's paper—chinensis should read chinense. Annals of the Missouri Botanical Garden Volume XL 1933 With 30 plates, and 41 figures Published quarterly at Galesburg, Illinois, by the Board of Trustees of the Missouri Botanical Garden, St. Louis, Mo. Entered as second-class matter at the post-office at Galesburg, Illinois, the Act of March 3, 1879. Annals of the Missouri Botanical Garden A Quarterly Journal containing Scientific Contributions from the Missouri Botanical Garden and the Henry Shaw School of Botany of Washington University in affiliation with the Missouri Botanical Garden. Information The ANNALS OF THE Missourl BOTANICAL GARDEN appears four times during the calendar year: February, May, September, and November. Four numbers constitute a volume. Subscription Price |... $10.00 per volume Single Numbers _____ 2.50 eac Contents of previous issues of the ANNALS OF THE Missouni BOTANICAL GARDEN are listed in the Agricultural Index, published by the H. W. Wilson Company. COUR! BOTAN 5 УЕ ne Rig t аак we MISSOURI BOTANICAL GARDEN STAFF Emeritus Director GEORGE T. Moore Acting Director Jonn S. LEHMANN Assistant Director EDGAR ANDERSON Executive Assistant LEICESTER В. Faust CARROLL W. DODGE, ROBERT J. GILL Mycologist Research К ROBERT E. WOODSON, GEORGE B. VAN SCHAACK, Curator of the Herbariu Honorary Curator of Grasses HENRY N. ANDREWS, JULIAN A. STE K, М н ist тенби ан d уле lite Rorra M. Tryon, Joun D. Dwyer, Assistent Curator of the Research Associate CUTLE ELL C. Hor Curator Museum of ае d | Editor Economic Plants of Publications GERALD ULRICI Business Manager BOARD OF TRUSTEES President JoHN S. LEHMANN Vice-President DANIEL К. CATLIN Second Vice-President EUGENE PETTUS DUDLEY FRENCH GEORGE Т. MOORE HENRY HITCHCOCK A. WESSEL SHAPLEIGH RICHARD J. Lock woop ETHAN A. H. SHEPLEY ROBERT BROOKINGS SMITH EX-OFFICIO MEMBERS ARTHUR Н. Сом RAYMOND К. Tuck Chancellor о of Washingt 1 Mayor of. d e City wy ARTHUR C. LICHTENBERGER, STRATFORD L Bishop ¢ ыы the Diocese of President of the Academy of Mis Science of St. Oscar EHRHARD President. of the Board of dn: of St. Louis GERALD ULRICI, Secretary ` TABLE OF CONTENTS Bluegrass Pasture Almanac. Alfred G. Etter The Popcorns of Turkey аваг Anderson and William L. Brown A Preliminary Survey of.the Milpa System of Maize Culture as Practiced by the Maya Indians of the Northern part of the Yucatan Peninsula. R. A. Emerson. А Contribution to the Lichen Flora of Arizona and New exico Emanuel D. Rudolph The Analysis of Suspected Hybrids, as Illustrated by Berberis X gladwynensis gar Anderson Variation in Cob Morphology among о. Archaeo- logical and Ethnological Races of M kontod H. Nickerson The Cytology, Morphology, and Systematic Relation- ships of Delphinium X Belladonna Hort. ex Bergm. Marilyn Amy Gage Some American Medullosas Henry N. Andrews and Sergius H. Mamay The Distribution of Diospyros R i с. arry R. e Wildwood—A Study in Historical Ecology |... Alfred G. Etter Some X-Ray Effects in Petunia. Henry A. McQuade Some Lichens of Tropical Africa... Carroll W. Dodge General Index to Volume XL PAGE 17-729 33— 48 51— 62 ег 73.78 795711 113—182 183—209 411-223 227-258 259—270 271—401 403—412 Volume XL Number l Annals e d. up rt Missouri Botanical | m n Annals | 4 о cof the о: врея Garden ar and the Henry Shaw School of dont of ity i in affiliation with. the Missouri Botanical Annals of the Missouri Botanical Garden Vol. 40 FEBRUARY, 1953 No. 1 BLUEGRASS PASTURE ALMANAC* ALFRED GORDON ETTER Malvern B. Clopton Experimental Farm, Clarksville, Mo. Washington University School of Medicine JANUARY The long summer leaves of bluegrass have turned brown, making the sere meadows and pastures of winter. In a few places, however, where these long leaves have been grazed off, pastures are still rather green. Bluegrass is also surprisingly green under certain trees and on the north slopes of hills. Isolated plants and small clones beneath apple trees in the orchard are brilliant green and are used by rabbits in preference to the tangle of brown plants in adjacent sod in full sun. Shading produces plants with leaves proportionately high in protein and low in fiber, which seem to satisfy the cottontails. Shading also restricts rhizome development suffi- ciently so that plants do not become crowded and thus sod-bound and nitrogen- deficient. After tree leaves are shed in fall, plants that were shaded all summer stay green well into the winter. When blades of these plants do die, they wither into very thin and delicate tufts, low in fiber, that contrast with the tough per- sistent dead leaves of sun-grown plants. Hogs on the hill pasture are working up moist areas into black patches of open soil. Where a bed of limestone outcrops above a bed of shale there are seeps which keep the ground soft and wet. Hogs have made the contact of these two forma- tions clearly visible across the hill (fig. 1). Their trails across the orchard-grass * The observations recorded in this study were made during bug! — s three-year residence on Brookhill не.) near Clarksville, Missouri. This farm and des pecial research fellowship were given ngton Univers ity н thé late Dr. Malve : S CH га? tes interest o Washin was in the касса of living things to the land. Grateful est is due Dr. Edgar Anderson of i Henry a ac y of dr of Washington University, and Dr. Arthur H. Compton, S Chan тА arranging this unusu niei Se = -— author. Much credit is also due the nis e "Aldo Leopold of the уник of Wisc hose inspiration err a background for these studies. nerous grant from the О. M. ro са Sons Company of Marysville, Ohio, =~ — it e As the author to organize his field notes and prepare the almanac for publica- e present pes is a companion pd to the author's "How Kentucky Bluegrass Grows, whic cb е eee in the September issue of the ANNALS for 1951 (Vol. 38, pp. 293-375). References to the — and орай 2 bits of toc tae will be made much more clear by referring to some of the diagrams in that p (1) [Уог. 40 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN pasture are also very apparent now. Їп a part of this pasture which was left un- mowed, the orchard-grass plants became rank and have been avoided by the hogs, not only in their feeding but also as they walk about the pasture. Consequently the pasture there has deteriorated into a rough tussocked run (fig. 2). During winter when the ground remains frozen for long periods, bluegrass leaves become thin, wiry and twisted, with tips curving inward, probably as a result of the difficulty of obtaining moisture from the frozen soil. January leaves are very short, only an inch or so long. In areas closely grazed since fall, stock are hard-put to it to find enough to eat. Horses are able to eat close enough, but hay is being fed cattle. The grazing habits of stock are distinctive and play an important role in determining the appearance of a pasture throughout the year. An important fact is that cattle and horses prefer grass which has grown in soil fed by the nutrients contained in their urine. They instinctively conserve the nitrogen, potassium, phosphorous, and trace minerals that urine contains and benefit from the stimula- tion which these substances, as well as dissolved hormones and vitamins, give the soil microorganisms and the herbage itself. This they do by actually seeking out the deep green grass which grows where urine has fallen (figs. 7 and 11). Analyses made by the author of grass in these spots has shown that it contains twice as much vitamin А (carotene) as ordinary grass, and Neven (1941) reports that the protein content of such fertilized grass may be nearly doubled. Horses graze small areas intensively, but cattle range widely. Urine is dis- tributed as the animals graze so that characteristic patterns of urine-fertilized grass develop. The pattern produced by cattle seems to be a rather random distribution over the pasture as a whole. With horses, however, urinations occur in small areas where the grass is kept closely grazed (fig. 7). The urine characteristically falls either on this short grass or, when grazing pressure increases, along the edges of these grazings (fig. 12) in which case it has the effect of continually enlarging the intensively grazed area (fig. 13). Of equal significance is the distribution of manure. Stock avoid grass around their own manure clumps as instinctively as they seek out that fertilized by their urine. Cattle manure is scattered rather indiscriminately though there is some considerable concentration in loafing and bedding areas. Horse droppings, how- ever, are somewhat controlled and deliberately concentrated in certain spots around the pasture. Often these spots are low places or fence corners; almost invariably they are weedy, full of rank vegetation. By keeping the manure thus concentrated and out of the way, their system of intensive grazing on small urine-fertilized areas is not interfered with. Since in summer, stock avoid grass around clumps of their own manure, "rough" areas of rank grass are protected from consumption (figs. 9 and 13). Considerable forage is thus saved for winter use. When the manure is frozen it no longer seems to be objectionable. In January, when new grass growth is at a 1953] ETTER—BLUEGRASS PASTURE ALMANAC 3 minimum, these areas are brought into use. Clumps of manure previously pro- tected by grass are now exposed, and to some extent are being scattered about the pasture. These "roughs" provide a convenient emergency supply of fodder when freezing rains and sleets fall. Urine grazings become a solid sheet of ice, but roughs, drifted with a little sleet, are still rather soft underfoot, and horses can paw them with their hooves, scrape away the crust of snow, and obtain all the grass they need (fig. 3). Cattle are not quite so adaptable and must be fed. No matter how heavy summer grazing may be, reserves of long grass around manure will usually be saved for overwintering. If fall months are very dry, however, grass growth will be slow throughout the pasture and “rough” reserves will be invaded during October and November so that before winter is over roughs may almost disappear. The past summer certain small blocks of pasture were enclosed with fence so that stock could not reach the grass within them (figs. 1 and 10). This protected grass has grown rank, and I can be almost certain that in each “‘exclosure” a rabbit will be spending the day. In fall and winter rabbits like the rich proteinaceous grass nourished by stock urine. By using these protected islands in the pasture they have access to much of that fertile grass, and their pellets are often found in closely grazed urine grass. The rabbit’s midwinter desire for proteinaceous food, which apparently ac- companies the advent of the breeding season, is satisfied by grazing intensively small plots of grass which have been fed urine by dogs, stock, man, or by the rabbits themselves. These grazing places are eaten down into the dirt with nothing but bare soil remaining (fig. 4). Such rabbit pastures are common sights along the runways of rabbits in winter orchards. They can also be seen occasionally in the summertime (fig. 20). In some places mice grazings are just as conspicuous. In October some urine was poured in a circle around a stake placed in an old Canada bluegrass meadow. Now the grass which received urine has been completely consumed and the ground is bare in a circle around the post (fig. 5). Even the rhizomes have been eaten. More striking still, where some of the urine had fallen on the post, the wood was chewed up by the mice. Gophers are no longer busy bringing up their piles of dirt as they were during fall (fig. 6). The extensive bulldozing they did then is slumping down with the frost and being scattered by treading hooves and being undermined by rhizomes of bluegrass. Most fall gopher mounds are built after temperatures are low enough to keep weed seeds from germinating. Also in fall and winter rhizoming and tillering of bluegrass are very active, and the new loosened aerated soil in the mounds encourages the vigorous vegetative development of bluegrass plants around its edges. The invasion and conquest of the mound go on to some extent all winter, for the loose soil does not freeze deeply. Revegetation of these mounds is [Vor. 40 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN also accomplished by plants which have been buried beneath the soil. When leaves and crowns are covered, internodes elongate and bring the crown up to the surface of the mound. Branching of rhizomes, while infrequent in most places, is the order of the day in gopher mounds. FEBRUARY Wherever the pasture thaws during the winter, stock cut the sod with their hooves and create large scars by sliding and slipping. These scars usually heal rapidly if made in early winter, but may remain for a considerable period if made in late spring. When the snow is soft, horses moving about build up pads of snow on their hooves. When these pads later slip off, many weed seeds and even vege- tative parts are carried about and planted. From a single pad picked up in Feb- ruary, 28 plants were germinated. Sixteen species were represented, half of which were grasses (Etter, 1948). During these cold months the rhizomes of bluegrass which began to grow in late October and November behave much as do the bulbs of such early spring plants as jonquils. Being situated near the surface of the soil, however, they are more subject to variations in the weather. They grow slowly in late fall, fitfully with thaws in the winter, and rapidly in the spring, shooting up and developing into new plants. At any time in winter one may find, in places where growth is rather vigorous and fall grazing was not too intensive, small rhizomes which are all white. Those which did considerable growing in fall have slightly green tips, and others may show a few short leaves. The dominant process, of course, during winter in every bluegrass plant is tillering. In old meadows this proceeds at a snail’s pace, but along streams, in loose, rich soil, where plants have room to stretch and root, it proceeds rapidly. More subtle is the secret development of the bluegrass seed head within enclosing sheaths. This process may have begun as early as November or may wait as late as March, depending on variety and environment. Winter thaws are important to bluegrass. With only a little rise in the soil temperature above freezing, fat white roots develop from the new tillers which have originated during the previous months. These roots, in turn, appear to stimulate new buds to develop into additional tillers. Where the soil is protected by an overburden of long leaves, thawing is slower and less frequent and roots do not develop very rapidly. Well-grazed or isolated plants, however, begin to send out roots a few hours after the soil thaws. This may play an important part in the earliness and the more vigorous growth of open-grown or spaced plants often observed along paths, in alluvium, in gardens, or near buildings. It is certainly true that bluegrass is often early to flower on the south sides of buildings and hills, above subterranean steam pipes, under eaves that drip warm water frequently in winter, or in places that are protected from snowdrifts as on the south sides of evergreens. 1953] ETTER— BLUEGRASS PASTURE ALMANAC 5 In meadows, now, there are well-camouflaged tunnels through which mice circulate in contempt of the winter weather. These are not mere pathways to and from, but feed alleys along which mice find their food. The stems of grass are cut near the ground to make the tunnel. When new leaves grow from the crowns of these severed plants, they stick up along the pathway. This growth is tender, bleached, and proteinaceous and is nipped off by the mouse as he navigates his pasture. These tunnels occur in heavy meadow grass where the overburden of leaves keeps the soil from freezing deeply, so that a small amount of blade growth takes place throughout the winter. Small feeding pockets extend short distances from the tunnels and are frequently filled with the grass-green pellets of mice. Undoubtedly the urine is also there. The floor of the tunnels and pockets is littered with clipped stems and pellets and is penetrated by a maze of bluegrass roots. When the ground thaws, the pellets disintegrate and a rich bed of mold develops, fed by these excretory products. The reserves of the rhizomes and crowns which feed the grazed plants are depleted by winter-long grazing and lack of sun, and tend, in spring, to produce weak growth which is often unable to reach through the overburden to the light. When plants such as chicory and plantain, which have bulky storage crowns, grow isolated in rank meadows they are apt to be consumed over winter by mice or gophers. Two plants, one of plantain, one of chicory, which I had marked last summer have been cut down and their roots consumed and transformed into a pile of pellets. MARCH A little warm weather in late February and March is soon followed by a slight greening-up of pastures. Earthworms return to the topsoil and robins arrive in time to take advantage of them. Flocks of starlings and purple grackles are often found feeding on these early spring pastures. During December and January bluegrass leaves tend to become short and prostrate, and leaves grow only about a half centimeter a week. When the days become longer and thaws are frequent, leaves and plants straighten up. New leaves shoot up from tillers and rhizomes which have developed during the winter. Since the snow, ice, wind, and treading cattle have broken the old tufts of grass in much of the pasture, these new leaves are not encased in the long sheaths of the previous year, and so become visible almost as soon as growth begins. Where leaves reach the light with a minimum expenditure of food reserves, they are in- clined to be more vigorous, broader, deeper green, and more attractive to livestock than those which have had to struggle up to find light. During this greening-up period, little further tillering occurs, and this is one time of the year when virtually no rhizomes are initiated. Any rhizomes still underground turn up and proceed to develop leaves. Often after this green-up first begins, the weather becomes cold again and leaf blades become reddish at the tips and growth almost stops. This redness, apparently anthocyanin accumulation, is especially common in the [Vor. 40 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN tips of grass recently grazed or cut. It seems to be a characteristic response of plants which are all ready to begin active growth and are prevented from doing so, either by injury, drought, low temperature, or excessively bright sun. During this spring renaissance, reserves in the crown, roots, and rhizomes are used up to produce new leaf growth. Sods become remarkably weak and may be pulled apart more easily than at any other time. | Wherever urine has fallen on the soil since last September, bluegrass shows ап early difference. Grass fed nitrogen does much more growing at low temperatures, below 40°, than does unfertilized grass (Blackman, 1936). At higher tempera- tures the nitrogen seems to have relatively less effect. Urine-fertilized grass is deeper green, thicker, and broader-leaved. Growth is also early in other sites where the ground is rich or moist, such as bare places once occupied by haystacks, near manure piles, in seeps, and around the edges of spots of sod burned out by urine last summer. Now that the sod is becoming soft, hogs are taken off the permanent pastures. Where they have been fed is a litter of corn cobs and mud which will become a rank bed of rich weeds in a few months. Horse grazings are now strikingly differentiated from the rest of the pasture which still is rather bleached (fig. 7). Tufts of grass fed by steer urine during fall and winter are now becoming visible, and cattle run in these early pastures seek out these patches. It is interesting that horse urine does not burn bluegrass when it falls on frozen ground. By mid-March, however, severe burning may occur. Cattle urine seldom burns grass even in mid-summer. APRIL In early April dormant buds on the bluegrass crown begin to swell, anticipating the spring flush of rhizomes that accompanies flowering. At the same time, blue- grass florets and panicles are growing, getting ready for the grand elongation of the stem which will occur in a few weeks. Long green blades of bluegrass are waving in the wind. Where pastures were not grazed last fall the blades are long, three or four inches, while in grazed places they are only an inch or so long and form small rosettes. There are many more blades in the grazed area, however. In rich soil, closely grazed, there may be twenty or more blades to a square inch. Trails and paths made through old meadows last summer and fall show up conspicuously now as stripes of green grass in a tangle of old whitish leaves. Break- ing down the sheaths and shading growth gave short fall and winter leaves access to winter sun. This stimulated tillering. A rough count of leaves along such a pathway showed twice as many new spring leaves as in undisturbed parts of the field. This path was used only one day, when it was walked over only five or six times. It demonstrates very simply how fall discing or mowing and raking of neglected pastures will do much to improve them. Down along the creek the succession of plants on newly built alluvial banks _ leads eventually, under grazing, to bluegrass. After gravel bars are stabilized by ee a eer мыд) 7999 1953] ETTER—BLUEGRASS PASTURE ALMANAC 7 sweet clover (Melilotus officinalis L. (Lam.), bouncing-Bet (Saponaria officinalis L.) and willow sprouts, coatings of alluvium begin to support a variety of weeds. Composites such as species of bur marigold (Bidens sp.), cocklebur (Xantbium sp.), sunflower (Helianthus sp.), as well as great ragweed (Ambrosia trifida L.), will also grow here, but will not withstand grazing and trampling. Rhizomes of Muhlenbergia racemosa (Michx.) B.S.P., broken loose from clones upstream, lodge on banks and begin to establish a rank stand of coarse grass which ties down the soil and speeds up deposition. As time goes on, bluegrass becomes established and invades and eliminates the other species. Without grazing, the succession would proceed from the coarse grass and weeds to shrub and tree growth. Grazed alluvial banks support excellent stands of bluegrass in the older established areas. Other grasses are developing rapidly now. Orchard grass (Dactylis glomerata L.) is flowering, and quack grass (Agropyron repens (L.) Beauv.) and smooth brome (Bromus inermis Leyss.) are beginning to develop rhizomes, while Canada bluegrass (Poa compressa L.) sods, devoid of new leaves, show many long white rhizomes which have not yet turned up. Redtop (Agrostis alba L.) is still almost dormant, a few green leaves beginning to appear. Purpletop (Triodia flava L.) has made virtually no growth, although its white dead stems are prominent in parts of certain pastures (fig. 8). On south slopes dandelions are just appearing, and in thin places a variety of small seedlings now appear. Old chicory (Cichorium Intybus L.) plants show only a few small leaves. Earthworms are abundant and are throwing up extensive castings within the secrecy of the grass. This subtle cultivation, accompanied by an increase in the numbers and activity of nitrogen-fixing bacteria in the soil, comes at the time when bluegrass needs it for rapid nitrogen-consuming leaf growth and inflorescence development. Moles are busy now too, tunneling in well-drained places, but gophers seem strangely absent. Cottontail rabbits are using wads of last summer’s long grass to hide their nests in. Young cottontails will be venturing from these nests very soon. By mid-April the blades on the flowering shoots of bluegrass have finished their development, but they are still telescoped within each other, waiting for the elongation of sheaths and internodes. The panicle or flowering head can be found just within the sheath of the newest leaf. It is striking to note that shoots of bluegrass which are going to bloom this spring can be easily selected now, because they have shorter and wider leaves than the blind or non-flowering shoots, and they are conspicuously a deeper bluer green. The short winter leaves are beginning to turn yellow, and this dulls the green- ness of the pasture very slightly. When stock graze the pastures now they are getting not only leaves but the developing flowers within. Mowing or close grazing at this time will greatly reduce the flowering of pastures. It will also produce a silage or feed as rich in protein as bluegrass can produce. It may, how- ever, be a little short on sugar. By waiting a few weeks for the panicles to develop [Vor. 40 8 ANNALS OF THE MISSOURI BOTANICAL GARDEN a little further a heavier yield will be forthcoming and the carbohydrate content will increase, though the per cent protein will drop accordingly. Old gopher mounds are now drying out and are very evident in pastures grazed continuously, for the grass there has not been given much chance to cover them (fig. 6). Bluegrass plants which grow on these mounds are very distinctive; the leaves are short and wide, deep blue-green, are prostrate, and often have much anthocyanin in their blades. Flowering shoots are consistently early to appear and grow almost horizontally. Winter pigs are on pasture now and they thrive on the high protein of the new grass. They are also busy rooting for worms and other foods in the loose soil under walnut trees. In a few weeks they will be seeking clover. Where hogs were pastured during the cold months, what grass remains is now a brilliant green. Hog urine has an especially strong, though brief, effect on bluegrass, and that combined with the well-distributed manure and treading stimulates the grass as it is stimu- lated in few other places. The excretory behaviour of swine is quite as interesting and significant as is that of cattle and horses. Hogs, when penned, are very careful to drop their manure only in certain corners. When on pasture they are careful to keep both urine and manure from contaminating the bedding area. Consequently, both waste products accumulate just outside the sleeping area, where sleepy hogs politely retire. Swine are just as careful not to defile their rooting areas, although here the prejudice is limited to the solid droppings. They do not hesitate to urinate in their excavations or in their wallows. Since hogs have need for various trace elements and antibiotics, it is quite possible that wallowing may be more than merely a way of keeping cool. The continual dabbling and rooting and munching that goes on while pigs search their wallows may provide them in a natural way with nitrogen, animal protein factors, and trace elements. Almost invariably, after rooting and wallowing, hogs leave along some much- used path, travel a few hundred feet, drop dung just off the path, and proceed with the day’s affairs. Where pastures are intensively grazed and where corn supplements, minerals, and water are provided all close together, these patterns are broken down and feeding areas are littered with manure. Frequent worming then becomes a necessity. In a pasture grazed last fall for a few months by steers at a rate of about two head per acre, there now appear as many as 20 urine spots in a plot 45 feet square, or about 1 square foot of urine grass to 40 unfertilized. In a pasture grazed longer and more intensively the spots are so close that I can easily step from one to the other. An important result of the stimulation of these pastures by urine is that bulky and less nutritious plants which might ordinarily be avoided are con- sumed to provide the necessary bulk. It is a common observation that the more rich proteinaceous food an animal has the more bulk it can handle. 1953] ETTER— BLUEGRASS PASTURE ALMANAC 9 It is rather easy to tell approximately when the urine fell which stimulated a certain piece of sod, for the vegetative response to fertilization depends on the season.when it is applied. Urine applied in fall promotes vigorous tillering which leads to short leaves and many short inflorescences with heavy loads of flowers and a decided blue-green color. Spring applications produce leaves as long as the in- florescences. ‘These shoots are weak, short, bear few well-developed flowers, and often lodge. With such information it is possible by examining urine spots to tell approximately when and how long an area was grazed. It is interesting that pig and calf urines have no such effect as do hog, beef, human, or mare urines. The difference in the reaction of the grass may be due partly to the characteristically lower nitrogen content of urine from a growing animal, or to a different hormonal content. The effect of urine on the grass of pastures is not limited to a single season, although the striking flush of growth in spring is the response most commonly seen. ere is a more lasting influence as a consequence of the fact that urine stimulates the development of grasses more than it does of weeds, especially when applied in winter. This rapid growth smothers out weed seedlings, clover plants, and results in a pure stand of bluegrass. Urine holds down weed development in a second way. From controlled tests made in flats kept in a greenhouse and planted to a variety of pasture weed seeds and grasses, it was found that germination of weed seeds was strongly inhibited in flats treated with urine, while grass seeds suf- fered little if any such inhibition. This effect was just as marked when urine was applied to plots of bluegrass in the pasture or on a lawn (fig. 14). In pastures grazed last summer by the horses there is much clover. Horse pastures characteristically contain more clover than those grazed by cattle. This is due in part to four facts: (1) Close grazing encourages clover, and horses graze quite close. (2) Compaction of the soil in certain areas maintains a favorable moisture condition for germination. (3) Breaking of the sod during winter in- duces germination of clover seeds, and horses cut the sod more than cattle. In a simple chopping test where a rank sod of pure bluegrass was cut with numerous strokes of a hand axe while the ground was frozen, clover germinated the following spring to such an extent that boundaries of the plot could easily be discerned. Winter discing also favors clover. (4) Most interesting relationship of all, how- ever, is that involving urine and grass. Horse urine that falls on growing grass usually kills it; cow urine does not. Burned patches soon become bare ground. While weather and soil conditions influence the flora that develops on these spots it is especially characteristic that spots which develop in dry mid-summer or fall remain bare until the following spring. At that time clover seedlings take over the bare ground and by April the patches are solid in clover. The attraction which urine-saturated soil has for earthworms and many other burrowing organisms may lead to extensive aeration of these spots and so encourage clover germination also. By the last week in April dandelions and peppergrass (Lepidium virginicum L.) are beginning to elongate, having already produced nutritious rosettes. Not only [Vor. 40 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN has bluegrass begun active production of rhizomes, but the panicles of early- flowering bluegrass plants have already been exserted. Silica deposition in the soft culms is going on and stock are avoiding them now. Where grazing is close and intensive the tough stem interferes with the efficiency of the grazing procedure. The stem does not tear when the cow pulls on the leaves, but instead the entire plant comes up by the roots. Dead plants pulled up in this manner litter the turf in places. At this season when growth is more rapid than at any other, a surplus of herbage soon develops and stock begin to pick and choose their preferences. А striking example of the attraction which urine-fed grass has for cattle now is shown in fig. 11. May During the first part of May flowering stems of bluegrass elongate rapidly, at least a centimeter a day. Ungrazed pastures become a sea of grass. Inflorescence elongation seems to have a depressing effect on the non-flowering tillers of the same plant, for many of them die during this period. They give the appearance of having been deprived of water. It is obvious that death of some plants and shoots must occur in a stand of bluegrass each year in order to maintain an even density over the years. In ungrazed meadow grass the greatest number of these deaths occurs at this time of year. There is also a gradual dying of the leaves on the inflorescence itself, all but the upper one or two having turned yellow at the tip. Beneath the soil new rhizomes are more abundant than ever. Some will turn up in the next few weeks; others may turn up sporadically throughout summer. All will turn up by early fall. It is a striking fact that new roots virtually cease to develop after April, though elongation of existing roots continues. Rank growth in the unknown meadows of last year, especially the prominent white stems of last summer’s purpletop (fig. 8), begins to blacken with mold, weaken and collapse. Lespedeza seedlings are coming up in patches of last year’s crabgrass, and both orchard grass and wild barley are showing their flowering heads. Cows drop their manure indiscriminately on the grazing places of horses, ignoring the careful system which the horses would maintain. Horses will avoid the grass around these cow-manure spots for a short time but in general the taboo against using grass around manure applies only within the species. In Germany, it was often the practice to apportion horses and cattle to pasture in such a ratio that rough areas were kept eaten clean, the horses cleaning up cow-dung patches, the cows the horse-dung grass. Cows are even now using some of the grass around piles of horse manure. Where horses graze with sheep, roughs are more evident (fig. 13); both species prefer the short grass, the sheep avoiding the grass around horse droppings almost as religiously as the horses themselves. Sheep drop most of their manure in loafing areas. What manure is dropped on the pasture is so easily scattered about that it seems to have very little residual effect. Grass fertilized 1953] ETTER—BLUEGRASS PASTURE ALMANAC 11 with sheep urine, however, has the same attraction as that fertilized by other stock. During the last part of April when bluegrass leaves are soft and green, aphids are active, sucking juices from them. Very soon, however, lady-bug larvae begin reducing the aphid population, searching every blade its entire length for their green prey. In two weeks more these larvae will fatten and develop into full- fledged lady-bugs. Now, in the last part of May, white clover (Trifolium repens L.) is flowering, and the inflorescences of Canada bluegrass are leaving the protection of surround- ing leaves. Cheat (Bromus sp.) is ripening, yarrow (Achillia Millefolium L.) has begun to bloom, ironweed (Veronia sp.) is 18 inches high, and the first blades of purpletop are beginning to show. Kentucky bluegrass has mostly flowered and seeds are ripening. Steers have shifted from bluegrass to rosettes of redtop and are grazing them intensively. Since redtop blooms later than bluegrass it is still in an early stage of development, growing rapidly, and consequently more palatable and proportionately higher in protein and lower in fiber than the more mature plants of bluegrass. This is a demonstration of the general tendency of stock to seek during each part of the summer those plants which are immature. After redtop begins to flower, the leaves of pigeon-grass, purpletop and paspalum will be sought, and new lespedeza will be consumed eagerly. By the time these late-developing species have matured, cool-weather species such as bluegrass will have begun to develop palatable new fall growt It is interesting to see young rabbits cutting the tender flowering shoots of bluegrass that grow on rich creek-bank soil, then eating them, flowers and all. Occasional they will prefer a dandelion stalk, which is consumed, stem, puff, and all. Dung beetles are actively digging their tunnels into the soil, then excavating fresh manure which they carry down into their burrows. The soil brought up is spread out over the top of the cow dung, serving to keep the dung from drying out, inoculating it with a variety of soil organisms, and hastening the course of disintegration and the incorporation of this organic matter into the soil. Dung beetles play a significant part in maintaining the fertility of a heavily used pasture for they keep the cycle of fertility operating efficiently with little waste of time or material. Where pastures are lightly grazed, or merely mowed occasionally, there is not enough manure to maintain a dung beetle population, and manure that falls often remains undisturbed and undecayed for years. Under those conditions little if any fertility is restored to the soil by way of animal waste products. It is also significant that stock on rank pastures, where there is little clover or rich green grass, produce manure which is bulky, hard, and low in nitrogen. This has little attraction for dung beetles hunting a rich medium for their eggs to develop in. Manure of stock on rich pasture is loose and moist, and decays readily. Machines which have been developed to cultivate intact sod by spiking or plugging the soil are following the precedent of the dung beetles, especially those devices which bring up small cores of soil and distribute them over the surface of [Vor. 40 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN the grass. Much of the benefit of this machine comes from the loose soil which it adds to the surface of the ground and mixes with the litter of grass stems, en- couraging their decay. In a small way, it simulates the natural process of flooding, whereby thin additions of alluvium from flooding streams are made to pastures on low ground. Plots of grass experimentally burned in February are now decidedly more yellow-green than adjacent grass not burned. Inflorescences are less green and less full. Since the ground was moist and frozen when the plot was burned and the fire not at all hot, there was little if any consumption of decayed nitrogen-rich litter from the soil surface. The only loss was the overburden of summer leaves. The evidence of nitrogen deficiency in the plants and flowering shoots of the burned plot suggests that the tangle of leaves makes an important contribution to the nutrient economy of an ungrazed meadow. A bluegrass pasture that is maintained by light grazing or mowing reaches a stability that is the result of a complex interrelationship of many factors. If we ook down at it closely, it is a veritable jungle with many kinds of insects walking in and out among the fallen stems. There are worm castings, mouse droppings, white molds, and minute fungi. A few attenuated weed seedlings try to reach the light. Bluegrass roots come up during the summer into this community and feed, and they are as dependent on a balance of these organisms as they are on rain. As a matter of fact, in this litter moisture is retained, and the molds and fungi, thus encouraged to develop, release nutrients and build up important organic com- pounds. Prolonged mowing experiments which remove leaves to be dried and weighed for assays discourage this natural process and inevitably interfere with production. Many such tests, repeated year after year, report unexplainable de- clining yields as years go by. There is little doubt that destruction of the reigning balance of nutrient release inhibits forage production. Summer roots have no place to feed. Disruption of the balance in a meadow sod by burning or continuous removal of clippings with consequent weakening of the cover encourages perennial summer grasses such as purpletop or broom-sedge which demand less nitrogen. With re- peated burning it has been shown (Curtis & Partch, 1948) that fire can weaken bluegrass enough to allow prairie grasses and forbs to become established. This weakening involves, in all probability, the discouragement of tillering. JUNE In early June bluegrass inflorescences are still mostly rather greenish-yellow, though some patches tend to be more purple, or red. There is also much difference in the heights and leafiness of the flowering shoots. At first glance one might believe that the different patches are clones, displaying inherent differences but such is not always the case. Bluegrass is a very flexible plant, and it is subject to an unbelievable variety of influences and environments. Urine spots alone cause the development of many false clones, either by stimulating certain patches, or by 1953] ETTER—BLUEGRASS PASTURE ALMANAC 13 encouraging grazing in certain places. The differences between fertilized grass or grazed grass, and adjacent grass which has not been treated is often so striking as to seem uninterpretable except on a genetic basis. Time and degree of fertilizing influence the length and number of inflorescences and leaves. Close fall and winter grazing will produce plants with numerous small inflorescences and short leaves. Lack of fall grazing results in fewer and much longer flowering stems and longer leaves. Spots entirely avoided by stock for one reason or another will con- trast with those which are grazed. Treading also has its effect. Where pigs pastured intensively in winter, blue- grass is hardly flowering; the few inflorescences present are much shortened. The combined influence of actual killing of crowns by hooves, and the weakening effect of continual injury and nitrogenous fertilization played a part in preventing blooming. It was found, however, that hoeing of all top growth of grass from the frozen surface of the soil during February did not prevent crowns beneath the surface from sending up flowering shoots in spring, but it did greatly shorten the inflorescence, and weakened recovery growth. This weakness persisted throughout the summer, and the area was invaded by many plants, such as horseweed (Erigeron canadensis L.) , milk purslane (Euphorbia maculata Г.) , Paspalum pubescens Muhl., boneset (Eupatorium sp.), purpletop, and white clover. Panicles on this plot ripened several weeks early, were abnormally white, and are already dropping seeds. During this flowering season inflorescences are often found which have barely appeared out of retaining leaves and then died. The heads are а gray-white color, and the condition is often called silver-top. It is caused, in many cases, by the larvae of a thrip which consumes the soft tissue of the developing stem at one of the upper joints. This severs the panicle from its source of water and food and prevents its further development. While this thrip sometimes causes trouble for seed-raisers, the severance of the seed-head at this early stage undoubtedly con- serves much vigor for the vegetative parts. Where silver top is common, the grass is commonly deeper green, but whether this is cause or effect is difficult to say. Chicory is beginning to flower now. In Missouri this plant is an abundant weed along highways and in pastures. It does not become especially noticeable until June, spending both spring and fall as a prostrate rosette. Stock like to graze it in its immature stages, and to some extent even after its milky fruiting stems have begun to elongate. The seed of the plant is used as an ingredient of some pasture seed mixtures in England and has even occasionally been tried in this country. It has a long large tap-root which penetrates the soil deeply and thus serves some of the functions of sweet clover and alfalfa. Where pastures are not grazed in spring, chicory plants develop into tall bushy weeds full of pale blue flowers. Although adding a touch of beauty to a pasture, they become tough and inedible and leave their resistant stems to clutter up fall and winter grass. In many cases where chicory has become objectionable, it is because the affected pasture was consistently not grazed in spring. If stock are allowed access to it at that time they will graze it closely and reduce its vigor. Fields along streams are LI [Vor. 40 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN especially subject to chicory infestation for the simple reason that floods frequently break through fences during the spring season. Procrastination in repairing these fences often gives the chicory time to develop and seed, and build up reserves. Then when pastures elsewhere are grazed out, the fences are repaired and stock turned in on the low fields full of mature chicory. This gets rid of the overburden of summer grass, so that in fall when the chicory plant again assumes a rosette type of growth and begins to store up food supplies, it has full access to sun. А single year's experiment to determine the effectiveness of spring grazing on restricting chicory growth was tried in some paddocks which had been ungrazed for some time and contained a heavy stand of chicory. Stock were allowed access to one paddock during April but were kept off the others till June. By July a strong reduction in height and vigor of the chicory stand was noted on the grazed paddock (fig. 18). The prominence of many weeds in certain fields can thus be traced back to the interrelationships of farm management and seasonal events. By mid-June the tedious mowing of permanent pastures has begun (fig. 15). Although it is customary to wait until the seed head reaches its full growth before any mowing begins, it is quite probable that earlier clipping might be advisable so that the resources expended on production of the inedible mature panicle might be conserved for vegetative development. The observations already made of tillers dying during the period of elongation indicate that the plant is under stress at this time. Earlier clipping would also prevent swaths from being so heavy. The blue- grass plant in the early flowering stage, when the panicle is just barely visible, is still fairly nitrogen-rich and soft and decays rapidly. As weeks pass, however, stems grow tough and siliceous, lie heavy on the new growth and disintegrate slowly. Even though the swaths are apparently obliterated in a couple of weeks by the rapidly growing June leaves, these leaves are forced to grow unnecessarily long through the weak light of the overburden and so deplete their food reserves. While it is essential to return this organic matter to the soil, it would be better to send it through the more efficient system, the animal, instead of trusting to the slow and expensive plan of decay. This is where the importance of grass silage comes in. Cut when grass is rich and generally in excess, and at the period when it is most beneficial to the plant, it is converted into manure and urine, rather than allowed to sink down slowly in an expensive disintegration. In well-grazed pastures there are now three main types of vegetation: (1) There are those areas in present use for grazing which are short, rich, green and non-flowering. These places indicate where urine fell this winter and spring.’ They are full of clover, buckhorn plaintain (Plantago lanceolata L.), and chicory, all of which are being readily eaten. The rich nitrogenous grass is eaten closely. (2) There are also those areas that were grazed last fall and winter, but are neglected for the time being. These show up now with short but well-developed inflorescences of bluegrass, mingled with chicory which has developed short flower- 1953] ETTER— BLUEGRASS PASTURE ALMANAC 15 ing shoots. Chickweed (Stellaria media (L.) Cyrillo) is common here, testifying to the fact that close fall grazing weakened the stand and exposed enough soil so that seed could germinate and the young seedlings receive sunlight all winter. Being winter annuals, they thrive best when winter competition is eliminated. Paspalum pubescens is also common in these places, since by the time fall and winter grazing occurs, this warm-weather species has already built up its reserves. Since fall-grazed bluegrass is slowed down in spring and its leaves shortened, the shoots of paspalum get off to a good start in May. Foxtail grass (Setaria lutescens (Weigel) Hubb.) is also beginning to become prominent in this short bluegrass. (3) A third kind of area distinguishable on the pasture is the rough about manure, where tall chicory and rank bluegrass grow, and little if any grazing is taking place. While these three conditions can be separately described as general tendencies where the pasture has been continuously grazed, they necessarily grade into each other to some extent. Heavy rains after a dry spell freshen up the bluegrass rapidly. Much of this rapid greening-up results from rhizomes which, caught underground during the dry period, were unable to move. With the rains and resumption of rapid growth many of them come to the surface. Most striking response is that of the grass along shallow drainages in the pastures. Much of the reaction in this case is due not only to the availability of moisture, but to the concentration of the nutrients washed down or leached from the pasture hills above. In response to the accumula- tion of soil and nutrients in these shallow drains, they commonly support a vege- tation distinct from that on adjacent slopes where removal of nutrient substances prevails. In one pasture, while the yellow soil of the slopes was thinly covered with a pure stand of Canada bluegrass the dark-soiled draws were distinctly marked off by tongues of Kentucky bluegrass. In another pasture, very poor and overgrazed, the hills were sparsely covered with triple-awn (Aristida oligantha Michx.), and lespedeza, with clumps of goldenrod (Solidago altissima L.) protecting a few re- maining patches of old top-soil. In the gullies were chains of undernourished cocklebur. Canada bluegrass is flowering now, and timothy heads are abundant in a few pastures and in the orchard. Redtop panicles are just now appearing. Ironweed flower buds are forming, lespedeza is coming up in pastures rooted by hogs this spring, and squirrel-tail grass (Hordeum jubatum L.) whitens the marshy swales in the hog pasture (fig. 16). This plant has the reputation of causing consider- able difficulty with stock, especially when its heads find their way into hay. The bristles on the grain cause mouth injuries and open the way for more serious in- fection through diseases. Occasionally now small grasshoppers appear in the grass, and the first horseflies are giving the saddle-horses trouble. In a few parts of a Canada bluegrass pasture which I fenced off, rabbits have grazed the grass very closely. In these spots, opened up to the full strength of the sun, seedlings of buttonweed (Diodia teres Walt.) and also of sheep sorrel (Rumex Acetosella L.) come up, giving the appearance of small clones. [Vor. 40 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN In a 15-acre pasture where a single horse had been grazing for 12 days I counted 18 newly burned urine spots on a plot 50 feet square. Since a horse averages around 7 urinations a day, in 12 days there would have been 84. If these urinations had been distributed evenly over the pasture there would have been only a third of a spot on 2500 square feet of the counted area. The concentration was actually 54 times that, indicating the extent to which grazing and urination are localized. The site of these counts was near the exclosure on the terrace in fig. 1. The soil was a rich fill and benefitted from subirrigation from the slope above. Cattle, after having been turned into a pasture where pigs had run all winter, found many of the rank weeds to their liking. They ate avidly of the lamb’s- quarters (Chenopodium album L.), tall smartweed (Polygonum lapathifolium L.), giant ragweed, and chicory (fig. 17). In a few days they had almost cleaned out even the Mexican tea (Chenopodium ambrosioides L.) and cockleburs (Xanthium sp.). They seemed to thrive on these weeds, even though the cockleburs are some- times considered quite poisonous in their early stages of growth. About the only plants that the cattle could not stomach were common smartweed (Polygonum Hydropiper L.) and lady’s-thumb (Polygonum persicaria L.). Consequently these two plants are taking over the old hog-feeding area. This indulgence in weeds took place at a time when plenty of good pasture was available. JuLy The grass of pastures which have gone ungrazed is rank and lax, and is dis- regarded or trampled underfoot by stock. Where bluegrass has been kept short by continuous grazing, weedy annual grasses such as foxtail and crabgrass (Digitaria sanguinalis (L.) Scop.) come in. The former is especially common in closely grazed urine places, and actually contributes considerable forage. It is beginning to bloom, however, and that reduces its palatability. In the pastures not so closely grazed, stock are feeding on summer perennials such as purpletop and Paspalum pubescens, new shoots of sweet clover (Melilotus sp.), prickly lettuce (Lactuca sp.) and new lespedeza. Pastures on low ground, partly as a result of extensive rains in June and July, are becoming much more rank and weedy than during previous drier summers. Bur marigold (Bidens sp.), Croton capitatus Michx., wild carrot (Daucus Carota L.), chicory, and Paspalum pubescens are coming up everywhere. When these weeds are cut during July mowing they leave a heavy swath which smothers out the grass and provides open areas where new weeds such as corn gromwell (Litho- spermum arvense L.) can come in. At the same time, in an exclosure on this moist and rich low ground where no mowing at all went on last summer, much grass has drowned itself out, and weeds such as bull nettle (Solanum carolinense L.), black nightshade (Solanum nigrum L.), sedges (Carex and Cyperus sp.), and pokeweed (Phytolacca americana L.) are invading. The new poke plants are lim- ited strictly to the vicinity of the fence, emphasizing that the chief means of 1953] ETTER— BLUEGRASS PASTURE ALMANAC 17 dispersal of poke seeds is by way of the alimentary tract of birds that eat the berries. Wherever convenient resting places such as fence rails, brush piles or fallen trees occur, there poke soon appears. As these rank weeds develop in the grass, they shade it out. They are soon followed by giant ragweed and other rank bottomland vegetation, and eventually by elms and walnuts. On such rich wet ground removal of some of the rankness of the bluegrass is absolutely necessary for its own perpetuation. This can be most profitably accomplished by grazing, though careful late summer burning occasion- ally can be used. Mowing and raking is an expensive alternative. Upland blue- grass is much more stable and less subject to this self-destruction. Although mowing in July is often done, it has little effect on pastures then except to shorten the grass for a time and postpone a little the flowering of certain prominent weeds. This postponement of flowering might help to reduce seed output were it followed later on with a second mowing. It often happens, how- ever, that such mowings are never accomplished. Actually, many of these weeds are rhizomatous species, and occasional mowing seems to help more than hinder their spread. St. John’s-wort (Hypericum perforatum L.) and chicory cut a few weeks ago already show flowers again. Ironweed (Vernonia sp.) comes back strong, and broad-leaved dock (Китех obtusifolius L.) is scarcely affected. When Canada bluegrass growing on exposed hills with little top soil is mowed, the hot summer sun has free access to the soil surface, which becomes dry and powdery. The slow process of restoring organic matter to these sterile places is consequently made even slower by indiscriminate mowing. Many farm practices such as manure spreading, fertilizing and liming, mowing and seeding, are done without reference to the natural conditions of the land. Мо effort is made to use ecological information to reduce the expense of wasteful blanket measures. Seed is not suited to the site, and treatment is not suited to the plant. The manure spreader is put into gear and manure flies over raw points and rich fills alike. As much seed is used on rich as on poor land. Alfalfa may be sown not only on suit- able slopes, but on the poorly drained swales between. Mowing is performed when and where the opportunity offers without reference to the developmental stages of the plants or the nature of the soil. Heavy swaths smother bottomlands, and on thin soils the shrivelling litter offers no protection from sun and rain. To achieve maximum production and balance on an uneven farm, the ecological factors must enter into the picture. Now that the spring drouth has been broken, rhizomes of bluegrass are elongat- ing again. The drouth period is recorded on the rhizomes in the form of several extremely short joints, or internodes which contrast with the new ones nearly an inch long. While the axillary buds on these rhizomes seldom develop under ordinary sod conditions, this new flush of growth has resulted in considerable branching from the first one or two nodes of new growth. This branching seems to be a characteristic response of many plants to a break in drouth. [Vor. 40 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN Gophers are throwing up mounds again. The fresh excavations are often dis- turbed and scattered by the mower bar during clipping of the pasture. They are also scattered by quail that use them for dusting places (fig. 21). These summer mounds usually develop a flora quite different from that of mounds pushed up in late fall. They are most commonly invaded by fleabane (Erigerom annuus (L.) Pers.), though also common are ironweed, boneset (Eupatorium sp.), thistle (Cirsium lanceolatum (L.) Hill), and other plants which germinate in summer, live for a fall and winter, and then flower the following summer. Old gopher diggings are often identified from a distance in summer pasture by patches of fleabane such as that shown in fig. 19. Bare soil areas are also prominent where horses stand head to tail, brushing constantly at the horse-flies that are now abundant. The continual stamping of their feet digs up the soil over a considerable area. On windy days horses usually retreat to some exposed point high on a hill where their efforts are assisted by the wind. In low shady places where the steer manure of last winter has remained moist, seed of honey-locust which passed through cattle undamaged have germinated, demonstrating one of the ways by which this plant spreads over pastures and waste land. Seeds of other plants also are able to navigate the alimentary canal of stock. From a small amount of hog manure planted in a flat with sterilized soil, bluegrass, nimble- Will (Muhlenbergia Schreberi Gmel.) , goosegrass (Eleusine indica (L.) Gaertn.) and Mexican tea germinated (Etter, 1948). AUGUST Although August, like July, is usually a hot month, its days are noticably shorter, and it is perhaps this difference that makes bluegrass change for the better toward the end of the month. Often following a good soaking rain leaves seem to become a deeper color and broaden out slightly. Stock will leave their summer diet of purpletop, foxtail, lespedeza and weeds to graze the bluegrass where it has | been kept short by summer traffic or close mowing, or where it has been grazed over by chickens for some time. A little anthocyanin may appear on the upper surface of leaves and on rhizome tips beginning to turn up. Some of the dormant summer buds on the crown of the plant have swollen. Most significant is the resumption of root initiation. As the month nears its end, short sprout-like rhizomes often develop, especially on meadow plants which have flowered. A rare plant or two, growing in a rich aerated nitrogenous soil, may even show the first development of a new tiller. This new activity depends on the availability of moisture, and if August is very dry resumption of growth may be delayed, to occur in a flush of activity after the first rain. While no amount of water in July can make bluegrass really palatable or healthy, irrigation in August and September is a different matter. By allowing fall growth to get away to an early start while days are still warm enough to encourage growth, total forage can be much encouraged and its nutritional 1953] ETTER— BLUEGRASS PASTURE ALMANAC 19 quality will be reasonably satisfactory. Where bluegrass pastures are depended on to provide overwintering forage in the fields, dry falls can be very serious. Other early signs that day length is getting short enough to produce results are the first fall flowering of dandelions and of annual bluegrass (Poa annua L.) and of the development from dry Canada bluegrass stems of new fall leaves which attract rabbits. The fall crop of grasshoppers also finds that this grass is begin- ning to have what they need to bring them to maturity, but many of them still prefer other more succulent sources of food. In lespedeza fields which were being saved for seed quite a few weeds were developing, especially prickly lettuce (Lactuca scariola L. and L. saligna L.). Grasshoppers invaded the field and while completely ignoring the lespedeza, they consumed leaves, seeds and stems of the weedy species, and almost entirely eliminated the weed problem before combining began. Grasshoppers are so uniformly castigated that it is interesting to observe also that they sometimes attack the giant ragweed with particular fury, consuming leaves and flowering heads before much pollen has had a chance to spill. Grass- hoppers are often just as choosy as cattle or horses about which plants they eat. In a field of ironweed, a very troublesome plant, grasshoppers may strip certain plants completely, leaving only the bare stems. At the same time adjacent plants may be untouched. lronweed is a highly variable species, and there are considerable differences in the leaf as to its hairiness, thickness, toughness, and greenness. There are probably equivalent differences in the chemistry of the tissues. It is small wonder that when they can afford it, the grasshoppers are particular. ugust is a time of spider webs, and in the morning dew-wet webs may be seen scattered widely over the pastures. They are also abundant in the woodland, and can sometimes be seen flying overhead on the wind. Mowing in mid or late August, while not a common practice, is very bene- ficial to pastures where summer grass has grown abundantly and deep. Mowing should be done just late enough to catch the last flowering shoots of the summer grasses such as purpletop, broom-sedge (Andropogon virginicus L.) crabgrass and foxtail so that no regrowth will occur. The removal of this overburden will per- mit the short fall blades of bluegrass to reap the benefit of all the fall sunlight which plays a big part in the amount of tillering that occurs as well as the width of leaves and the nutritiousness of the grass. This mowing should be done early enough, on the other hand, to allow rears fall top growth to occur. In souri, the third week in August seems a good time. Later mowing will reduce the crop of grass which may be used for fall grazing or saved for winter subsistence. SEPTEMBER The change which bluegrass began to feel in August has now brightened up the pastures except in ungrazed places where long summer leaves have begun to die and turn brown. Growth is rapid, summer rhizomes are turning up and leafing out, and in good soil a few plants may begin to form tillers. Just as in March, this September rebirth of activity results in a temporary absence of rhizomes. [Vor. 40 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN Except for a few short sprouts in the meadow or on wounded plants, they are almost impossible to find. In good soil which has had plenty of air, water, and sun during August vigorous plants develop, and these often begin producing rhizomes in September and continue all fall. This fall growth is at the expense of stored reserves, and the growth it produces is rather proteinaceous, though low in sugars. Summer rhizomes in the soil are depleted and weakened. Sods tend to fall apart rather easily as new roots develop. New leaves are definitely shorter and broader and stock show renewed interest in bluegrass. They may lose weight, or gain very little during this period as a result of the low sugar-high protein composition of the forage, and at the same time dairy cattle often show signs of acetonemia, or sugar shortage. Аз the days get shorter and the nights cooler, however, sugars will begin to accumulate and by November bluegrass will have a fairy high carbohydrate content provided the pasture has not been excessively grazed in fall. While work horses take readily to this new fall grass after being turned loose, they seem generally to prefer to consume considerable quantities of the more mature higher carbohydrate growth of nimble- Will and crabgrass before settling down to bluegrass and clover. Winter annuals such as wild brome or cheat are now developing, just in time to replace the summer grasses such as purpletop, now flowering, and crabgrass and setaria, which are becoming rank or dying out. Cheat is now four to five inches high and furnishes good pasture. It is much sought after by stock and will con- tinue to provide pasture until well into winter. It will also return early in spring to give stock an early bite. This is the wild counterpart of such crops as winter rye or wheat, often planted for late fall and spring grazing. Аза matter of fact, while man has laboriously devised through trial and error pasture systems involving winter annuals and hot-weather annuals such as lespedeza or Sudan grass to sup- plement permanent grass, nature does it automatically on pastures, and stock take advantage of it instinctively. On a hill which was seeded several years ago to lespedeza and timothy there is now a striking demonstration of the advantage of mowing at the correct time (fig. 22). Part of this field was mowed the latter part of June, while the timothy heads were still green and the stalks tender. The lespedeza seedlings were just getting a good start. The other part of the field was not mowed until more than a month later. The heavy strawy swaths can still be seen on this pasture. The lespedeza seedlings, shaded during their early development and then covered with grass, have scarcely produced any fall forage. Lespedeza in bluegrass also benefits by early mowing in late May or early June. OcTOBER If the fall has been moist enough, October pastures may be as green as those of spring, but drouth in September and October, a frequent occurrence, cuts seriously into fall grass production. The greenest grass, as usual, is to be found in disturbed areas, where vehicles have passed, animals or men have walked, or chickens 1953] ETTER— BLUEGRASS PASTURE ALMANAC 21 scratched, in gardens or where new soil has been deposited either by flooding creeks or eroding rains. In these places fall bluegrass plants produce many rhizomes and tiller early and are sought by stock. If the plants have been extensively injured during late summer, however, as by burning, or by the cleats of farm machinery or by hooves of swine, the recovery growth may be very deep green and actively tillering, but very few if any rhizomes will develop. In general, such rhizomatous pasture or meadow species as quack grass, smooth brome, or redtop follow patterns of development similar to bluegrass, though slight differences in time of maturity are reflected in vegetative responses. None of these species is actively developing rhizomes now. Hogs are feeding on bluegrass again, as they did during spring, testifying to the return of high protein forage. Cool weather and short days cause bluegrass to produce short leaves and prostrate plants. Stock graze closely seeking the nutritious foliage. They are invading grass protected during summer by rank foxtail or purpletop (fig. 23). Areas where urine fell last spring are still preferred, though little holdover effect is apparent. In pastures grazed during September spots of green grass are beginning to show where cattle urinated, but it is apparent that urine spilled in the hot weather of July and August produces virtually no results. The combined heat and dryness of summer permit the nitrogen to escape. Plots on which urine was applied artificially in August and September gave an amazing demonstration of the difference a month made (fig. 24). This fact has important meaning for grazers, for while it is commonly felt that animals return to the soil most of what they take from it, the truth is that much of the value of urine is lost when it goes on summer pastures. This might well influence the type of man- agement which the farmer chooses to practice during that period, especially the dairy farmers who return the waste products of their cows to the fields during summer. It is quite possible that summer applications of manure lose so much nitrogen as to be unprofitable. Storage under protected conditions until cold weather might be more efficient. Where pastures were not mowed in late summer the grass is very slow to develop and few if any tillers and rhizomes have formed. Fresh horse urine still kills the grass, but invasion of these burned spots is rapid now, at least in moist soil where adjacent plants have begun active rhizoming. Stock are avoiding the tall green grass around manure clumps. The cessation of root initiation during summer causes an accumulation of buds without roots. During fall this backlog is reduced, and by October roots develop as soon as buds mature. These fall roots are fat and white and contrast with the slender wiry roots of summer. There is now much mole activity, and gophers begin to build mounds out into closely grazed places seeking the succulent plants of dandelion and chicory. Clover is blooming a little, and aphids are back on the soft grass leaves. Under the oak trees where horses stand, fallen acorns are cracked beneath their hooves, and the soil becomes dry and powdery. Quail come to these trees, enjoy dust baths, and [Vor. 40 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN pick up fragments from the cracked acorn shells. In the woods pasture they follow the hogs that crack hickory and walnuts and acorns with their teeth. Frag- ments dropped to the ground provide the quail with a convenient meal. Bluegrass is often noticably superior under walnut trees. While this may be a complicated relationship involving microorganisms and obscure biochemistry, it also involves in part the fact that walnut leaves are shed early in fall, sometimes by mid-September, when bluegrass begins its important fall growing. Other trees often retain their leaves until November when all warm weather has passed. The shade cast in summer by walnuts is not dense enough to weaken growth, but is sufficient to limit rhizome development so that the grass does not become sod- u The sun of fall then allows tillering to progress and winter reserves to build up. To test the influence of shade on bluegrass between September and November, a shade of muslin sack was placed a foot above the sod at the same time that walnut leaves were shed. This shade was left on until late-shedding trees such as maples had become bare. It had a noticeable effect on grass leaves, making them more lax, waxy yellowish green, and delayed the death or browning of summer leaves. In addition it was later noticed that the body of the fall leaves was much different. The shaded ones were fragile and shriveled and gray, while those in full sun retained their shape well and died to a tough yellow brown. The repercussions of this treatment were not followed the next year, but undoubtedly influenced the quality of recovery growth in spring, as well as the size and shape of the florescence. NovEMBER In November bluegrass begins to suffer from the short days and the cold. Following the first frosts and the first freezes the blades shorten and cling to the ground. Rhizomes in vigorous grasses continue to be initiated, and to turn up, as they will, more slowly through the early winter. Nearly every new bud that develops now becomes a tiller. Gopher mounds are being invaded by rhizomes from adjacent plants. When the fall has been dry, stock begin to graze the rough areas where long grass remains. Clover leaves have been killed by the frost and swine are feeding on bluegrass, young chicory, and acorns. Canada bluegrass is just beginning to send out its rhizomes, which will continue growing off and on all winter and will turn up next spring to produce flowering shoots. astures where sows and pigs were running during August were so softened by heavy rains that the unringed sows were able to attack the sod and uproot much of it in their search for worms, roots and other food. These rough bare areas still remain, testimony to the results of delaying too long ringing of the sows (fig. 25). In spring these rootings will be a maze of weeds, DECEMBER Occasional frost has little effect on bluegrass, but in December when the soil freezes, bluegrass growth slows down greatly. With several inches of the top-soil frozen the leaves often turn dark, become flushed with anthocyanin. Wherever 1953] ETTER— BLUEGRASS PASTURE ALMANAC 23 bluegrass would like to grow but can’t, as in frozen ground, dry ground, intense sunlight, or because of injury to growing leaves, it has a tendency to develop this purple coloring. Uprooted rhizomes become pink at the tips or occasionally for their full length. This is especially noticable in Johnson grass (Sorghum hale pense (L.) Pers.) where the large fat rhizomes often turn a brilliant cerise when dug up and allowed to remain in the sun. Many other species seem to obey a similar compulsion. Around manure clumps, the grass remains surprisingly green, and bluegrass rhizomes work beneath the clumps with indifference to the cold. The manure insulates the ground and prevents its freezing, thus allowing bluegrass to continue its growth and to build up its reserves. As a result of this advantage, the grass in these clumps gets away to an early start in spring. Bluegrass leaves produced in December are very short, only an inch or two long. Except where the grass has been kept short, they are lost within the longer fall and summer leaves. When these old leaves die, pastures turn brown. In heavily pastured places, however, the grass is still green and stock still seek the sugar-rich grass. In many places, they are eating cheat seedlings in preference. Gophers are very active, throwing up mounds overnight, especially in the longer grass areas where the soil is not so solidly frozen. Mice are once again building extensive tunnels in the sod, and when snow melts away, elaborate feed- ing runs bordered with cut leaves can be found in the shorter grass, ending here and there in large “haystacks” of reserve food. Most fall rhizomes have turned up and now rest at the surface of the ground, though a few have been caught underground in the frozen soil. Tillering is still occurring, although the new shoots take several weeks to become large enough to be seen outside the retaining sheaths of their axillant leaf. Even more important than tillering, however, is the beginning of flower initiation. This process began on a few early plants last month, but is now general. A careful examination of the growing point shows a slight accumulation of extra plant segments at its tip, and it is from these that the panicle will develop. So it is that preparation for June pastures, full of waving bluegrass, is made even during this cold month half a year in advance. BIBLIOGRAPHY mec G E. (19 36). The ice cam of pro ger - available nitrogen supply on the owth of pasture in the spring. Jour. Agr. Sci. 26:6 Curis, Ja Au and Max L. App (1948). чаи of fire be Mere competition between bluegrass and ain prairie plants. Amer. Midl. Nat. 39:437-4 Etter, А. С. (1948). Seeds Hat db livestock. r M Gard. Sera 36:170-172. — — —, (1951). How Kentucky bluegrass grow . Gard. 38:293-37 ne У. В . (1941). Cow's urine as a fertilizer fe quies e Jour. De Sci. 24:761- [Ver. 40, 1953] 24 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 1 Fig. 1. January. On a pasture hill near the top, rooting hogs have defined the con- tact of a limestone and shale bed. Scepage along the contact favors their activity. Result- ing changes in the vegetation will mark this contact for some time. Fig. 2. Janu ry. Gilts in an orchard-grass pasture that was not mowed last summer beat paths among the resulting tussocks. Fig. 3. February. During periods when snow and ice cover the ground horses feed on tall grass preserved in manured or weedy areas. There they can easily scrape the snow away with their hooves. Fig. 4. February. Rank grass in orchards provides cover for cottontails. For sus- tenance, however, the rabbits prefer special spots where the grass is short and deep green. hese spots usually show evidence о aving received applications of urine. often grazed completely bare, roots, rhizomes and leaves all being consumed. Fig. 5. March. Urine applied in a circle around this post in fall resulted in com- plete utilization of all grass, including subterranean parts, by mice. Even the post was aten where urine had fallen on it. Fig. 6. April. Old gopher mounds of last fall are partly covered with grass, but new ones remain barren and spring winds whiten them. These mounds, especially the spring diggings, provide opportunity for weed seeds to germinate, but also produce grass which is darker green, productive, and preferred by stock. : ANN. Mo. Вот. Garb., Vor. 40, 1953 PLATE 1 ETTER—BLUEGRASS PASTURES [Vor. 40, 1953] ` 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 2 18. 7. March. At this time of year the pattern of horse grazing is very apparent. - Rank grass remains uneaten while small areas are consistently grazed. "These spots received continual applications of urine and remained green all winter. Fig. 8. April. Stems of last summer's purpletop stand in the new rank grass of spring. When temperatures rise sufficiently to encourage molds and mildews, these stems will drop into the surrounding grass. Fig. 9. April. Stock do not graze the rank grass round clumps of their own manure : and rough areas thus appear in closely grazed pastures. Where cows and horses are grazed | together these roughs are not so apparent since no prejudice seems to exist for grass con- | taminated by droppings of other animals. 1 ig. 10. May. The spots of darker and taller from steers during fall and winter. The resultin is preferred by stock. This pasture has grass in this picture received urinations — $ grass growth is more nutritious and — not yet been grazed. Fig. 11. June. When stock are turned in on a pasture such as that in fig. 10, they seek out the urine spots and consume the grass that grows there. Fig. 12. June. Horses graze limited areas intensivel these areas gradually expand. Urinations which burn the periphery of grazings. The dead grass will be replaced b will be grazed closely. Nutrit y. Under heavy grazing pressure - y rich weeds and clover which - ious bluegrass will soon invade the spots and restore the turf. ANN. Mo. Bor. GARD., Vor. 40, 1953 PLATE 2 ETTER—BLUEGRASS PA [Vor. 40, 1953] 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION ОЕ PLATE PLATE 3 Fig. June. When horse and sheep graze together stable rough areas are built up where horse manure accumulates, Sheep, like horses, prefer short grass. As grazing pressure increases rough areas are gradually whittled away. Fig. 14. June. This illustration shows the effects of urine on the vegetation of a lawn. Not only does the grass thrive in the right-hand plot, but weeds are eliminated. Application here was about two-thirds gallon per square yard and was applied in June, the previous year, on moist soil. 8. e. Mowing of bluegrass pastures commences. The heavy swaths will soon disappear beneath long summer leaves, but earlier mowing would have been desirable and would have maintained the food value of the grass at a higher level. Fig. 16. June. In the swales of an old hog pasture squirrel-tail grass heads out. This grass often causes mouth injuries to stock that consume it in hay. Fig. 17. June. Cattle turned into a new pasture in June sought first the rank growth of weeds that had grown up in areas where hogs had fed and urinated. The lamb's-quarters _ in the picture was cleaned out as were other weed species. Fig. 18. June. Chicory is a bad Pasture weed in some parts of Missouri. It can be controlled to a considerable extent by grazing it in s i er t grazed in spring at all while paddock in foreground was Grazing chicory while it is in the rosette stage weakens it and reduces the height of the coarse flowering stems. on zs га > < E un» o < 24 O 2 = р са а н H m i rol + EAE ۴ и [Vor. 40, 1953] 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 4 Fig. 19 у. Gopher mounds in rank pastures are often ries at a distance by a white growth of annual fleabane which germinates in the disturbed so Fig. 20. “August. Rabbits are choosey about where they graze. This small spot of crabgrass was grazed into the dirt. The grass was visibly darker pas and had apparently benefitted from urine of some species Fig. 21. July. Quail use gopher mounds for dusting places. Fig. 22. September. The time to mow is critical in pasture management and it varies We the еске In foreground а field of timothy and lespedeza was mowed while timothy heads were young and green. In distance where fall herbage is much inferior, mowing had bern pois. till July. Fig. 23. October. In late samme and fall ricus oat comes into rich heavily grazed pastures. For a while it provides some good mids azing, but, when seeding, is unpalatable. In fall, hungry stock invade the old foil beat it to the ground ^ November. Urine has à striking effect on bluegrass та in both fall and spring. In this picture, it is shown that this effect dep ends time of application. Grass in the left plot was given urine in late August; on da is a Loisir application in late September. There was no visible effect from the earlier is on. Fig. 25. December. Late s mmer rains softened the soil and unr ringed sows ie. the turf. If this oc. were disced and harrowed slightly i in early fall it odi diced recover. If allowed to go into spring in this condition it will be covered with weeds. ANN. Mo. Вот. Garp., Vor. 40, 1953 PLATE 4 ETTER—BLUEGRASS PASTURES THE POPCORNS OF TURKEY EDGAR ANDERSON, Missouri Botanical Garden N WILLIAM L. BROWN, Pioneer Hi-Bred Corn Company The special significance of popcorns in the history of maize has been recognized by many students of that perennially fascinating problem (see, for instance, Sturte- vant, 1894, Mangelsdorf, 1948). To scientists at large, however, the classification of popcorns and the description of the various ways in which they are used have been matters of such little moment that we were eventually driven to making our own world survey of popcorns and their uses (see Anderson and Cutler, 1950, for a general discussion). It was not until a decade of research had taught us to inquire in some of the most unlikely places that we learned (through Volney Jones) that popcorn is commonly used throughout Turkey and is a characteristic feature of many Turkish villages. At the very moment when we were endeavoring to have a comprehensive collection made for us there, Dr. Jack Harlan returned from that country with a collection of economic plants made for the Division of Plant Exploration and Introduction of the United States Department of Agricul- ture. Fifty-four collections of popcorn made by Dr. Harlan and his collaborators form the basis of the following report. We are indebted to Dr. Harlan for various observations supplementing the unusually complete data turned over to us by the Department of Agriculture, and to Dr. M. M. Hoover and his staff of the Regional Introduction Station at Ames, Iowa. It is a pleasure to acknowledge the efficiency and courtesy of this entire organization. We were welcomed to the increase plots in good weather and in bad, our attention was called to other collections of possible significance, and we were supplied promptly with viable seed of all the cultures we wanted to grow. For a few varieties of particular interest pertinent informa- tion was quickly produced from the files, and remnant seed of the original collec- tions was made available to us. The 54 Turkish popcorns collected by Dr. Harlan were grown in the experi- mental plots of the Pioneer Hi-Bred Corn Company at Johnston, Iowa, 20-30 plants being grown of each collection. А few varieties of particular interest were grown in replicate, and flint varieties and flint-dent mixtures from Dr. Harlan's collections were available for comparison in another plot. Each variety was scored for morphological uniformity, for season, for tassel type. Detailed measurements were taken on plant height, ear height, ear number, number of leaves above the ear, leaf length, leaf width, tassel exsertion, internode length and internode pattern, the width of the central spike of the tassel, glume length, the number of tertiary branches on the lowermost secondary branch of the tassel, the number of branch- lets of the fourth order (if any), the number of secondary tassel branches, and the pubescence of the leaf sheath. Ас harvest time the plants were scored for husk number, for number of apparent nodes in the shank (the difference between these two numbers represents the number of condensed [telescoped] nodes of the (33) \ Уот. 40 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN shank), shank length, shank width, kernel row number, kernel thickness, pith diameter, basal enlargement of the ear, pointing of the kernel, and color of peri- carp and endosperm. At silking time regulation photographs of representative plants were made against a measured background. “Inclusive” herbarium specimens (Anderson, 1951) were made as a permanent record of each collection, each speci- men including a photograph of the entire plant against a measured background and pressed central spikes and lower tassel branches. Sporocytes of each culture were pickled in aceto-alcohol and smeared in the cytological laboratory. The number of chromosome knobs (a variable and diag- nostic feature in maize—see, for instance, Mangelsdorf and Cameron, 1942) were determined and wherever possible knob positions were worked out. The results of these various measurements and scores are presented in Tables I and II. Photo- graphs of representative plants are shown in pls. 5 and 6. As soon as the collection began to tassel it was clear that maize is as hetero- geneous a mixture in Turkey as are many other crops in that area (see Harlan, 1951). The variation within collections and between collections exceeded that in any of the collections of exotic maize from various parts of the world which we have had under observation. Three extreme types and three mixtures and inter- mediates between these extremes were apparent in the collection. One of these extremes was a popcorn identical with the variety known as Japanese Hull-less Popcorn. Since it came from Ankara, the capital city, and since most of the intermediates between it and the other two types came from the vicinity of Ankara or Istambul, the older capital, it probably represents a comparatively recent introduction into Turkey. It was strikingly different from the other two extremes. It had pubescent leaves, a heavy tassel with an extremely thick central spike, and the ear was elliptical in cross-section with a high number of rows of kernels, all of these features being characteristic of the variety Japanese Hull-less. The other two extreme types were unlike рорсогпѕ commonly grown in the United States, and the two extremes, though connected by various intermediates, were quite unlike one another. One was very early to mature; the other was very late. The early type was short, with an exserted tassel, large glumes, and stiff upright tassel branches. Characteristically, the type had a low number of chromo- some knobs, usually from none to three with either no knob on chromosome No. 6, or merely one small one. The late extreme was tall with numerous short inter- nodes above the ear, which was borne high on the plant. The tassel was charac- terized by small glumes and lax branches, at maturity even the central spike remaining drooped over. The tassel was so included in the sheaths of the upper leaves that the lowest tassel branches were never completely free from the sheath of the uppermost leaf. Characteristically, it had seven to eight chromosome knobs, including two on chromosome No. 6 and one on No. 8. All these facts are set forth in the tables and the plates. Figures 1 and 2 show diagrammatically, but to scale, representative plants from the three most extreme collections of the two types. 1953] ANDERSON AND BROWN-—TURKISH POPCORNS 35 The early extreme bears no close resemblance to any known race of maize though it is vaguely similar to several. It is rather like a somewhat degenerate form of the Northern Flint corns which were once characteristic of eastern Nort America. It is shorter and earlier than most of them, with somewhat smaller ears, higher row numbers on the average, and without the more or less enlarged base to the ear which is one of their most distinctive features. The late extreme, on the other hand, belongs to a group of little-known pop- corns with an intriguingly peculiar distribution. They are commonly grown by various aboriginal tribes along the borders of China and India; they were in early prehistoric times the prevailing type of maize on the coast of Peru and Chile; and they are still to be found here and there in various out-of-the-way places in South America. We received some years ago from Lorenzo Parodi, of Argentina, a pop- corn of this general type and have a few scattered collections from Peru, Chile, and Colombia. Mangelsdorf and Oliver (1951) described and illustrated one from Caldas, Colombia. The collections from Turkey agreed with these Asiatic and South American popcorns in various technical characteristics of the inflorescence (see Alava, 1952)—in their knob numbers, the numerous ears high on the plant, the short upper internodes, and the drooping, included tassels. In its height, leaf shape, etc., the collection made by Harlan at Samsun was similar to varieties we have grown from the mountains of Siam, from the Naga tribes of Assam and from the Lushai Hills. Not only did it agree in general with these Oriental and ancient uth American popcorns; with a popcorn collected in the mountains of Nepal (P.I. 166,162 called to our attention by Dr. Hoover), it was virtually identical. The collections from Nepal and from Turkey differed not as much as would two strains of any open-pollinated (ie. non-hybrid) popcorn variety (such as Japanese Hull-less or South American) grown in the United States. The three extreme types described above were apprehended by repeated study of the plants in the breeding plot. The exact association of characters was worked out more objectively by means of pictorialized scatter diagrams. Опе of these is presented in fig. 3. It has been set up to demonstrate the relationship for the seven characters which best differentiate the early flint-like popcorns from the late “Asiatic” extremes. Аз fig. 3 demonstrates, there is in the Turkish collections an over-all association for these seven characters, tassel exsertion, percentage of total height in the internodes above the ear, glume length, chromosome knob number, shank width, plant height, and ear height. At the upper right-hand corner of the diagram are the Asiatic extremes. Аз the diagram demonstrates, they are all plants with ears high on the plant, the internodes above the ears being propor- tionately short. They have narrow shanks, small glumes, and high knob numbers. At the lower left-hand corner of the diagram are the opposite extremes, a more variable lot. They tend to be short, low-eared plants, with exserted tassels and proportionately long internodes below the tassel. They have wide shanks, long glumes, and few or no knobs on their chromosomes. It will be seen that though the Asiatic extremes seem to be somewhat set off from the rest there is a gradual Fig. 1 Three examples of the “Asiatic” race of popcorn, highly diagrammatic but exactly to scale CX 25). The lengths of the internodes, of the leaves, of the shanks, and of the branches of the tassel are all to scale, and the shape of the tassel is copied from photographs of fully mature tassels. Each diagram represents a different collection. Note the short internodes, the drooping included tassels, and the multiple ears high on the plant. NHGWVO 'IVOINV.LOH MNOSSIN AHL AO SIVNNV 10A]‏ ض0 1953] ANDERSON AND BROWN-—TURKISH POPCORNS а. EF /,. Ex d TM TUN da ы Lr а c. и hree examples of the “Aegean” race of popcorn, at the upper part of the plant, and the low ears F ү т p Fig. 2 У e| Lm тү, —. a to the same scale and prepared in the same way as fig. 1. Note the exserted tassels, the longer internodes [Vor. 40 58 ANNALS OF THE MISSOURI BOTANICAL GARDEN EAR HEIGHT IN DECIMETERS : v 13] * v У + 12} y éd 11 є Ф я xi i QN Ф, 6 Ri và 4 ‚ ёе at Ф é * $ у $ Ф, é | ө. 199 % «o é & eood 1 6% % rates VY we 5 € ò 6 © 6 4 e 6 e n „а. نہد ن‎ 11 12 13 14 15 16 17 18 19 20 21 22 23 24 PLANT HEIGHT IN DECIMETERS TASSEL EXSERTION GLUME LENGTH (MM) 0.3 -1 7-9 é Fos Ф v e 6-2 ы и - PERCENT TOTAL HEIGH IN UPPER INTERNODE Е "i 0-1 X li Ф 2-4 w. 5-6 e 35- 9 3-4 e 0-2 Fi Pictorialized scatter dem ее. ue variation in irs n of Turkish popcorns for seven hese age Each d a dia of a single plant for each collection, the plant having € osen in the field as pa e n e ds EK. aee ery Зет: Further explanation in the text. 1953] ANDERSON AND BROWN-—TURKISH POPCORNS 39 transition from one extreme to the other. The diagram presents an exact picture of the way in which two diverse races of maize have been brought together and have intermingled, forming a variable set of intermediates. Some of these are extreme recombinations in which characters of one race have been combined with characters from the other. For the whole group of varieties, however, there can be demonstrated the continuing association, om the average, of those characters which went in together. For those not familiar with the recent work on races of maize it should be pointed out that just such a mingling of correlated variables is characteristic of maize. It has been shown for Mexico y Wellhausen, Roberts, and Hernandez in collaboration with Mangelsdorf (1951, 1952), who have greatly extended the preliminary studies of Anderson and Cutler (1942) and Anderson (1946). For the maize of the American corn belt where purposeful mixing of two races has been carried on intensively, Brown and Anderson (1947) were able through these correlations to work out the history of the mingling by these indirect methods before carrying on the historical and archaeological research which proved that it had indeed occurred in just such a way (Anderson and Brown, 1952). There can be little doubt then that the heterogeneous popcorns of Turkey were largely derived from the mingling of popcorns from the two widely different races of maize, the resulting mixture being made somewhat more diverse by the com- paratively recent addition of still other types of popcorn. After the racial affiliations of the 54 collections made by Dr. Harlan had been worked out, their distribution in Turkey was determined from his collecting notes. The resulting picture is a simple one. The Asiatic race in its purest form is char- acteristic of the mountainous areas in northeastern Turkey. The other race is characteristic of the Aegean shores of Anatolia and we are accordingly referring to it as the Aegean race. It is possible to turn the pictorialized scatter diagram of fig. 3 into a crude sort of index running from 0 to 14. By plotting these index numbers in four grades it is possible to demonstrate the mingling of the Aegean and Asiatic races of popcorn as reflected by Dr. Harlan's collections (fig. 4). Conclusions: From the evidence presented above we conclude that popcorn is widespread in Turkey. Though heterogeneous there even for maize, it can be assigned to two provisional races, the Aegean and the Asiatic. The former is commonest along the coast, the latter in the northeastern mountains. Most of the popcorns of Turkey are various intermediates between these two extremes, occasionally modified by more recent introductions of modern commercial varieties. In the above report there is little or no evidence to indicate when or where or by whom these popcorns were introduced into Turkey. То answer such questions we have begun a series of ethnological, linguistic, and historical inquiries. These investigations have proceeded to the point where we are confident that though the problem is a complex one, it is not chaotic. It seems probable to us, in the . 4, Distribution of ав. pape — made by Dr. чи апа ove тай in fig. The numerous = in and around gre the capitol i т; been ous values o E n turned in crude - runn or D for typical "Aeg varieties to 14 for typical “Asiatic” varieties. Solid bla k dots eR ent c ollectio ons si like the Asiatic мча. e (those in de upper right-ha ad corner of dn ; open circles, Pa most extreme Aegean types; lightly stippled dots, intermediates ыы Аереап; heavily verae dots, "змс ав resembling Asiatic race. N3GWVO 'IVOINV.ILOH ГЯПО$$ПА AHL AO SIVNNV OF TOA] 1953] ANDERSON AND BROWN-—TURKISH POPCORNS 41 light of all the evidence, that the Asiatic and the Aegean popcorns were brought to Turkey at different times, and by different routes. Further discussion must be postponed until other evidence can be presented. LITERATURE CITED Alava, Reino, O. (1952). Spikelet variation = Zea Mays І. Ann. Mo. Bot. Gard.39:65—96. Anderson, Edgar (1946). Maize in Mexico: preliminary study. —€— 33:147-247. ; (1947). Field studies of Guat smalan maize. Ibid. - Ош). Inclusive herbaria. Jour. Ind. Genet. & РІ. 2 11: :1-3. , and William L. Brown Eu The history of the common maize varieties of the United States corn belt. Agr. Hist — — ——, and ‘Hugh С. Cutler ( Ts Races of Zea Mays: Their recognition and classification. Ann. Mo. jo Gard. 29:69-88. ————, — — ——, (1950). CE of corn popping and their historical significance. South- t. Jour. ae 6:303—3 FEES William L., and Edgar A read (1947). A Northern Flint corns. Ann. Mo. Bot. Gard. 34:1-28 Harlan, J. R. (1951). Anatomy of gene centers. ys Nat. 85:97-1 е ret Раш С. (19 si iion role of pod corn in she origin d evolution of maize. Ann. t. Gar “ ч :377 —_ ae s W. E (1942). Western си. 2 е аа center of origin of dire maize varieties. Bot. Mus. Leafl., Har жг 10:217-252. — —— —, and D. L r (1951). Whence came m Aper) ‘bid. 14:263—291. Sturtevant, E L. (1894). Notes on maize. Bull. ne Bor. Club 21:319-343, 503-523. ен п, Е. Ј., et al. (1951). Razas de maiz en Mexico. Foll. Tec. 5, Ofic. Estud. Езрес:, "M y Ganad. Mexico, D. F. 952). Races of maize in Mexico. Bussey Inst. Harv. Univ. TABLE I Turkish popcorns listed in the order of rod dcum pla seed w nt introduction number ach rity, Ja у мы For collection there is given the place where the ted, the season of matu of the plants grown from that dir genda the "bendi tae lant in decimeters, the height of the uppermost ear in decimeters, the number of ears, the y^ gth of the u pou internode in centi- meters, е M of фет! male чур: in millimeters and the Index number on a seven-fold index maa values of 0 nai treme “Aegean” types and of 14 for extreme “Asiatic” types (further ie ine in з s 8 K > ч > ч ta ч ЕР : Be toe c asdaM ia. d 5 sii g Е 2а |а 2 | 25 | iii) BEBO) D$ 4 #23 Я © i © i af | Нес ES ЕЕ - REA Е di - Mok) Mss | 25 |9258 | Wwe | ^^ Б 164,994 | Cayirhan, Ankara Mid Uniform 14 8 2 2.3 10 3 g 165,036 | Nallihan, Ankara Mid Uniform | 15 8 2 2.4 0 1 165,057 Beypazare, Ankara Mid Variable | 16 9 2 1.8 11 1 "i 167,030 | Kirik-Khan, Hatay Late Variable | 18 11 1 2.2 12 5 le 167,9 dd Balikesir Mid Uniform 15 5 3 1.9 10 5 m 167,948 | Bursa Early Uniform | 15 6 2 7 12 0 < 167,949 | Balikesir Early — 16 6 2 1.9 13 3 | 167,963 | Bucak, Burdur Late Uniform 18 9 2 1.8 8 8 Ф 167,967 | Osman Вики, Aydin Mid Uniform | 15 7 2 1.7 9 8 o 167,983 | Koycegiz, Mugla : Very Late pem 23 15 2 1.6 9 12 c 167,989 | Akhisar, Manisa 16 9 2 2.4 9 2 E 168,000 | Istanbul Mid A 16 8 2 24 9 4 Lc 168,006 | Kirklareli Mid Uniform | 15 10 2 2.0 10 4 O 168,007 | Kirklareli Mid Uniform | 16 10 1 ZI 9 4 H 168,008 | Kirklareli Mid Uniform | 18 11 2 Zi 9 5 > 168,016 | Tekirda Mid Uniform | 15 6 2 2.4 10 1 2 168,017 | Tekirdag Mid Uniform | 15 4 2 22 10 2 Ф 168,019 | From a seed s Early 16 9 1 2.3 10 2 р 168,027 | Ayvacik, Canakkale Early — 14 6 2 2% 12 0 168,032 | Edremit id 16 6 2 2.4 9 2 o 168,033 | Balikesir Early Usiforin 12 5 2 24 10 2 > 168,039 | Kulak, Balikesir Early Uniform 15 6 1 1.3 11 2 6 170,877 | Manisa Mid Uniform | 14 6 2 2.5 11 1 tr 170,879 | Soma, Manisa Early Uniform 13 4 1 Pi 12 2 2 170,880 | Istanbul Mid — 13 6 2 2.7 12 2 170,881 | Cayirova, rtr Mid Uniform | 18 10 3 1.6 11 6 171,896 | Inceoglu, Bartin — 13 10 3 1.4 10 8 e 171,912 | ' Unifor 22 13 3 2.5 9 14 = 171,915 Uniform | 23 13 3 1.2 8 14 3 171,91 Tokat Uniform | 22 15 3 1.0 10 13 è 172,600 indir, ата = 13 8 2 2.0 10 3 173,830 | Malatya Uniform 14 6 2 22 11 1 174,414 | Mardin 3 2 1.7 11 6 ани os " 75,976 167,030 170,881 170,881 171,904 Biga, Canakkale Biga, Canakkale rzincan Ayfon, Karahisa Araplar, ДАДЕ Balikesir > Ss o'm m > 25 B. = > ч p ecik, Ankara \lpagut, Sakarya Eskisehir Altinova, Ayvalik, Balikesir Araplar Sehir, Sebin Karahisar, Giresun Bolce Agac, Manyas, Balikesir Elbistan- us Igdir, Karahis Gemlik, Barsa Karga ili, Kastanou Ka are t-S Zo nguldak Kozam, i Kilic, Kesan, Edirne irne Lalapasa, ым eer a (near) Mani = чора Bafra, Sam Akashic J Kh isifite te, from farmers at Kirik-Khan, Hatay eli 0 Cayirova, Kocaeli Seed dealer in Samsun Early (Mid) Late Very Late Uniform for Uniform Variable Uniform Variable Variable Variable Uniform — ` чаи м سم‎ mm T nw m м олоо ое мм Ч eee NNN © NNN et ЫБ Ne Рә Рә Мә МӘ NEN NUN ж ie ies ка NIN NINN Ri Д wenn о œ N ее о — о їч Umea оошо ооо ооо МЧ Фо (о М фо їл оло? н m n س‎ ROW m | eS each] ide eR T ЧЫТЕ 6 ©‏ سم سر Rt‏ ي ی س طب ما \л‏ ++ m boo م‎ SNOOdOd HSIMYNL—NAOUT аму NOSYAGNY tv [£561 [Vor. 40 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE П Numbers and positions of chromosome knobs for the collection of = from Turkey. The ern at ti left are > eges ris introduction numbers of the Bureau of Plant Pu d Introduction. Numbe to the heading t the right refer pe the 10 chrom es of Zea r as rec бшш! a maize cytogenetics The final column, “В”, records the ct EE of “В” chromosomes, if an Wher mber is given, there were no "B" chromosomes. —1 indicates a s kicb on аб one of $ed E Seep, “ү” — small T" = terminal, эш минах Knob position 2 3 4 5 6 7 8 9 10 Р.І. 164994 4 2 —1|— IT >. I. 165036 1 1 >. I. 165057 4 1 1 1 fT P.1. 167030 4 1 1 1 DI P.1. 167947 Ж 4 2 1 LE >. |. 167948 2 0 1 1 0 0 » 1. 167949 3 0 1 1T >, I. 167963 6 1 1? 2 1 1 >, I. 167967 5 0 1 1 1 1 0 —1 |— >, I. 167983 6-7 2 1 Dr >, 1. 167989 2 0 1 0 0 › 1. 168000 4 1 0 —1 | — HT > I. 168006 1 1 0 ›. I. 168007 2 1? 1 >, I. 168008 3 0 1 ҮТ >, I. 168016 3 1 1 1T › I. 168017 4 1 1 1 rr >, 1. 168019 3 1 1 iT P.L 168027 1 1 >». I. 168032 3 1 1 0 1T 0 >, I. 168033 2 1 0 1 >, 1. 168039 0 >, I. 170877 3 1 1 fr >, I. 170879 1 0 1Т >. I. 170880 3 1 0 1 4 > 1. 170881 = 2 1 1 . L 171896 5 1s 1? 2 1T 5$ 171912 8 0 1 0 1? 2 1 1 2 0 » 1: 171915 7-8 15 2 1 1 >, 1. 171917 7? 2 1 1 HI 0 >. I. 17260 6 1s 1 1 2 PT 0 . 173830 3 0 1s 0 1 то '. I. 174414 3 1 1 0 1 0 0 0 „ 1. 175976 4 1 1 1 ae >. I. 175978 3 1 1 Ge 0 . 1. 176801 6 1 2 1 1 1T . I. 176804 1 1 0 . 177114 2 1? 1T . L 177583 5 1? 2 1 Жеш лч. 177598 3 1 0 1 IT] 0 І. 177622 2 1 1T . 1. 177624 4 1 0 1 TES . I. 177625 2 0 Is 0 1 0 0 Р. I. 177626 2 1? 0 1 Р. L17762 6 0 1 0 1 1 2 1? 0 0 Р. 1. 177631 5 1 1 2 1 . L 177637 3 1 1 0 . 1. 177641 5 1 i 2 с : . L 177643 2 1 0 " M а „179127 1 1T . L 179131 5 1 1 0 { iT 1953] ANDERSON AND BROWN—TURKISH POPCORNS 45 Number Knob position — of knobs |" 2 3 7 8 9 10 | B P. I. 179566 2 0 iT 0 P. I. 179568 2 2 P. I. 182324 2 0 1 1T 0 Р. I. 182326 4 1 1 2 P. I. 182327 2 0 1 1T Р. [. 182785 1 sd Р. 1.183750 4 1? iT P. I. 183751 5? IT P. I. 183756 4 2 1 IT P. I. 183764 1 1 P. I. 183771 3 0 2 IT 0 P. I. 183783 5 1 0 0 2 IT 0 P.1. 183787 5 2 P. 1. 183790 5 0 1 1 2 0 Р.І. 183798 4 1 1 1 1 1 P. I. 185045 3 1 1T P. I. 185063 3L 1 1 ІТ Р.І. 185073 2? 1 т P.I. 167030 2 1T P. I. 167030 5L 2 1 1 1T 0 P. I. 170881 3L 1 1 1 P. I. 170881 3 1 1 1 P. I. 171904 8 1 2 0 1 2 0 [Vor. 40, 1953] 46 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 5 Four plants of Dr. Harlan' s collection from Samsun, Turkey, photographed against a measured background, at the time when the X: were actively shedding pollen. Note the drooping tassels, barely exserted from the eaves, the bs ih on the plant. Specimen No. 4 has had its leaf blades and on $i its leaf күк сы аи The lines іп the background аге 25 cm. apart. PLATE 5 Mo. Вот. Garp., Vor. 40, 1953 ANN. ANDERSON AND BROWN—TURKISH POPCORNS : | Vor. 40, 1953] 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN ExPLANATION OF PLATE PLATE 6 The same “Asiatic” popcorn of pl. 5, photographed in the field at harvest time. e ants photographed in the experimental plot. Note a: ears high on the plane, -de ber g tassels, and the plant- to-plant uniformity. These plants were grown а actually collected in Turkey. The uniformity ot this variety is сысы ot collections made both in Nepal and in Turkey 2. Three plants from the same row as those in fig. 1, laid down on the earth of the дый plot. е the sare ears, high on the Slat: the leaves continuing to the verv base of the tassel, the drooping tassel кан the slender stem, and the short silks. These are all VETTEN А of Asiatic popcorn Mo. Bor. Garp., Vor. 40, 1953 ANN. u5 Z 22 О о a е a m ^ z ра р 1 Z 2 о ec © A Z < Z О Ze] га ыы A Z < А PRELIMINARY SURVEY OF THE MILPA SYSTEM OF MAIZE CULTURE AS PRACTICED BY THE MAYA INDIANS OF THE NORTHERN PART OF THE YUCATAN PENINSULA R. A. EMERSON FoREWORD In January and February of 1935, өн А. Emerson, of von Celery) an МЕ T. HE REDE, then of the U. S. Dep nt "| Agricult ture, studied m e farming a Maya Indians at the request of the Car beg а о My reins тие їп mimeographed form under Dr. Emerson’s name, is of the ta ir e results of this preliminary Though it has been cited in at least two bibliographies its very existence has been unsus- inl by many students of maize. indebted to Mrs. Ida K. he ee р the А Natural e of Philadelphia, for supplying us with a y, and to Dr. Н. E. D. Pollock, of the E of Archaeology of the Carnegie ій "tee Hort ag а to reprint the Up uii the time of his merson was the heart np center of resear rch in maize e on his part, has been identified with much of the research on maize which was ca mid o in the U Department of Agriculture and had taken the lead in зра maize and its dnd in rica. oe the “pe decade, Mexico, in age and Mexico’s — of growing maize, in particular, have changed a n ever- EEE v» e. Dr. Emerson's report, though preliminary and informal, ives us an itu of maize in an out-of-the-way part ie Mexico ат years ago as seen through the eyes of Б. rein maize scholar of his time,.—EpGAR AN PURPOSE OF THE SURVEY In the course of archeological studies of the ancient Maya civilization, con- ducted by the Carnegie Institution of Washington and other agencies, it has be- come important to estimate the density of population that could have been main- tained by the agricultural system then in use. Is it possible that the Mayas of one or two thousand years ago could have used a more intensive type of agriculture than that employed by Mayas of the present day? Could the fact that large cities and their contributing territories were abandoned, some of them to be rehabilitated only after some hundreds of years, have been due even in part to the inadequacy of the agriculture of these regions? Could a possible decline in the productivity of the agricultural land have necessitated abandonment after a century or more? Was such postulated decline in soil fertility related casually to the system of agriculture in vogue? It was not so much to find conclusive answers to such questions as these as it Was to form an opinion of the possible value of a more thorough and sustained agronomic study as an aid in answering them that a preliminary survey of parts of the former Maya region was undertaken. REGION COVERED BY THE SURVEY The survey was conducted under the auspices of the Division of Historical Research of the Carnegie Institution of Washington and under the tutelage of Dr. S. G. Morley of the Chichen Itza project of that Institution. It was begun the latter part of January and continued to the end of February, 1935. The (51) [Vor. 40 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN personnel consisted of Mr. J. H. Kempton of the United States Department of Agriculture, and of the writer. "Throughout the study they had the help of an interpreter and usually of a local milpero. The study of milpas was begun at Piste in the central part of the state of Yucatan near Chichen Itza and about 120 kilometers from Merida. Trips between Merida and Chichen Itza by automobile and by train over different routes, a trip by train from Merida southeast to Peto about 150 kilometers, and another by train from Merida southwest to Campeche about 180 kilometers, afforded glimpses of the western half of the state of Yucatan and the northwestern part of the state of Campeche. From a number of places along these lines of railroad, milpas at dis- tances ranging from a few up to 40 kilometers were visited by automobile, tram- car, mule cart, on horse-back, or afoot. Such points of departure were: Piste, Oxkutzkab, and Peto in the state of Yucatan, and Campeche in the state of ampec TYPE OF COUNTRY VISITED Topography.—The parts of Yucatan and of Campeche that were visited, as is said to be true of all of the northern part of the Yucatan peninsula, are a low, slightly undulating plain, broken in the southwestern part by ranges of low hills themselves only a few hundred feet above sea level. There is not even a suggestion of a river or brook in all that part of: the peninsula visited. Limestone rock.—The soil is underlaid throughout by a rather soft and porous limestone. Everywhere there appear outcrops of this rock. The sides and tops of small low knolls, often seen to be all rock, and the intervening areas, only a few meters less elevated than the knolls, exhibit many outcrops of rocks. In fact, it is never far from the surface to stratified rock. Although this soft limestone seems to harden on exposure after being taken from quarries, there are abundant evi- dences of its weathering to produce the usually scant soil. Everywhere exposed rocks are pitted with holes from the size of pin-heads to a meter across and half that deep. Both vertical and horizontal crevices of variable sizes are to be noted greater or less distances apart. Such crevices extend downward for considerable distances, perhaps even to the permanent water table which lies from a few to 50 or more meters below the surface, the distance apparently depending on the rela- tive elevation of the surface. Numerous caves and even large caverns exist. Not infrequently these extend down to or considerably below the water table. When such caverns are open to the surface, they constitute the natural wells, cenotes, of the region, upon which the Mayas depend for their water supply. Soils.—Evidences of the weathering of limestone rocks to produce soil are so universal that one is inclined to speculate on what would have been the nature of the land surface, had these rocks been exposed for ages to the alternate freezing and thawing in latitudes far to the north. Presumably the low plain would have been covered with some meters of fine loam instead of the scant soil covering now seen. And this might have resulted in a very different type of agriculture from that now prevailing. 1953] EMERSON-——MILPA AGRICULTURE IN YUCATAN 53 On the numerous low knolls the soil is seldom more than a few centimeters deep and on many of them soil is seen only in pockets, or crevices in the rock. Crevices seem to be filled with surface soil and humus to considerable depths, as can be readily observed in recently worked stone quarries. Between the knolls the soil may be a half meter in depth, in some places more, and in many places much less than that. The only relatively deep soils observed are those bordering the ranges of low hills in the southwestern part of the area visited. At Oxkutzkab such border soils were said to be as much as five meters deep. The near absence of surface stone and the luxuriant growth and productivity of citrus trees in this region make such an estimate seem not unreasonable. Evidently the heaviest rains have carried down to the plain enough sediment to build up a relatively deep soil at the foot of the hills. Yet, there is little evidence of even temporary waterways on the hillsides and no evidence whatever of brooks paralleling the base of the hills or leading away from them. The water presumably spreads out in a thin sheet over the plain, depositing much of its sediment before disappearing into rock crevices. These bands of deeper soil at the base of the hills are of very minor importance for the region as a whole, for they are seldom as much as a kilometer wide and in many places are very narrow. The surface soil, to a depth of a few centimeters on land on which the bush has been undisturbed for from ten to twenty years, is made up largely of decaying vegetable matter. During the dry winter season there is a sparse covering of dry leaves, nowhere the heavy covering seen in northern forests. Beneath this layer of leaves is a shallow layer of partly decayed vegetable matter and below that the red or brown loam. Evidently the relatively high temperature and considerable moisture of the region induce such rapid decay of the fallen leaves that no deep layer of decaying vegetable matter ever accumulates. Climate.—The entire peninsula of Yucatan lies within the tropics, the north- ernmost part being somewhat below the 22nd parallel of latitude. The rainfall is light in the extreme northern and northwestern parts of the peninsula. Near Merida and along the gulf coast as far south as Campeche, the annual rainfall is said to be not over 18 to 20 inches. At Chichen Itza the annual precipitation approaches 50 inches. The year is divided into two seasons, a summer rainy season of about five months, usually from May to September inclusive but some- times beginning nearly a month earlier or а month later, and a dry winter season of about seven months from October to April inclusive. Some of the weather records kept at Chichen Itza for the past nine or ten years were examined. The total precipitation for 1933 was 50.1 inches and for 1934, 46.6 inches. During the five months, May to September, of 1933, 86 per cent of the year’s rain fell; during the same months of 1934, only 73 per cent of the annual rainfall was recorded. Of the 730 days of the two-year period, rain fell on 275 days, 185 of them in the two five-month summer seasons. The maximum rainfall for one day, 4.75 inches, occurred in September, 1933. One-half inch or more of rain fell on 56 of the 275 rainy days, and 46 of the 56 were in the Мау- [Vor. 40 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN September periods. During the two-year period there were 168 days on which .2 inches or less of rain fell, 102 of them having been in the five-month summer seasons. The mean of the maximum daily temperatures for the years 1933 and 1934 at Chichen Itza was 91.6? F., and the mean of the daily minimum temperatures was 65.6° Е. The highest temperatures occurred in March, April, and early May, maximum temperatures of 101°, 103? and 105° Е. respectively having been re- corded for these months in 1933. When the rainy season begins the maximum temperatures are not as high. Minimum temperatures are lowest during the period from late November to early March. During 1933 the lowest temperature re- corded was 47^ F., which occurred twice in December; in 1934, 44? was recorded twice in March, and 43° once in December. The lowest temperature on record was 40° Е. on January 27, 1935. Vegetation.—Except for land in crops, the whole of the northwestern part of the Yucatan peninsula is covered by "bush". "This consists for the most part of trees and of vines which cling to the trees. In land not in crops for ten to twenty years, the dominant trees are 10 to 15 meters high and.10 to 20 centimeters in diameter. Under these are many smaller trees that have died or been suppressed by the shade of the larger trees. Relatively few annual weeds and almost no grasses are seen in any heavy bush. Both of these occur, however, along trails and in the more open places in the bush. In bush of only a few years’ growth, many annual weeds and small vines are to be seen. Among these are amaranths, numerous composites, and several kinds of bindweed. As the trees grow and shade the ground more and more, these weeds are less and less common. Even in very recently abandoned milpas little grass is seen except in the extreme northern and western parts of the area visited. Неге the many large henequen haciendas, in which the bush has been kept cut for ten to fifteen years, apparently return to bush less rapidly when abandoned and more grass is seen. The surface outcrop of rock does not seem to interfere much with tree growth. It is not uncommon to see large trees with roots exposed on the rock surface for several meters. Evidently the roots grow out in the thin cover of soil until a crevice is reached, then turn abruptly downward. In rock quarries roots 5 centi- meters in diameter have been found 2—3 meters beneath the land surface. THE MILPA SYSTEM OF AGRICULTURE Location of milpas.—Difficulty was experienced in learning why a Maya Indian selects a particular site for his milpa. The Mexican Government, in some regions at least, has assigned areas of bush around the several villages for the use of the inhabitants of those villages. If a milpa is made beyond the designated area on Federal land, a tax is exacted by the Government. Milpas have been visited which were 8 to 12 kilometers from the village in which the milperos making them lived. The factors that seem to govern the selection of a site are: type of soil, ease of 1953] EMERSON—MILPA AGRICULTURE IN YUCATAN 55 cutting the bush, nearness to the village, the presence of water, etc. There are undoubtedly differences in bush soils, though it is not easy to see the difference tween what the milpero calls good and poor soils. There appears to be a size range in bush trees on either side of which cutting the trees is more difficult. Trees of more than say 15 centimeters diameter are not easily cut with the small axes of milperos. Likewise, small trees sometimes bend under blows of the axe and must be cut with machetes. Moreover, and perhaps of more importance, in bush made up largely of small trees, there are many more trees per unit area. When milpas are made near a village, the milperos return to the village after their day's work. When a milpa is more than an hour's walk away—5 or 6 kilometers—the milpero usually moves to his milpa during the periods when he must work more or less steadily. In such instances, nearness to water, a cenote or a well, is of prime importance. The necessity of building a fence around a milpa, if near a village, where cattle and horses have free range, may often be a factor influencing the selection of a site far enough away to obviate the necessity for this extra labor. Making the Milpa.—A site having been selected, the milpero cuts the bush in the early part of the dry season, November to January, and allows the trees to lie as cut until they are dry enough to burn. The largest trees are usually left stand- ing. Among these are the ramon tree, the leaves and small branches of which are much used as forage for domestic animals, and the zapote, from which the chicle is obtained. The few palm trees, the leaves of which furnish the thatch for Maya huts, are almost never cut. The trees are usually cut at about waist height. Where fences are necessary, many of the smaller trees near the periphery of the milpa are left to be cut as needed for construction of the brush fence. In some milpas, some of the smaller trees are cut at heights of 2 or more meters, the stumps later serving as supports for the bean vines grown with the maize. In March or April, before the rainy season, the cut and then dry bush is burned. The milpero waits for a day of relatively strong wind and starts his fire on the windward side of the milpa. By this time many of the stumps left standing have ‚ Sprouts a meter or more long. This new growth is killed by the fire, and if the bush were a heavy one, the stumps and the large uncut trees may be killed, but usually, though considerably burned, many of them are left alive. Lundell, in his ‘Preliminary sketch of the phytogeography of the Yucatan peninsula,” refers re- peatedly to devastating fires which not only burn the felled bush but also sweep through the whole countryside. In this visit to northwestern Yucatan, absolutely no evidence of such forest fires was seen. Particular attention was given to this Possibility throughout the trip; and in no instance was evidence found of milpa fires having burned for more than a few meters into the surrounding green bush. This study was made before the season of milpa burning, but one large milpa fire near Chichen Itza was watched. Even with a strong wind, the fire burned only the dry leaves and small dead trees for only a few meters beyond the felled bush; and the fire had ceased to burn in the uncut bush while it was still burning hotly ; [Vor. 40 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN in the trunks of the larger felled trees. Sometimes а milpero takes the precaution of removing the cut bush around the edges of his milpa for a few meters before setting the fire, but most often this precaution is to protect from fire his newly erected brush fence. And perhaps this measure is only apparently one of pre- caution; the fence is most easily built with the cut bush that is nearest it. When the rainy season begins, which may be in April or as late as the end of May, the maize is planted. А sharpened stick, usually with a metal point, is used to make a hole in the ground. In this are dropped four or five kernels of maize. Seeds of a native squash and of the common bean (always a black-seeded one) or of a lima bean are mixed with the maize and all are planted together at random. No attempt is made to arrange the hills in rows, and they are spaced irregularly averaging probably a little less than a meter and a half apart. Not infrequently hills are planted in narrow rock crevices or in the deeper pockets in the surface limestone. If this were not done, considerable areas on the low knolls could not be planted at all. Маше seems to grow in rock crevices as well as trees do. In the deeper soils at the base of the range of hills in the western part of the area visited, maize is planted in rows about one and a half meters apart with hills not over half a meter apart in the rows. Sprouts of tree stumps and weeds that appear in a first-year milpa are cut with a machete from once to three times during the growing season of the maize. Very rarely is this done three times and probably more often once than twice. By harvest time the weeds and sprouts have grown to a length of from 1 to 3 meters. When the maize ears begin to mature, the stalks are bent over below the ears, so that the upper part of the stalks, including the ears, hangs down; this is done, it is said, to protect the ears from rain and from birds The maize harvest extends over a considerable period of time. It is not un- common for a milpero to harvest a basket of ears whenever the previously harvested basketful has been used up. By the early part of February, however, most of the maize has been harvested and stored. The ears are gathered into palm-leaf baskets, with the husks on, and piled near a thatched hut in the milpa. The part of the crop that is to be used or sold at once is husked and shelled. Ears that are stored for any considerable period are left in the husks. A crib of stocks is built a little above the dirt floor of the storage hut, and the ears are packed in it tightly in a vertical position with the tips downward. This method of storage is said to lessen injury from rodents and insects. In preparing a milpa for a second crop of maize, the weeds, maize stalks are not cut early in the dry season as was the b The cutting is delayed until toward the end of the dry season, it was said because: (1) The small cut stems dry out quickly and therefore need no long drying period; (2) if cut early, more weeds and stump sprouts would grow before burning; and (3) the relatively small amount of trash does not produce a sufficiently hot fire to kill late-grown weeds and sprouts. As a matter of fact, this trash has to be piled as Cut in order to insure its burning at all. A serious difficulty arising from this stump sprouts, and ush the year before. 1953] ЕМЕК$ОМ—МПРА AGRICULTURE IN YUCATAN 57 delayed cutting, seemingly overlooked by the Maya milpero, is that it affords time for the ripening and dispersal of an abundant crop of weed seeds which are not destroyed in any large measure by the fires which burn the piled trash. The first-year milpa, having been cleared and burned, a second maize crop is planted and tended just as the first one was. For the reason noted above, the weeds on a second-year milpa are much more abundant than on a first-year опе. On visiting a milpa it was rarely difficult to decide at once whether it was a first- or a second-year one, using relative abundance of weeds as the sole criterion. Occasionally, maize is grown on the same milpa for three consecutive years. On the narrow belts of deeper soil at the base of ranges of hills, maize is some- times grown on the same milpa for four years. But, in an overwhelming per- centage of cases, two years is the limit. The milpa is then abandoned to return to bush and a new one prepared by cutting and burning another area of bush. The length of time that a piece of land is left in bush before being cut again for a milpa varies greatly. Rarely, this period is as short as four or five years, and in some instances it is as long as fifteen or twenty years. There is insufficient evi- dence upon which to base a positive statement of the average length of this bush period, but it is probably not far from ten years. It would seem that the essential factor is a length of time sufficient for the dominant trees to reach a size satis- factory for cutting, and to choke out the smaller bush, and incidentally the annual weeds also. YIELDS OF MAIZE GROWN BY THE MILPA SYSTEM As in other places, maize yields in Yucatan vary from season to season. Throughout much of the area visited 1-1 cargas per mecate is a good yield for а first-year milpa, and one or а little less than one for a second-year milpa. It should be explained that a carga is 42 kilos, or about 93 pounds, approximately 125 bushels, and that a mecate is a plot of land 20 20 meters, or 400 sq. meters, almost one-tenth of an acre. While yields of 15 to 20 bushels per acre might we be regarded as unsatisfactory to a farmer in the corn belt of the United States, the surprising thing is that so good yields are obtained under the conditions pre- vailing in Yucatan. DECLINE IN MAIZE YIELDS That in general the yield of maize is somewhat less in a second-year than in a first-year milpa is everywhere evident. The bearing of this fact on the customary abandonment of a milpa after the second crop and its possible bearing on the withdrawal from a whole region by the ancient Mayas makes its cause or causes worth more than passing notice. [Vor. 40 58 ANNALS OF THE MISSOURI BOTANICAL GARDEN Soil depletion—An opinion, apparently somewhat generally held, is that the system of agriculture now practiced by the Mayas results in rapid depletion of soil fertility. Lundell, in his "Phytogeography of Yucatan,” says: The ashes are the only fertilizer the soil receives, so it is quickly exhausted. Little humus ept the land. The cr the s remains after fires have swep op is best the first year; econd year it falls off, and in either the third or fourth year the clearing is abandoned and another site he ab i i in a la is chosen. years, during which | the fertility of the soil is partly restored by the rank vegeta- tio: i EO. uos interva. Apparently continued rotation leads to complete soil exhaustio: Since the trees of northern and western Yucatan are hardwoods, and ashes from their burning undoubtedly provide readily available mineral nutrients for the first crop of maize, it seems unlikely that there can be much leaching of these nutrients, for, even after the maize is ripe, the weeds and stump sprouts continue to grow and are not cut until late in spring. It is true also that the fire which burns the dried bush in preparation for the first-year crop burns the few leaves then on the ground and destroys some of the scant layer of humus. Rarely, how- ever, does this destruction of humus extend for more than a couple of centimeters below the soil surface, and that far only where relatively large tree trunks have burned. The burning of stalks, weeds, and stump sprouts preparatory to the second crop has little effect on the remaining humus. This vegetation is so scanty that it has to be piled to insure its burning at all. One with an agronomic background finds it difficult to believe that milpas, after two crops of maize, have been abandoned because of soil depletion, and equally difficult to conceive of soil fertility, once depleted, being restored by a few years of tree growth. Even without an agricultural background, one might reason- ably question how such weeds as amaranths in a second-year milpa could grow to a height of 2-3 meters with a spread of branches nearly equal to their height if the soil were nearing exhaustion. Weed competition.—In bush of several years standing, such as is felled for a first-year milpa, there are relatively few annual weeds, and these are found mainly along the narrow trails and in the more open parts of the bush. Many of the seeds of these annuals are undoubtedly killed by the intense heat of the burning slash. But some certainly escape, for there is always a considerable number of weeds in a first-year milpa. Since the earlier appearing weeds are cut usually only once, and that during the maize-growing season, weeds continue to grow long after the crop is ripe and are still flowering and only beginning to ripen seed in ebruary; and since these late-growing weeds, along with the maize stalks and stump sprouts, are not cut until near the end of the dry season, there is oppor- tunity for the ripening and dispersal of an abundant crop of seeds. Only a few large weeds may produce enough seed to stock the whole milpa. The burning of scattered piles of trash late in spring can have little destructive effect on the weed seeds which by that time are widely spread over the soil. The system is an ideal one for stocking the second-year milpa with noxious weeds. It seems almost too 1953] EMERSON—MILPA AGRICULTURE IN YUCATAN 59 obvious to require statement that weed competition rather than soil depletion is the factor primarily responsible for the lessened yield of the second-year milpa. It seems equally clear that tree growth after abandonment of a milpa functions primarily in choking annual weeds rather than in restoring depleted soil fertility. The labor differential between first-year and second-year milpas.—Statements of milperos agree that more time is required to cut the weeds during summer in a second-year than in a first-year milpa. Moreover, it takes materially more labor to cut, pile, and burn the maize stalks, weeds, and stump sprouts in preparation for a second-year milpa than to cut and burn the bush for a first year milpa. Only from careful records of labor requirements can this differential be accurately determined, but it would not be surprising to find that the preparation and care of a second-year milpa necessitate from one and a half to two times the amount of labor that a first-year milpa requires. THE MILPA SYSTEM OF AGRICULTURE IN RELATION TO DENSITY OF POPULATION Dr. Steggerda’s study of the dietary of the Maya Indians of the village of Piste indicates that maize furnishes about 85 per cent of the Maya’s diet. е problem of food supply adequate for a given population is, therefore, largely one of how much maize can be grown in the region. Dr. Steggerda's records indicate that an average Maya family of five requires annually about 30 cargas of maize. In so far as can be determined from a hurried survey such as this, the usual yield of maize is 1 to somewhat more than 1 carga per mecate, and an average size of milpa for one family is probably not far from 40 mecates. It would seem, there- fore, that the Maya now grows enough maize for his family plus what the chickens, dogs, and other animals eat, with some to sell to provide the family with the cheap cotton clothes they wear and the few other purchased articles they require. Since a milpa is used ordinarily for only two years and then abandoned, and if about a ten-year interval in bush is necessary before the land is again used for maize, a crop can be produced on the average about one year in six. In other words, about 17 per cent of the entire area might be in maize each year. This statement is based on the assumption that none of the land is unsuited to maize production—an assumption that is almost literally true for the parts of the Yuca- tan peninsula covered in this survey. The water problem could be solved by artificial wells, such as are now in use. What percentage of this area is now in milpas in any one year is not known. But going over the country by rail, auto- mobile, mule cart, and on horse-back, one is impressed by the very small fraction of the country in milpas. This impression was strengthened by observations made in going by air over the entire northern part of Yucatan from Merida to the east coast of the peninsula. While only by accurate records of sample areas can one Obtain the necessary information, it would surprise the writer if such records, when obtained, show so much as 1 per cent of the land in milpa at a given time. [Vor. 40 60 ANNALS OF THE MISSOURI BOTANICAL GARDEN If this guess is not too wild to be credited, 15 to 20 times as much maize might be produced each year as is now grown, thus supporting that many times as large a population as is now found in the Yucatan peninsula, and this without changing the present system of milpa agriculture. Мо evidence was found in this short- time survey to indicate that the present system of milpa agriculture could not maintained indefinitely. It would seem, therefore, that 15 to 20 times as great a population as the rather sparse one now in existence could find an adequate food supply. Even this figure would probably fall far short of the tremendous density of ancient population postulated by some archeologists in moments of ultra en- thusiasm. The factual bases for such estimates are unknown to the writer and are perhaps beyond the powers of comprehension of a mere maize specialist. They do, however, call for a consideration of the possibilities that, in ancient times, a more intensive type of agriculture may have been practiced. In the territory covered by this survey, there is no evidence of the type of in- tensive agriculture inferred from mountain-side terraces such as those seen by the writer some years ago near Inca remains in Peru. Of course, no such things are to be expected in a low and flat country such as northern and western Yucatan. Whether the ancient Mayas employed fertilizers other than the ashes from burned trees or used partly rotted vegetable matter as a manure—as the present- day Maya Indians certainly do not—it is idle to guess. The advanced civilization of the ancient Mayas would certainly have been capable of inventing tillage implements. They had no beasts of burden, but the postulated population should have furnished man power enough to obviate the need of horses or oxen. Even though the ancient Mayas had no metal tools, they could have made plows and cultivators, as they did axes and knives, from stone and hard wood. But why waste time on such details? Even our modern steel implements of tillage could not possibly be used anywhere in Yucatan except in the deeper soils at the base of the ranges of hills. Even if the outcropping rock did not incapacitate such implements in the first half hour of use, the rocks would keep them from reaching the intervening soil pockets. The main basis for the opinion that ancient Maya agriculture was much like present-day milpa agriculture is the fact that no other type can be regarded as having a chance of successful use. The Mayas of today use the only method available; and the ancient Mayas presum- ably used the same system for the same reason. There remains one possibility not yet discussed. Could the ancient Mayas have kept the land in maize more than two years at a time, perhaps many more than two? Or, could they have shortened the interval during which the land was abandoned to go back to bush? Ву either of these ways the percentage of the whole area in maize at any one time could have been correspondingly increased. The agronomic problem involved in such longer or more frequent use of the land for maize is largely one of greater weed competition resulting in lessened yields or of more labor per unit area to hold the weeds in check. 1953] EMERSON—MILPA AGRICULTURE IN YUCATAN 61 Information obtained from Maya milperos indicates that it requires from one and one-half to two times as much labor to care for a second-year as for a first- year milpa. Cutting the weeds, stalks, and sprouts in a milpa as soon as the maize is mature enough to harvest, and thus before the weed seeds have ripened and been scattered, instead of waiting as at present until near the end of the dry season, should materially lessen the weeds to be fought the next season. But there is no way of knowing whether the ancient Maya's routine differed from that in vogue at present. How much time does a Maya spend in caring for a milpa large enough to supply his dependents with maize for the year? If the ancient Mayas used materially more than one-sixth of the land for maize each year, could the milperos have devoted enough time to their milpas to meet the weed problem involved? А milpero cuts about two mecates of high bush a day. In cutting the trash in prep- aration for a second-year milpa, he covers about the same amount of land, but this takes the greater part of a day, while cutting two mecates of bush for a first- year milpa seldom requires all day. Two mecates seem to be the day’s stint, irre- spective of how many hours of work are required. Rarely is more than one mecate cleaned of weeds in one day’s work during the growing season of maize in a second-year milpa. Under the present system, therefore, to prepare land enough for the maize needed by one family—say 40 mecates, or about 4 acres—and to weed it once in summer requires about sixty days of labor for one man. Burning the felled trees or the weeds and stalks, planting, and harvesting could hardly require more than twenty days. It follows that the present-day Maya milpero has about three-fourths of his time free from the labor necessarily connected with his milpa. The ancient Maya milpera grew cotton as well as maize, but the Maya of today grows enough more corn so that he can sell some in order to buy cotton cloth. Moreover, the ancient milperos must have grown enough maize for their priests and rulers in addition to the requirements of their own families. But even so, the ancients could have spent more time on their milpas than do their descend- ants of the present time, and still have had much time in which to build pyramids, temples, and the like. All this, if true, suggests that the present milpa system of agriculture, with only minor modifications, could have supported a very much greater population than is found today in the Yucatan peninsula—how much greater only careful long continued studies can determine. at effects longer and more frequent use of the land for maize growing would have on soil fertility is beyond the writer’s ability at guessing. Certainly soil erosion could have played no part in bringing about the abandonment of a low, flat region like that around Chichen Itza. There is no evidence whatever, so far as one who is not a geologist can determine, to indicate the presence of such “washes” as one sees commonly in the much drier regions, deserts, of California. But these regions are bordered by real mountains, not merely the low hills of Yucatan; and the steepness of their slopes is a surprise to any automobile driver who is likely to have his engine stall on what seems almost level ground. [Vi с. 40, 1953] 62 ANNALS OF THE MISSOURI BOTANICAL GARDEN The absence of erosion in northern and western Yucatan is no argument against the possibility that erosion was an important factor in causing the abandonment of the higher and more broken regions farther south where the old-empire Mayas flourished for a time only to abandon the area later and migrate to the low lands of Yucatan. Whether or not the erosion theory is the correct answer to the aban- donment of the territory of the old-empire Mayas, it is now serving a useful purpose as the goblin of the present erosion campaign in the United States. RECOMMENDATIONS The survey here reported was only a preliminary study. Although statements of milperos are perhaps correct, many of the figures given here are at best on y estimates and some of them are guesses, perhaps very poor guesses. At best a hurried survey could not be expected to do more than to indicate what the elements of the problem are and to suggest the kinds of accurate information needed. he writer is convinced that weed competition and the labor involved in weed control is a more important factor than soil exhaustion, and yet, the latter might enter to a greater or less degree, particularly if the land were used for maize cul- ture for relatively long periods and at relatively frequent intervals. The only way to get reliable information concerning the possible depletion of soil fertility by maize culture is to conduct, for several years, a carefully planned and carefully executed field-plot experiment. But this would require the services of a man trained in modern agronomic technique for at least part of each year, and of a trustworthy helper to keep detailed records during the growing season. But there are problems of perhaps more immediate importance which should be undertaken at once, if their solution is of enough importance archeologically to warrant the expense involved. And, for the solution of these problems also, it is of prime importance to have a well-trained man with a well-founded agronomic background. Such a study should provide accurate information to take the place of the hear-say evidence, the estimates, and the guesses presented in this report. It would seem sufficient to select six or eight sample areas, such as the territory contributory to Piste already studied in part by Dr. Steggerda. It is important to know the number of people gaining their support in a given area, the amount of maize produced and used for food by them and their domestic animals, or sold in order that other necessities may be purchased, the number of mecates cultivated in order to provide this maize, and the relation that the area of all the milpas bears to the entire area. Accurate information should be obtained also respecting the hours of labor per unit area devoted to each separate operation involved and in the preparation and care of milpas. If a field-plot experiment is undertaken, one series of plots should be kept free from weeds at whatever cost, while another series is cared for as the Mayas now tend their milpas. Such an experiment should afford information not only in respect to possible soil depletion, as measured by maize yields, but the labor records should be of value in appraising the possibility that the ancient Mayas may have kept the land in milpas a greater percentage of the time than is now done. ee eer STAFF .- OF THE MISSOURI BOTANICAL GARDEN Director GEORGE T. Moore Assistant Director EDGAR ANDERSON CARROLL W. DODGE, HUGH C. CUTLER, Mycologist Curator Museum of Economie Plants ROBERT E. WooDsoN, JR., Gong P- B. Van AN $снАлск, Curator of the Herbarium of neo N. ANDREWS, JULIAN A. нме botanist Honorary Research Associate Rorra M. Try Nett C. Horner, gat istant Gate ‘of Len Librarian and Editor erbarium of Publications —~ GrRALD ULRICH .. Business Manager | BOARD oF TRUSTEES ОЕ ТНЕ MISSOURI BOTANTAL GARDEN 5 bd | RICHARD J. Lock Vice-President Dant К, К. CATLIN Second Vice-President EucENE Perros . ^. оны S. dcs DE qe ` ETHAN H. SHEPLEY Er um : Romer Brookes: Sur ; о a EX-OFFICIO ‘MEMBERS 3 Автыча Н. Compton, _ : dH _5тадтово L. маек. Chancellor of Washington pe d kt “у = ЕЕ | THUR © LICHTENBERGER, | os Volume XL Number 2 Annals of the Missouri Botanical Garden MAY, 1953 А Contribution to the Tiden Flora of Arizona and New Mexico $i Emanuel D. ). Rudolph The Analysis of Suspected Hybrids, as s Illustrated by | Berberis X Sladwynensis. - . iens Anderson Variation in Cob Morphology among Certain Archaeo : Nort Et The Cytology, a and Systematic Relationships of e map cm p сарат Hort. ex Berg: om = = Marilyn Ашу age n dem and Ethnological Races of Маше . . - A ee 79-1 ш» issouri Botanical Garden. A e A С м L B r ic 1 Annals of the Missouri Botanical Garden Vol. 40 MAY, 1953 No. 2 A CONTRIBUTION TO THE LICHEN FLORA OF ARIZONA AND NEW MEXICO EMANUEL D. RUDOLPH* The lichen flora of the states of Arizona and New Mexico is much less known than that of many other sections of the United States. In addition to the references on distribution given in the standard works of Tuckerman (1882, 1888) and Fink (1935), there are only scattered papers in the literature which are concerned with the lichens of these states. Tuckerman, in 1860 and 1862, reported on the collec- tions made by Fendler near Santa Fe. When these records were brought together in 1866, several collections made by Wright in New Mexico were added. Tuckerman (1878) also wrote up some Arizona collections made by the U. 5. Geological Survey of the 100th Meridian. It was not until some time later that Fink (1909, 19092) studied the lichens of the vicinity of Tucson, Arizona, from where some new species were described by Zahlbruckner (1908,'09). In 1932 Bouly de Lesdain published an account of lichen collections from the vicinity of Las Vegas, New Mexico, made by Brother Arséne Brouard, and, in 1942, on further collections from the vicinity of Santa Fe. Magnusson (1929, 1937) treated the Acarospora of Brother Агзёпе Brouard. Herre (1944, 1950) has provided additional informa- tion on the lichen flora of New Mexico from San Miguel and Sierra counties. Finally, Darrow (1950) has reported on the arboreal lichen flora of southeastern Arizona. The present paper includes forms from some new localities in Arizona and New Mexico of which three are new species and a number are additions to the flora. This compilation is the result of a study of two collections: (1) that of Dr. Robert A. Darrow made in various localities in southeastern Arizona in 1933 and 1934, and in the 1940's which includes only saxicolous and terricolous forms; (2) that made by Mr. and Mrs. Francis Elmore at Chaco Canyon National Monu- ment in northwestern New Mexico in 1952. The latter is for the most part saxicolous, but a few corticolous forms are included. Thus most of these lichens are crustaceous rock-inhabiting ones which form a conspicuous element in the vegetation of these arid mountainous regions. * Missouri Botanical Garden. (63) [Vor. 40 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN The arrangement and nomenclature, wherever possible, follows that of Fink (1935). The treatment of Acarospora is in accord with that of Magnusson (1929a). А complete set of specimens is on deposit in the herbarium of the Missouri Botanical Garden. I should like to thank Dr. Carroll W. Dodge for his valuable guidance and assistance throughout this study. LIST OF SPECIES VERRUCARIA FUSCELLA (Turn.) Ach.—ARIZONA: Tucson, on caliche, 2500 ft., Dec. 4, 1933, Darrow 572; Tucson Mts., on volcanic rock, 3000 ft., March 11, 1934, Darrow 746. NEW MEXICO: Chaco Canyon, on sandstone, 6200 ft., Jan. 16 & 20, Feb. 28, 1952, Elmore. STAUROTHELE UMBRINA (Ach.) Tuck.—NEW MEXICO: Chaco Canyon, on sandstone, 6200 ft., Jan. 20, 1952, Elmore. DERMATOCARPON POLYPHYLLUM (Wulf.) Dalla Torre & Sarnth.—arizona: Pima Co., Rincon Mts., on shale, 4500 ft., March 4, 1934, Darrow 697. DERMATOCARPON RUPICOLA Zahlbr.—ARIZONA: Tucson Mts., on andesite, 3500 ft., March 15, 1934, Darrow боо. ENDOCARPON PUSILLUM Hedw.—ARIZONA: Tucson Mts., on caliche, 3000 ft., March 11, 1934, Darrow 745. RTHOPYRENIA HALODYTES (Nyl) Arn.—NEw MEXICO: Chaco Canyon, on sandstone, 6200 ft., Jan. 16, 1952, Elmore. PYRENULA NITIDA (Weig.) Ach.—Nrw MEXICO: Chaco Canyon, on grease- wood (Sarcobatus vermiculatus (Hook.) Torr.), 6200 ft., Jan. 20, 1952, Elmore. URCEOLARIA SCRUPOSA (Schreb.) Ach. var. BRYOPHILA (Ehrh.) Ach.— ARIZONA: Whitehouse Canyon, Santa Rita Mts., on Cladonia primary thalli, Dec. 24, 1933, Darrow 537. LEPTOGIUM APALACHENSE Nyl.—arizona: Pima Co., Rincon Mts., on shale, 4500 ft., March 4, 1934, Darrow 608. LEPTOGIUM BURGESSII (L.) Mont.—arizona: Whitehouse Canyon, Santa Rita Mts., on soil, Dec. 24, 1933, Darrow 533. PLACYNTHIUM MICROPHYLLIZUM (Nyl) Hasse—arizona: Tucson, on soil, 2500 ft., Nov. 26, 1933, Apr. 5, 1934, Darrow 527, 691; Pima Co., Cortaro, on soil, 3000 ft., Nov. 26, 1933, Darrow 528; Pima Co., Rincon Mts., on soil, 3500 ft., March 4, 1934, Darrow 740, 741. LECIDEA BRANDEGEI Tuck.—NEW MEXICO: Chaco Canyon, on sandstone, 6200 ft., Jan. 16, 1952, Elmore. LECIDEA PARASEMA Ach.—NEW MEXICO: Chaco Canyon, on sandstone, 6200 ft., Jan. 16, 1952, Elmore. LECIDEA PLANA (Lahm) Nyl.—new MEXICO: Chaco Canyon, on sandstone, 6200 ft., Feb. 28, 1952, Elmore. LECIDEA VORTICOSA (Floerke) Koerb.— NEw meExic о: Chaco Canyon, on sand- stone, 6200 ft., Feb. 28, 1952, Elmore. 1953] RUDOLPH—LICHENS ОЕ ARIZONA AND NEW MEXICO 65 RHIZOCARPON DISPORUM (Naeg.) Müll. Arg.—arizona: Mt. Lemmon, Santa Catalina Mts., on marble, 8500 ft., April 1, 1934, Darrow 721. ACAROSPORA AMERICANA Н. Magn.—ARIZONA: Tucson Mts., on volcanic rock, March 11, 1934, Darrow 747; NEW MEXICO: Chaco Canyon, on sandstone, 6200 ft., Feb. 28, 1952, Elmore. ACAROSPORA COLORADINA Н. Magn.—arizona: Tucson Mts., on volcanic rock, 3000 ft., March 11, 1934, Darrow 746. ACAROSPORA OXYTONA (Ach.) Mass.—NEw MEXICO: Chaco Canyon, on sand- stone, 6200 ft., Jan. 20, 1952, Elmore. ; ACAROSPORA PELTASTICA Zahlbr.—arizona: Tucson Mts., on volcanic rock, 3000 ft., March 11, 1934, Darrow 736; NEW MEXICO: Chaco Canyon, on sand- stone, 6200 ft., Jan. 20, 1952, Elmore. ACAROSPORA sTRIGATA (Nyl) Jatta—NrEw MExico: Chaco Canyon, on sand- stone, 6200 ft., Jan. 16 & 20, 1952, Elmore. ACAROSPORA TENEBRICA Н. Magn.—NEw MEXICO: Chaco Canyon, on sand- stone, 6200 ft., Feb. 28, 1952, Elmore. ACAROSPORA WASHINGTONENSIS Н. Magn.—ARIZONA: Рипа Co., Rincon Mts., on schist, 4500 ft., March 4, 1934, Darrow 766. PERTUSARIA FLAVICUNDA Tuck.—arIzoNa: Santa Rita Mts., on igneous rock, 7200 ft., Aug. 14, 1934, Darrow 820. LECANORA ATRA (Huds.) Ach.—arizona: Patagonia Mts., on granite, 5500 ft., Oct. 6, 1946, Darrow 4252. ECANORA CENISIA Ách.—ARIZONA: Santa Rita Mts., on igneous rock, 7000 ft., July 16, 1934, Darrow 796. LECANORA CONTORTA (Hoffm.) Stiz.—arizona: Mt. Lemmon, Santa Cata- lina Mts., on quartz, 8500 ft., April 1, 1934, Darrow 724. LECANORA DIFFRACTA Ach.—ARIzONA: Whitehouse Canyon, Santa Rita Mts., on granite, 6500 ft., Dec. 24, 1933, Darrow 470. LrEcANORA (ASPICILIA) elmorei E. Rud., var. nov.—TYPE: NEW MEXICO: Chaco Canyon, on sandstone, 6200 ft., Jan. 20, 1952, T. & Е. Elmore. Thallus determinatus, laevigatus, obscure olivaceus, areolatus, areolis crassis convexisque, 1-2 mm. diametro; cortex 31—41 p crassitudine, K—, fastigiatus, dimidia parte corticis crassitudine strato gelifacto tectus, hyphis circa 3 р diametro; algae ad Trebouxiam pertinentes, cellulis ad 38 p diametro, in strato non continuo, 100-170 р crassitudine sub cortice; medulla hyphis laxe contextis circa 4 p diametro. Apothecia in areolis crassioribus immersa, unum vel plura in quavis areola, amphithecio prominente, persistente, crasso, thallo concolore, disco concavo obscure olivaceo-alutaceo pulverulento; hypothecium indistinctum sed densum; thecium 150—171 р altitudine; paraphyses simplices, septatae, circa 2 p diametro, apicibus clavatis; asci 71.5-100.0 X 14.3-15.7 р, clavati, apicibus incrassatis; sporae uniseriate, 3—4-nae, sphaericae, 17.4—24.4 y diametro, granulosae, episporis tenuibus. Thallus determinate, smooth, deep olive, closely areolate, the areolae thick and convex, 1-2 mm. in diameter; cortex fastigiate, 31—41 p thick, covered by a 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN BPS | 988009. GP coto o OE. 0° | > p 2999 9 Кк 8528 оъ we Vd ا‎ T RA N AS itu E = —— Y; Ee eS < c Fig. 1. Lecanora elmorei E. Rud. 1. Section of thallus. ni Enlarged section showing cortex. w . Section of an apothecium. 4. Азсиз with associated paraphyses. [Vor.- 40 1953] RUDOLPH—LICHENS OF ARIZONA AND NEW MEXICO 67 gelatinous layer about half the thickness of the cortex, KOH—, the hyphae ap- proximately 3 р thick; algae Trebouxia, forming a layer 100—170 p thick below the cortex, broken into vertically rectangular patches separated by fungous hyphae, the cells globular to slightly angular, 10-38 р in diameter; medulla of loosely woven hyphae about 4 и in diameter. Apothecia sunken, the margin formed of the remaining part of the aerolae, one to several per areolae, the disc very concave, dark olive-buff, appearing powdery; margin persistent, raised and thick, thalline, colored like the thallus; epithecium of gelatinous tips of paraphyses; hypothecium indistinct, slightly denser than the underlying tissues; thecium 150—171 p thick, the paraphyses simple, filiform, septate, about 2 д thick, the tips swollen. Asci clavate, 71.5-100.0 X 14.3-15.7 и wide, hyaline tips present in many; spores uniseriate, 3—4 per ascus, globose, 17.4—24.4 и in diameter, mostly coarsely granu- lar, the epispores thin. This new species is easily distinguished by its 3—4 globose spores per ascus and by having Trebouxia as its alga. The globose-spored species Lecanora praecrenata Nyl, with its diffuse, whitish, indeterminate thallus and sessile brown apothecia, contrasts easily with the thick, closely areolate, deep-olive thallus and sunken apothecia of the present species. LECANORA EPULOTICA (Ach.) Leighton—arizona: Santa Rita Mts., on ig- neous rock, 6500 ft., July 1, 1934, Darrow 614. LECANORA FRUSTULOSA (Dicks.) Ach.—arizona: Carr Canyon, Huachuca Mts., on marble, 5600 ft., June 12, 1945, Darrow 4236. LECANORA MELAENA (Hedlung) Fink—arizona: Tucson Mts., on calcareous rock, March 25, 1934, Darrow 682. LECANORA PARISENSIS Му|.— МЕХ МЕХ!СО: Chaco Canyon, on greasewood (Sarcobatus vermiculatus (Hook.) Torr.), 6200 ft., Feb. 28, 1952, Elmore. LECANORA POLYTROPA (Ehrh.) Rabh.—arizona: Santa Rita Mts., Pima te a оп igneous rock, 4500 ft., Aug. 14, 1934, Darrow 810. LECANORA THAMNOPLACA Tuck.—ARIZONA: Santa Кіса Mts., on igneous rock, 7000 ft., Aug. 14, 1934, Darrow 822; NEW MEXICO: Chaco Canyon, on sandstone, 6200 ft., Jan. 16, 1952, Elmore. PARMELIA Novo-MEXICANA Gyeln.—NEw MEXICO: Chaco Canyon, on sand- stone, 6200 ft., Feb. 28, 1952, Elmore. CaLoPLAcA arizonica E. Rud. sp. nov.—TYPE: ARIZONA: Cortaro, оп rhyo- lite, 3000 ft., Nov. 26, 1933, В. A. Darrow 521. Thallus areolatus, fuscus; areolae 0.5—1.0 mm., dispersae; cortex circa 18 po fastigiatus, strato gelifacto tectus, K obscure purpurascens; algae protococcoideae ad 10 р diametro, cellulis sphaericis vel subangulosis, in strato 90—130 p crassitudine sub cortice; medulla hyphis laxe intertextis crystallis saxi impleta. Apothecia 0.2— 0.6 mm. diametro, orbiculata, disco ferrigineo-aurantiaco, amphithecio distincto, Persistente, tenui, thallo concolore; epithecium crystallis runneo-luteis inspersum; hypothecium hyalinum, centro circa 70 и altitudine; thecium 63—87 р altitudine; Paraphyses filiformes, 1.7 р diametro, simplices, septatae; asci 24—42 X 7-11 p, [Vor. 40 Xx e WES SS ee Se eT om APA y; НИ A AMET n a N| VEU 1 MN. ^ Те СА d и (и М ub KU OYE ә КОЖО ж d T 9 ak. -T - sem Fig. 2. Caloplaca arizonica E. Rud. 1. Section of thallus. Enlarged section showing cortex. Section of an apothecium. Ascus with associated paraphyses. “ ЖУ и Ascospores, 1953] RUDOLPH—LICHENS ОЕ ARIZONA AND NEW MEXICO 69 clavati; sporae 8-nae, 8.7—12.2 X 3.5-5.2 y, polari-biloculares, isthmo circa 3 p longitudine, uni- vel biseriales. Thallus indeterminate, aerolate, fuscous, the areolae 0.5—1.0 mm., often quite far apart; cortex about 18 y thick, with a gelatinous covering, fastigiate, KOH dark purple; algae protococcoid, forming a layer 90-130 p thick below the cortex, the cells round to angular, 5-10 и in diameter; medulla of loosely woven hyphae about 3 p thick, containing numerous small rock crystals. Apothecia circular, 0.2—0.6 mm., disc brownish-orange, the margin persistent, thin, of the same color as the thallus; algae in the margin and in a layer below the hypothecium; epithecium incrusted with brownish-yellow crystals, about 13 u thick; hypothecium hyaline, about 70 p thick at center; thecium 63—87 p high; paraphyses filiform, septate, about 1.7 р thick, unbranched. Asci clavate, 24—42 X 7-11 и; spores 8, uni- or biseriate, polar-bilocular, 8.7—12.2 X 3.5-5.2 p, the isthmus about 3 y long. This species somewhat resembles Caloplaca cinnabarina (Ach.) Zahlbr., but can be distinguished by its relatively fewer and more dispersed apothecia, darker thallus, and narrower spores. CALOPLACA CERINA (Ehrh.) T. Fries—NEw MEXICO: Chaco Canyon, on sand- stone, 6200 ft., Jan. 16, 1952, Elmore. CALOPLACA CINNABARINA (Ach.) Zahlbr.—arizona: Whitehouse Canyon, Santa Rita Mts., on granite, 6500 ft., Dec. 24, 1933, Darrow 471. CALOPLACA ELEGANS (Link) T. Fries—NEWw MEXICO: Chaco Canyon, оп sand- stone, 6200 ft., Jan. 16 & 20, Feb. 28, 1952, Elmore. CALOPLACA Festiva (Ach.) Zwackh.—arizona: Miller Canyon, Huachuca Mts., on igneous rock, 6300 ft., June 14, 1945, Darrow 4243. CALOPLACA FULGENS (Swartz) Koerb.—arizona: Nogales, on calcareous rock, 4000 ft., Oct. 6, 1946, Darrow 4261; NEW MEXICO: Chaco Canyon, on sandstone, 6200 ft., Feb. 28, 1952, Elmore. CALOPLACA LOBULATA (Floerke) Hellb.—arizona: Рипа Co., Rincon Mts., on granite, 4000 ft., March 4, 1934, Darrow 785. CALOPLACA MODESTA (Zahlbr.) Fink—arizona: Tucson Mts., on calcareous rock, 3000 ft., March 19, 1934, Darrow 730; Pima Co., Coyote Mts., on granite, 3200 ft., Feb. 16, 1945, Darrow 4266. TELOSCHISTES PARIETINUS (L.) Norm.—NEW MEXICO: Chaco Canyon, оп sandstone, 6200 ft., Jan. 16, 1952, Elmore. BUELLIA BLUMERI Zahlbr.—arizona: Whitehouse Canyon, Santa Rita Mts., on igneous rock, 6500 ft., Dec. 24, 1933, Darrow 477. BUELLIA RETROVERTENS Tuck.—aARizoNA: Whitehouse Canyon, Santa Rita Mts., on granite, 6500 ft., Dec. 24, 1933, Darrow 489; Tucson Mts., on volcanic rock, 3000 ft., March 19, 1934, Darrow 736; NEw MEXICO: Chaco Canyon, on sandstone, 6200 ft., Feb. 28, 1952, Elmore. Rinopina darrovii E. Rud. sp. nov.—TYPE: ARIZONA: Santa Catalina Mts., on ground and moss, Nov. 12, 1933, R. A. Darrow 498; PARATYPE: Santa Cata- lina Mts., on soil, 8000 ft., Nov. 12, 1933, Darrow 503. ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 3. Rinodina darrovii E. Rud. Section of thallus. — . ого» Enlarged section showing cortex. Section of an apothecium. Ascus with associated paraphyses. [Vor. 4 1953] RUDOLPH—LICHENS OF ARIZONA AND NEW MEXICO 71 Thallus ochraceus vel cinnabarinus, granulosus, K—, diffractus; cortex geli- factus, erodus; algae protococcoideae, cellulis sphaericis vel angulosis, 5—10 p diametro, in strato continuo 57-71 p crassitudine sub cortice; medulla densa, hyphis indistinctis circa 1.5 u diametro intertextis. Apothecia nigra, 0.5-1.0 mm. diametro, plana vel convexa; margine thalloideo, tenui, mox emarginato, thallo concolore; hypothecium hyalinum, in centro 80 џи altitudine; thecium 27-34 p altitudine; paraphyses 2 и diametro, septatae, simplices; asci 70-71 X 10-16 p, clavati; sporae 8-nae, biseriatae, 17.4—24.4 X 8.7-11.5 p, primo non-septatae, deinde tri-septatae, brunneae. Thallus ochraceous-tawny to cinnamon-brown, granulose, broken in places into irregular pieces, KOH-—; cortex gelatinous, eroded in parts; algae proto- coccoid, forming a continuous layer 57-71 р thick, the cells spherical to angular, 5—10 р in diameter; medulla of tightly woven, indistinct hyphae about 1.5 p in diameter. Apothecia black, 0.5—1.0 mm. in diameter; disc flat to convex; margin thalloid, thin, of same color as the thallus, soon disappearing; hypothecium hyaline, of closely woven hyphae, about 80 д thick at center; thecium 27—34 р high; paraphyses filiform, septate, about 2 p thick, swollen at tips and incrusted with yellow crystals, unbranched; asci clavate, 70—71 Ж 10—16 р; spores 8, biseriate, brown, at first 1-celled becoming 4-celled, 17.4—24.4 X 8.7-11.5 p This species has the general appearance of Rinodina оныг Dni, ) Vainio (R. nimbosa (E. Fr.) T. Fr.) but the four-celled spore places it in the subsection coNRADIA Malme as recognized by Zahlbruckner (1926). Its affinities seem to be with В. conradi Koerb. from which it can be separated by its larger black apothecia, its indistinct cortical hyphae, and its thicker hypothecium. The spores in this species have not been found to have more than four cells and also to be slightly smaller than those of R. conradi. ODINA EURYSPORA Zahlbr.—aAmizoNA: Santa Rita Mts., on igneous rock, 7000 ft., Aug. 14, 1934, Darrow 821. ODINA NOVOMEXICANA B. de Lesd.—ARIZONA: Santa Rita Mts., оп igneous rock, 4500 ft., June 5, 1934, Darrow 707; 7000 ft., Aug. 14, 1934, Darrow 823. BIBLIOGRAPHY — d iMac M (1932). topa de зи de New-Mexico (U.S.A.) recueillis par le Frére С. “т ids een У. na. Su im ent. da t Lichenol. 12:44—66. e nA . (1950). niim wd А fora of southeastern Arizona. Amer. Midl. Nat. Fink, Bı Mn EE и aati са Si The — ad of a desert кг -— y с "eti Distribution and movements of desert plants. Санга» Inst. Washington Publ изи. 3 EN The Lichen flora e the United States. Univ. ме an 426 he (1950). New Lichens from California, New Mexico and the Philippines. Bryologist 53:296-299, [Vor. 40, 1953] 72 ANNALS OF THE MISSOURI BOTANICAL GARDE кыл ча. AHO veo The Lichen genus Асатозротв in New Mexico. Meddelanden Gótesborgs tT. ved -72. Vien a ч Janoti of the genus Acarospora. Kungl, Svenska Vetenskapsakademiens $ Az , (193 ‘dona notes on ын и экы ЫА 1. Species from New Mexico. Meddelanden 2:87-9 7). Götesborgs Bot. Trad M Бар (1860). Den cial on North American and some other Lichenes, 1. Amer. cad. A 7007. з & Sci. Proc. 4 4 (1862) Part П. Ibid. 5:383—4 ‚ (1866). ie of California, cecus and the Rocky Mountains; as far as yet known. em › (18 25. | йай. In Rothrock's Report upon the botanical vore made in portions of Nevada; oni California, Colorado, New Mexico, and Arizona. Geol. Surv. W. 100d Meridian 6:3 > К ins) A synopsis of the North American lichens; Part I. 262 pp. Boston; art П. 176 pp. New Bedford, Zahlbruckner, Alerander (1808). ay меа American сакен Bull, Torr. Bot. Club 35:297-300. —————, (19 eue Flechte Mycol. 7:472-478. ————, а. Lichenes (Pacht): gue se Teil. ‘Ta Engler & Prantl’s Nat. Pflanzenfam. 8:61-270. THE ANALYSIS OF SUSPECTED HYBRIDS, AS ILLUSTRATED BY BERBERIS Х GLADWYNENSIS EDGAR ANDERSON During the last fifteen years a considerable portion of my time has been spent in attempting to measure the importance of hybridization in the evolution of natural populations. More than a score of genera have been studied intensively in the wild (and whenever feasible in the breeding plot as well) with students and associates, though as yet only a fraction of these studies has been published. Various techniques have been worked out for dealing with such problems (Ander- son, 1949, chap. 6), and as their validity has been more widely recognized (Stebbins, 1952) they are being more widely adopted. In the course of helping workers elsewhere to adapt these methods to their own problems I have become increasingly aware that in part they are based upon a scrutiny of the original material more exhaustive than is customary in many laboratories. It has accord- ingly seemed expedient to describe the procedure in more elementary detail than hitherto. Instead of discussing the complexities of hybridization in natural popu- lations I have chosen the relatively simple example of an apparent hybrid which arose spontaneously in the garden of Mrs. J. Norman Henry, of Gladwyne, Penn- sylvania. This seems a particularly happy choice since it also serves to demonstrate the usefulness of these methods in dealing with horticultural material of unknown ancestry. А vigorous young barberry was found coming up in the shelter of a large bush of Berberis verruculosa in Mrs. Henry's garden. Berberis verruculosa is a dense, evergreen-leaved plant, certainly the most distinctive of the hardy barberries. The seedling, so obviously related that it must have been a seedling of B. verruculosa, was nevertheless so different from it that Mrs. Henry had supposed it was prob- ably a spontaneous hybrid. It was different from any barberry known to her or to me and it exhibited the hybrid vigor which is the mark of so many hybrids. Furthermore, many garden hybrids of Berberis are known and Mrs. Henry at Various times had grown a number of other species of Berberis in her garden. With er permission a precise examination was made both of the putative hybrid and of Berberis verruculosa to establish the probable ancestry. The following is the procedure established in this laboratory for such examina- tions: l. Choose comparable material of parent and hybrid. 2. Examine, describe, and measure each feature item by item. Use great Care to work with truly comparable material. Do not attempt comparisons of a branch grown in the shade from one specimen and one grown in the sun from the other. If one is from a fruiting branch, then the other should be also. Use great саге in distinguishing between long shoots and short shoots. Many plants without such conspicuously heterogeneous shoot systems as the Ginkgo have a more or less well-defined short shoot-long shoot system which requires careful examination to Perceive, (73) [Vor. 40 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN INTERNODE LENGTH S № о А e = n L T T T , 3 4 5 6 1 '8 9 NUMBER OF SUCCESSIVE INTERNODES - Fig. 1. Internode diagrams of comparable branchlets of two species of barberry and their putative hybrid. Solid line, Berberis verruculosa; long dashes, Berberis X gladwynensis; short dashes, Berberi апае. Аз explained in Anderson А made by plotting the lengths of successive internodes from the base to the tip an ^ i i im гу line. The first internode at the base of the branchlet in В. Julianae, tc. e o Е 5 e = 5 > =. 8 ts wi for instance, is 45 mm. long, the second is 28 mm., the third, 33 mm., e 3. (And, most important): Conduct the entire examination, if possible, against a plain neutral background. Cover the table top with clean wrapping Paper or some such material. Remove all distracting objects such as pencils, paper clips, erasers, etc., out of the immediate field of vision. 'The eye can work much more efficiently in the perception of resemblances and differences if it can con- centrate upon the problem in hand. Work in a good light and have a good dis- 1953] ANDERSON —HYBRID ANALYSIS IN BERBERIS 75 cussion, and yet in assisting other people to use my methods I nearly always find that these precautions have not been followed and that it is difficult to convince other scholars of their supreme importance. This is one of those ridiculously simple matters which are far more important than they seem and which are indicative of true scientific precision. Specific differences are frequently subtle and they can be apprehended much more readily if one uses comparable material, a neutral background, and a good light. 4. Make measurements with good steel calipers and a steel metric rule. 5. Pay particular attention to pubescence, pubescence pattern, internode pattern (Anderson and Schregardus, 1944), branching pattern. Careful analysis will show that nearly all closely related species of the higher plants differ sig- nificantly (and with enough study, definably) in their internode patterns. Comparable branches were accordingly chosen from Berberis verruculosa and the seedling. Both were selected from well-developed branches of flowering age, growing in almost full sun and borne upon the upper parts of their respective. plants. After these two branches had been examined carefully against a neutral background, comparable branchlets were chosen for further study as follows: Proceeding from the tip, the first strong secondary branch (all of the current season's growth) was selected for study. The internode lengths of this branchlet were measured to the nearest millimeter, the results being plotted as an internode diagram (fig. 1). Hybrids are usually intermediate between their two parents when due allow- ances are made for heterosis, for growth-pattern differences, and for possible dif- ferences in ploidy. From a study of fig. 1, it is possible to make several predictions as to the putative male parent of the seedling barberry. B. verruculosa is shown to have more and shorter internodes than the seedling. Its basal internode is only about twice as long as the other internodes and the remaining internodes are sub- equal. The seedling has a basal internode more than twice as long as the others and the remaining internodes are not at all uniform in length and seem to have a definite pattern of decreasing length toward the tip. We therefore predict that the putative parent (when comparable material is studied) should have conspicu- ously non-uniform internode lengths with a very long internode at the base of such branches and decreasingly short ones as the tip is approached, probably in a well-defined pattern. a Leaf comparisons were then made. Choosing comparable leaves proved to be difficult but not impossible. Leaf size and shape were uniform in B. verruculosa but variable in the seedling. Care had to be taken, therefore, to choose exactly comparable material. Most of the primary leaves in these barberries are spines. The leafy leaves are borne on short secondary branches in the axils of the spine- leaves, giving the appearance of little clusters up and down the branches. In B. verruculosa all the leaves on the plant were superficially similar in size, shape, and serration. In the seedling they differed markedly. Those in clusters arising from the older wood were variable. Among them were leaves which were narrower ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vor. 40 PORE M 6.5 ; | 5.0 i | H | i | 5.5, I | : | 5.0] I j CF = Н = } Н / .0 H : 4.0] / “= Н Ё 3 H ; ^ 3.5. ; д Н / Н / |! / ^l Ly Н i № H H 2.5]; Н 2.0 3 ` 1.0 1.1 ; 1.3 ecu LEAF WIDTH CM. Relationships of size, shape, an of marginal setae in leaves from young and ies of estu their duin. hybrid in the previous figure, 2 hes, Ber x , h, Fig. 2. mature spurs for two spec solid line indi : young to ad. Ai rs hang matic, e. Th { size and shape are all to the same scale Perge represent ы values beris X gla | and seta number of e lines ien indicates the change pe and seta number Berberis ана ны es conspicuously i in size, $ hybrid is intermediate in degree of ; their ed t to the diagrams, while highly diga from actual measurem 1953] ANDERSON—HYBRID ANALYSIS IN BERBERIS 77 in shape, larger in size and with more strongly developed setae at the tips of the serrations along their edges. Careful examination then showed that B. verruculosa had a very slight tendency in the same direction though not strong enough to be readily apparent without examination. Measurements showed that while Berberis verruculosa and the seedling had about the same number of setae per leaf on the new wood, the seedling had nearly twice as many as verruculosa on the old wood. Figure 2 shows all these relationships. The putative parent for which we are searching should have leaves on its new wood as small as those of verruculosa, or even smaller, with greatly differentiated leaves on the older wood. Some of the leaves on the older wood should be very long, proportionately narrow, and with many setae along their edges. As, in this fashion, one proceeds feature by feature in the careful comparison of the one parent with the hybrid, he becomes progressively better acquainted with the general ground plan of the plant and is increasingly capable of making precise comparisons and valid inferences. Before long it is possible to draw up precise descriptions not only of B. verruculosa and its hybrid seedling but also of the putative male parent. Such a comparison is presented below in tabular form, including the predictions as to the male parent. verruculosa glad wynensis unknown (Julianae) Branches arching, Branches sub-arching, Branches straight, sub-horizontal some nearly erect erect to horizontal боса, short, Internodes medium, Internodes long, uni variable M Branchlets densely Branchlets E most Branchlets eglandular ew dular sub-glan ew growth scarcely New growth ча брон strongly Prem ated differentiated (bo iated Mature bark Mature Gá Mature Pan strongly barely ribbed definitely ribbed ribbed and furrowed Wood greenish Wood yellowish-green Wood bright yellow-green Spines up to 1.2 cm Spines up to 1.3 . Spines up to 1.5 cm. Leaves glossy Leaves somewhat glossy, Leaves not glossy, dark green above, -dark above, bright green above, glaucous bel sub-glaucous below light green bel Fall color a more or Fall color stronger on Brilliant fall color less general purple newest growth, occasional eaves о and d growth reels dior: ted Largest send on old spurs 7 mm. X 12 mm. 10 marginal setae at most Flowers borne singly in pairs Stigma sessile розаны н кы RN, Se DRIN T RING "o owth definitely аа Largest leaves on old spurs 43 mm. X 14 mm 20 ае setae at most Flowers in fascicles of 5 to 8 Stigma sub-sessile Leaves > new and old growt differen ciated Largest lave on ан spurs 65 mm. X 11 mm Over 30 ыа, setae Flowers in fascicles of 15 to 25 Stigma definitely not sessile, style of at least 1 mm. [Vor. 40, 1953] 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN As the description began to resolve itself my knowledge of barberries was suf- cient to suggest that Berberis Julianae was probably the barberry we were looking or. As soon as I made this suggestion Mrs. Henry informed me that a large bush of this species had once stood just a few feet away from B. verruculosa but it had been damaged in a windstorm and removed. Many of the technical points in the description, however, involved matters which were completely outside my knowl- edge of barberries. Predictions as to the numbers of setae on the mature leaves, the number of flowers per fascicle, the presence of a style, etc., were drawn up with no knowledge of what they might be like in B. Julianae. The entire hypothetical description was then run down in Rehder's key (Rehder, 1940) to the barberries just as if one had the specimen actually in his hand. It led to B. Julianae and fitted the description of that species precisely, down to the most technical detail. There can be little doubt, therefore, that the seedling was a spontaneous hybrid, B. verruculosa X B. Julianae. I am accordingly describing it as: BERBERIS X gladwynensis hyb. nov. Intermediate between its parents B. verruculosa and B. Julianae. Flowers in fascicles of 5 to 8, stigma subsessile, leaves subglaucous below, nearly evergreen. Type: Henry, in the herbarium of the Philadelphia Academy of Natural Sciences; from the type bush, E. Anderson 5. 9., It is a ens to name this handsome barberry after the site of Mrs. Henry's garden which has long been a mecca to botanists and gardeners alike. LITERATURE CITED Anderson, е рн у: MSS aene Hybridization. = Wiley & Sons, Inc. New ا‎ d ork. پا‎ — 3 1944). : os thod for recording and analyzing ай of inter- node t. Gard. а Rehder, А "a 1940). ЫТ? кө of "Cultiv vat id pus | Shrubs, The Macmillan Co. York. New Stebbins, G. Г. (1952). The evolution of совет plants and weeds. рае 6:445—448. 1 BERBERIS быку nsis hyb. nov. inter parentes В culos, Flores in fasciculis 5-8, stigmate subsessile. Folia fere = » E Јанета, verruculosam semperviridia, subter subglauc VARIATION IN СОВ MORPHOLOGY AMONG CERTAIN ARCHAEOLOGICAL AND ETHNOLOGICAL RACES OF MAIZE NORTON Н. NICKERSON* INTRODUCTION Zea Mays L., one of the most highly evolved of all grasses, is still botanically much of an enigma. Commonly known as Indian corn or maize, it was a very important plant to the peoples of the New World long before the arrival of Columbus. It formed the basis of the highly developed Inca, Maya, and Aztec civilizations and was the staple crop from Canada to Chile for several thousand years. It is, of course, quite important to the present-day inhabitants of Central and South America, and agriculture and industry in the United States uses three and one-quarter billion bushels annually. Botanists have until recently followed almost exclusively Sturtevant's (1899) classification of contemporary races of maize by kernel types. Although this classification is an artificial one, it has proven to be of practical commercial worth. Archaeologists have long known the value of maize kernels found at excavation sites for determining types of maize grown and the uses to which they were put. Until the last few years, other parts of the maize plant have been quite generally neglected as a source of historical data, chiefly because of the complexities involved in determining and evaluating such evidence as they contain. Evidence from maize tassels has been used with promising results (Alava, 1952; Anderson, 1944b, 1944c, 1949a, 1951; Anderson and Brown, 1948; Anderson and Cutler, 1942; Brown ef al, 1952; Cutler, 1946; Wellhausen e£ al, 1951, 1952). Prat (1948) showed that in maize and other grasses hairs and other epidermal emergences can be used as a basis for identification and classification. Internode patterns have been studied by several workers (Anderson, 1943, 1949а; Anderson and Brown, 1948; Anderson and Schregardus, 1944; Stonor and Anderson, 1949; Wellhausen e£ al, 1951, 1952). Ear and tassel ontogeny have been studied by Bonnett (1940, 1948) and Kiesselbach (1949). Esau studied the ontogeny of the maize vascular bundle. Surprisingly little has been done to measure and evaluate those morphological structures which are present on a maize cob after the kernels have been removed. Weatherwax, a pioneer in the study of the maize cob, pointed out (1918) the need and value of accurate morphology in understanding this structure, and dem- onstrated (1920) the orderly spikelet behavior underlying changes in kernel row- ing. Fujita (1939) observed that an even number of pairs of kernels resulted in straight rows, and an odd number in spiral rows. Cutler (unpublished work) made many histological studies of the cobs of both North and South American maize, as well as of several closely related genera. Lenz (1948) indicated the types eer "Special Assistant in Botany, Henry Shaw School of Botany, Washington University. The stimulating advice of Dr. Edgar Anderson and the technical erai of Mr. Ding Hou throughout the course of this investigation are hereby gratefully acknowledge [Vor. 40 80 ANNALS OF THE MISSOURI BOTANICAL GARDEN of evidence available from a histological study of the maize cob. Alava (unpub- lished work) modeled in great detail small portions of the exterior surface of a maize cob. Mangelsdorf and Smith (1949) tabulated a number of external and internal maize ear characters, and their general procedure was used in character- izing 25 present-day races of maize in Mexico by Wellhausen e£ al (1951, 1952). These studies were based on the evaluation of many characters which did not in- volve exact measurements. In the present study, the value of several morphological characters used by these investigators has been increased by employing exact measurements. Other measureable morphological characters have been which are of significance in tracing historical influences of one race of maize upon another. : There are two reasons why evidence present on the maize cob is important. The first one is based on the generally accepted fact that distinguishing characters are not distributed at random among plants; with regard to their occurrence in the Maydeae, agrostologists have found the female inflorescence to be of particular significance. The second reason, already pointed out by Lenz (1948), is that there are more archaeological remains of maize cobs than of any other plant material. The importance of these remains can perhaps be appreciated when one considers that maize, unlike most cultivated crops, is completely dependent upon man for its preservation. This fact can only mean that it has had a constant associatio with man since its adoption as a food plant, and that in its manner of origin, in the course of its migrations, in its development into a myriad of races, and in its intimate association with ancient religious symbolism, the history of maize becomes intimately tied to the history of man. Increased insight into the one will certainly add to our present understanding of the other. For a reasonably complete investigation of the maize cob, the following three courses were deemed necessary: (1) to measure certain easily-recognized features of the maize cob and to interpret their morphological nature; (2) to set forth one possible scheme of analysis of the variation patterns of different races of maize based on these measurements; (3) to discuss the resulting indicated trends as to their validity and applicability in extending our present understanding of the history of maize. MORPHOLOGY oF THE EAR Bearing in mind that an adequate treatment of variation in maize ears would be more complete if the nature of those parts under discussion were understood, 2 detailed study of morphological structures found on the maize ear was first under- taken. However, since the results are of a technical nature quite different from those enum from a study of variation, they are treated elsewhere (Nickerson, in press). 1953] NICKERSON—VARIATION IN СОВ MORPHOLOGY 81 MORPHOLOGICAL STRUCTURES EXAMINED One of the purposes of this study in variation is to determine what morpho- logical structures expressed in the cob are of significance in differentiating between races of maize. To accomplish this, several characters had to be included which ultimately proved to be of little use. A total of 526 cobs was examined for each of the three external and five internal characters discussed below. EXTERNAL CHARACTERS Shape of Ear.—This character is the only one used in the investigations which is dependent upon a subjective grading. Ears were classified into one of four types: straight, cigar-shaped, tapered, enlarged butt. Two of these types—straight and tapered— were used in analysis of each sample. For reasons discussed else- where (Nickerson, in press), a straight-eared race exhibits less condensation than a tapered- or enlarged butt-eared race, and a race with cigar-shaped ears is inter- mediate between these extremes. Shank Diameter.—Yhe diameter of the shank was measured in millimeters at a point close to the base of the ear, above the last apparent husk node whenever possible. In specimens with elliptical shanks, two measurements were made and an average of these was used. Shank diameters are fairly consistent within any one kind of maize and are often markedly divergent between different kinds. Row Number.—Row number was determined by counting the number of vertical rows of glumes in the middle area of the cob. The middle area was used for this and all subsequent measurements because of its uniformity of size, stage of development, and freedom from growth irregularities common at both base and tip of the ear. Row number is a readily-observed character the importance of which is not yet fully understood. Anderson and Brown (1948) were able to correlate the condensation index of the tassel with kernel row number in the ear. Cutler (1952) used row number in his preliminary study of cobs from successive layers of cultural remains found in Tularosa Cave. His tables indicated the pres- ence of high row numbers in lower strata and progressively lower row numbers in successively higher strata. These data show the reverse of the situation reported for the stratified remains of Bat Cave by Mangelsdorf and Smith (1949), who stated that in general the older (and more pod-corn-like) the ear, the lower was its row number. INTERNAL CHARACTERS To study internal characters, the ear was broken in two at the approximate middle; the exposed ends were then observed under a dissecting microscope. Width of Lower Glume.—This character was measured in millimeters with the calipers held perpendicular to the widest point on the abaxial surface of the glume. measurement is not a direct expression of a particular gene such as Mangels- dorf and Smith (1949) reported for the relationship between lower glume length and various alleles of the Tu gene, but reflects the width of the basal portion of the kernel which it encloses. [Vor. 40 82 ANNALS OF THE MISSOURI BOTANICAL GARDEN Height of Rachis-flaps——This measurement in millimeters was made perpen- dicular to the axis of the cob. In instances where the rachis-flaps were of unequal height, an average measurement was used. The rachis-flap is a part of the cupule (Nickerson, in press) and its significance is somewhat minimized by its variability I in some otherwse homogeneous samples used in the present investigation. Mangels- dorf and Smith (1949) reported the same situation for maize cobs from Bat Cave, but Wellhausen e? al (1951, 1952) found rachis-flaps to be extremely useful in characterizing races of maize in Mexico. Deptb of Cupule.—This measurement in millimeters was made perpendicular to the cob axis, and was given a positive or a negative value, depending upon whether it was out from, or down into the rachis. It represents the extent of adnation of the cupule-forming prophyll (Nickerson, in presa) to the axis, and is well correlated with ear shape. Width of Cupule—This measurement was made by holding the calipers at right angles to the cob surface and measuring the distance in millimeters between iN See ри MN dnm A A NORTHERN FLINT 1 HUACA PRIETA HEIGHT OF RACHIS . FLAPS N DEPTH OF CUPULE - POSITIVE 3 DEPTH OF CUPULE - NEGATIVE > WIDTH OF CUPULE 5 KERNEL THICKNESS : Text-fig. 1. Drawings of two representative maize races ques distances in deter- mining four internal cob characters. А, transverse section of c have bes ч ch a transverse cut was m. i i n ings are to scale; arrows at 2 and 3 are each 1953] NICKERSON—VARIATION IN COB MORPHOLOGY 83 any pair of outside rachis-flap edges across their cupule. It is in part an indication of kernel width, in that wide cupules always mean wide kernels. Thickness of Kernel.—The distance along the cob axis from the base of one set of lower glumes to the upper edge of their cupule is exactly equivalent to the thickness of each kernel produced there. The measurement is useful in making comparisons between productivity of different cob samples, especially those in- volving fragmentary archaeological remains. These last four characters were recorded in outline drawings carefully made to scale (working with calipers under a dissecting microscope) for each of the 526 cobs examined (text-fig. 1). Other characters measured but not used include height and texture of the lower glume and longitudinal profile drawings of each cupule. Diameters of the pith, cob, and rachis were measured and the cob/ rachis index (Mangelsdorf and Smith, 1949), as well as the rachis/pith index, was com- puted for all specimens. However, neither the averages nor the individual vari- ations of these indices were found to be particularly characteristic among the maize samples studied in this investigation. NAMES, SOURCES, AND ANTHROPOLOGICAL BACKGROUND OF SAMPLES Maize samples examined were of diverse origin both geographically and temporally. Races of contemporary maize here included were chosen as typical for those parts of the world. Archaeological cob samples were largely from the southwestern United States, but included one sample from Lower California and two extensive collections from sites excavated by Junius Bird in Peru and northern Chile. In listing the samples they are grouped arbitrarily into geographical units; no further inferences should be drawn from this arrangement. Samples, except where otherwise noted, are from the maize collection of the Museum of Economic Botany of the Missouri Botanical Garden, St. Louis. NORTH AMERICA PRESENT-DAY RACES: Iroquois Sacred Flour.—This sample was from material originally grown by Professor Frank P. Bussell from seed collected by Erl Bates from the Iroquois Indian tribes of northern New York. The Indians grew it under conditions of Strict isolation for ceremonial use (Anderson, 1947b). Ears are straight, 20—28 cm. long, with heavy shanks and 8 rows of wide white or yellow (sometimes pale blue) kernels. Fourteen ears were examined. Northern Flint.—A composite sample, this group included flint corns from the Northeast (Parker's, Canada, Mammoth Yellow, Longfellow) and the - Great Plains (Winnebago, Bear Island, Brownell, Mandan, Tama, Golden Chip- Pewa). Brown and Anderson (1947) studied these and other representative flint Varieties and reported that the ears were long and slender with 8—10 rows of wide, crescent-shaped kernels, heavy shanks, and a tendency toward a large cob base. wenty-three ears were examin [Vor. 40 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN Hopi White Flour.—This sample was furnished by Dr. William L. Brown, Pioneer Hi-Bred Corn Co., Johnston, Iowa, who collected it at the Hopi town of Hotevilla, Arizona. Carter and Anderson (1945) considered Hopi White Flour to be an example of Puebloan maize. Brown еЁ al (1952) reported that this variety is one of their main sources of meal. The straight ears are 18—24 cm. long, with wide shanks, no basal compression, and wide white kernels in 12—14 rows. Seven ears were examined. Hopi Blue Flour.—Also furnished by Dr. William L. Brown, this collection has substantially the same history as Hopi White Flour. However, Brown ef al (1952) reported that it is probably more important as a source of food. e cigar-shaped ears are 14—20 cm. long, with small shanks, a tendency toward basal compression, and 10—20 rows of narrow blue-aleuroned kernels with white endo- sperm. Eleven ears were examined. : Papago Flour.—This race of maize is grown by the desert-dwelling Papago Indians, and was reported by Carter and Anderson (1945) to be quite similar to prehistoric Basketmaker maize. 'The ground kernels make a meal of excellent quality. Ears are 20-25 cm. long, cigar-shaped, with narrow shanks and 12-14 rows of isodiametric yellow kernels with white endosperm. Twenty-five ears were examined. ARCHAEOLOGICAL REMAINS: Basketmaker.—Cobs included in this collection were recovered from Kinboko and White Dog Caves, both of which are located in Tsegi Canyon, Marsh Pass, northeastern Arizona. The samples were submitted by Mr. Harold S. Gladwin, who estimated their age to be circa 200-300 A.D. On the basis of there being no pottery associated with these cobs, they are believed to belong to the cultural level known as Basketmaker П. The cigar-shaped cobs are 6—12 cm. long, and bore somewhat isodiametric kernels in an average row number of 14. Six cobs were examined. Marsh Pass.—Cobs included in this collection were recovered in West Hack- berry Canyon and Turkey Cave, both of which are located in Tsegi Canyon, Marsh Pass, northeastern Arizona. These samples were also submitted by Mr. Н. S. Gladwin, who estimated their age to be from 300 A.D. to about 1000 A.D. The collection represents cultural levels from late Basketmaker П or later up through Pueblo П. The predominantly cigar-shaped cobs are 6—14 cm. long, and bore kernels wider than thick in an average row number of 10. Thirty-one cobs were examined. Turner Site.—This sample consisted of 50 cobs and fragments recovered from the Turner Site, near Cisco, Utah, by H. M. Wormington, Curator of Archaeology, Denver Museum of Natural History. Wormington estimates its age to be around 1000 A.D. (personal communication, 1953), and believes that the site was in- habited by peoples who were culturally a later manifestation of the Fremont Basketmakers. Remains of their culture were recovered from Castle Park in the 1953] NICKERSON— VARIATION IN COB MORPHOLOGY 85 Yampa River Valley, Utah, and are dated at 400-800 A.D. by Burgh and Scoggin (1948). There are nearly equal numbers of tapered and cigar-shaped cobs in this collection; some were not used in this study because of their fragmentary nature. Cobs are short, 3—8 cm. long, charred, with medium shank diameters and medium- wide kernels in an average number of 12 rows. Twenty-seven cobs were examined. Luster Cave.—Ten cobs were recovered by Dr. Robert Lister, Department of Anthropology, University of Colorado, Boulder, from a cave located in Utah, just west of the Colorado state line, in the Glade Park area. A preliminary report (Lister and Dick, 1952) described the archaeological finds of this cave and of other nearby sites. On the basis of pottery types, the maximum possible age assigned to Luster Cave by these workers is 900-1000 A.D., but it may be much more recent. The straight and tapered cobs are generally uncharred and fairly complete with small shank diameters and wide kernels in an average number of 10 rows. Nine cobs were examined. Tularosa Cave.—More than 30,000 maize cobs were recovered from this site in Pine Lawn Valley, northeast New Mexico, by archaeologists from the Chicago Natural History Museum. The Museum has submitted these cobs to Dr. Hugh C. Cutler, who has generously made available for inclusion in the present study samples representing three levels of Square 3R2. Maize cobs from the lowest levels of Tularosa Cave were dated at 2300 + 200 years by the Carbon 14 method. The cave is estimated to have been abandoned somewhere around 1000—1200 A.D., and thus was continuously occupied for about 1500 + 500 years (Martin ef al, 1952). Material from Square 3R2, Levels 3, 6, and 11 was studied as three separate sam- ples, each of which was chosen so that the cobs analyzed reflected the proportionate row numbers determined by Cutler (1952) from all cobs found in each of these layers. "There was a decrease in row number from the lower levels to the surface, average row numbers for Level 11 being 12—14, for Level 6, 10—12, and for Level 3, 8-10. Culturally, Level 11 is the oldest (400 В.С. + 200 years) and represents the Pre-pottery Phase. Level 6 dates back to 500—600 A.D., and represents the Georgetown Phase. Level 3, at 1000—1200 A.D., represents the San Francisco Phase. Twenty-five cobs from each of the three levels were examined. Point of Pines.—This sample consists of about 250 ears of charred maize. It Was recovered from Room 50 of a large ruin at Point of Pines, Arizona, numbered Ariz. W:10:50, and was submitted by Dr. Emil W. Haury, Director, Arizona State Museum, Tucson, Arizona. He believes its age to be 1250 A.D. (personal communication, 1953). It can be assigned to the Pueblo II cultural phase. The straight and cigar-shaped ears are 5-8 cm. in length, with small shanks and smooth kernels wider than thick in an average number of 10 rows. Seventy-five cobs Were examined. [Vor. 40 86 ANNALS OF THE MISSOURI BOTANICAL GARDEN MEXICO AND CENTRAL AMERICA PRESENT-DAY RACEs: Chapalote——This race of popcorn is considered by Wellhausen её al (1951, 1952) to be one of the ancient indigenous races of Mexico. It was mentioned by Anderson (1944a, 1946) as being allied to the primitive Maíz reventador. Well- hausen e£ al pointed out its resemblance to archaeological maize finds at Painted Cave (Haury, 1945) and Cottonwood Cave (Hurst, 1948; Hurst and Anderson, 1949). They considered it to be one of the most distinctive races of maize in Mexico. Mangelsdorf (1948) and Mangelsdorf and Smith (1949) regarded it as possessing the "weak" allele for pod corn. The cigar-shaped ears are 10—15 cm. long, with small shanks and smooth rounded chocolate-brown kernels in an average row number of 12. Ten ears were examined. Guatemala Flint.—A common variety of the Guatemala highlands, this race was described by Anderson (1947a), who considered it to represent one of the basic elements in Guatemalan maize. Ears are 10—25 cm. long, with a conspicu- ously enlarged and irregularly rowed butt, heavy shanks, a heavily sclerenchyma- tized rachis, and wide flinty kernels (some of which are capped with soft starch) in 8 (occasionally 10 or 12) straight rows. Eleven ears were examined. ARCHAEOLOGICAL REMAINS: Lower California.—This sample of 52 maize remains was recovered from Cave B.C. 100, located 8-10 miles east-southeast of Comondu, in central Baja Cali- fornia, and was submitted by Dr. William С. Massey, Dept. of Anthropology, University of Washington, Seattle. The cobs were recovered from a layer dated by Dr. Massey at 1697—1750 A.D. (personal communication, 1953). The cigar- shaped and straight cobs are 6-12 cm. long, with small shanks, and bore nearly isodiametric kernels set in 10-12 rows. Thirty-five cobs were examined. SOUTH AMERICA PRESENT-DAY RACES: Peru Flour.—This sample was collected by G. Edward Nicholson at Huancayo, Peru, where it was being offered for sale in one of the native markets under the name of "Maíz de Color.” It belongs to the race Cutler (1946) called “Valle Maize.” Ears are short, tapering, constricted at the base, with small shanks. The race exhibits little sclerenchymatization. Kernel color varies from ear to eat; brown, red, yellow, and various delicate striped combinations are most common. Kernels are pointed, as wide as thick, and arranged in prominent rows averaging 10 per ear. Twenty-three ears were examined. Coroico.—This race, described by Cutler (1946) as "the most unusual race so far known," has the odd characteristic of brick-like arrangement of “‘alicoles” (cupule plus pair of associated spikelets; Nickerson, in press). Cutler described the tapered ears as long, slender, and flexible (25-30 cm. lengths were common in the collection examined), with a light brown cob, small pith, and brown-orange 1953] NICKERSON—VARIATION IN COB MORPHOLOGY 87 kernels. He stated that the row number averaged 9, but he now believes (personal interview, 1953) that two successive rings of alicoles constitute a single diametral whorl; thus the row number of Coroico is generally high, averaging 18. Whether 9 or 18 rows are present is governed by the state of condensation of the cob axis. Twenty-five ears were examined. ARCHAEOLOGICAL REMAINS: Arica.—These samples were recovered from two midden sites, Playa Miller and Quiani, located on the coast in the vicinity of Arica, Chile (18° 30’ S. Latitude), by Junius B. Bird, Associate Curator of Archaeology, American Museum of Natural History, New York. Cobs from both sites were treated as one sample following Bird’s (1943, 1948) suggestion that Playa Miller represented a continu- ation of sequences started at Quiani. The oldest maize remains from this area are dated at about the time of Christ (Bird, personal communication, 1952). It is thus younger than material from the Huaca Prieta sites discussed below. The cigar-shaped and tapering cobs are 5-12 cm. in length and 1—2.5 cm. in width, with small shanks and an average row number of 14. Twenty-one cobs were examined. Huaca Prieta.—These samples, also from the American Museum, were recovered by Junius B. Bird from Huaca Prieta, a site on the Peruvian coast at the south of the Chicama Valley (app. 8° $. Latitude). These comprehensive samples consist of 604 cobs from Test 1 and 171 cobs from Test 5. The two groups were treated separately, since Bird regarded cobs from Test 5 as older than those from Test 1, and were designated Huaca Prieta 5 and Huaca Prieta 1, respectively. Cultures responsible for the Huaca Prieta middens began about 3000 B.C. (Bennett, 1948; Bird, 1948), but maize did not make its appearance there until 850 B.C. (Bird, personal communication, 1953). Implications as to archaeological and botanical Significance of these finds are further discussed by Anderson (1947b), Carter (1950), and Whitaker and Bird (1949). The well-preserved, tapered and cigar- shaped cobs are remarkably uniform, especially those found in the lower levels, which have small shank diameters, an average row number of 14, and are horny rather than brittle or bony in texture. Thirty-five cobs from Huaca Prieta 1, representing 7 layers, and 44 cobs from Huaca Prieta 5, representing 5 layers, were examined. ASIA Assam and Lushai.—The samples from Asia were contributed by Stonor, who collected them in the hills of Assam. The history and morphology of varieties in this collection have been described by Stonor and Anderson (1949). The cobs used here were those actually raised in Asia. For this investigation they were Separated into two groups: (1) Assam, which includes 21 cobs from tribes other than the Lushai; (2) Lushai, which includes 8 cobs representative of the maize Varieties grown by the Lushai tribe. Cobs of both groups are sufficiently alike to TABLE I AVERAGE VALUES OF NINE ca amm CHARACTERS AND TWO INDICES FOR EACH MAIZE COB SAMPLE STUDIED (M urements in mm. except where otherwise indicated % cobs Height of Kernel % cobs | Lower gl. | Cob/rachis | Rachis/pith Name at Sanpla straight | rachis flaps | thickness | tapered width index index Iroquois Sacred Flour 43 2.0 3.7 43 8.7 1.7 2.1 Northern Flin 35 2.0 4.4 32 7.0 1.8 2.1 Hopi White Flour 35 2.0 4.7 65 6.2 17 1.8 Hopi Blue 35 1.6 4.2 0 5.8 1.9 1.8 Papago Flou 0 1.2 4.0 28 5.4 1.8 1.9 Basketmaker 9 0 1.5 4.3 16 4.4 1.6 2.0 Marsh Р 1 16 1.5 4.1 20 5.5 1.5 2.1 Turner Site 8 12 1.0 3.0 50 3.7 1.3 1.7 Luster Cave = 33 1.6 4.5 33 3.4 1.8 1.8 Tularosa Cave—L. 3 9 0 1.5 4.1 4 5.9 1.8 2.4 Tularosa Cave—L. 6 7 4 1.3 4.7 48 5.5 17 2.3 Tularosa Cave—L. 11 8 0 1.0 4.0 4 5.5 1.7 2.3 oint of Pi 8 20 t2 3.6 3 5.5 1.7 2.0 apalote - 0 1.0 4.0 10 5.4 zl 2.5 Guatemala Flint 15 0 13 4.5 44 6.8 1.9 2.9 er California 8 37 1.3 5.8 27 5.9 1.8 2.1 Peru Flour 8 8 .9 4.0 92 5.0 1.9 24 roic 13 0 P 6.6 95 5.9 1.9 11 i 6 0 5 3.2 30 3.5 1.6 2.1 Huaca Prieta 1 4 5 .3 Ai 31 2.4 1.6 1.8 Huaca Prieta 5 6 9 2 3.0 13 а 1.9 1.9 Assam 12 0 .9 4.9 75 5.0 1 2.3 Lushai 10 0 s 4.0 75 6.0 1.9 2.0 МЯҤачу5 'IVOINV.LOdH IWüOSSIW AHL AO SIVNNV 0p тод] 1953] NICKERSON—VARIATION IN СОВ MORPHOLOGY 89 зр ne Ш 5 SACRED MOVE il tower QVATEMALA ном BLUE ном WHITE Е CAUFORNIA MNT nove nour P] N AUSMAI ASSAM TULAROSA PAPAGO TULAROSA MARSH LUSTER uvm + = МУН 2 PASS CAVE w P о $075 H BASKETMAKER eu, o se ive n % * v COROINCO CHAPALOTE шее эд Хы” 3P e 2 A Г a L A a n Г n Г B. —————— Sl 2.00 1.50 1.00 .50 0 .50 1.00 150 200 neg pos CUPULE DEPTH MM. PERCENT OF COBS 8- ROWED $ FY a 100 . 51 50.1 0 SHANK DIAMETER é 4 * . 16 15 . 10 9. PERCENT OF СОВЅ STRAIGHT 9 . . 30 29 . 10 9.0 HEIGHT ОҒ RACHIS - FLAPS v ә . 20 19.9 Ж. KERNEL THICKNESS » E . 244 41. 36 35. PERCENT OF COBS TAPERED » bd . 6l 60.31 30.0 LOWER GLUME WIDTH ~ ~ а „60 59.40 39 . sa б Further posal tion in скалой л. in ug 1, showing relation- of maize. Each dot in that с groupi horizontal‏ ا ters are diagrammed by rays,‏ NHGWVO 'IVOINV.LOS MNOSSIW AHL AO SIVNNV vw К y yy v w + м y » 3 м LUSHAI MODERN IROQUOIS SACRED FLOUR MODERN V L y $ es ON у w E dy % v% v% « + €, зу W N зуу ws Е" > ч ASSAM MODERN NORTHERN FLINT mere Ob TOA] 5 * ~ 7 № S v, | л, ws $$ * > 4 rs è . vee o,f E k^ ` e а r e Ы L HUACA PRIETA 1 850 ВС.- 1000 AD. ARICA 0 - 1250 SINGLE SPECIMEN SCORES NUMBER OF ROWS 4 $ ° 8 10 2 STRAIGHT СОВ « e PRESENT ABSENT М TAPERED СОВ ° E ° PRESENT ABSENT HUACA PRIETA 5 850 B.C.- 1000 AD. Text-fig. Te Pe 3, 4, 5. Twenty-three eee — diagrams dita. ie variation and relationships of nine measured maet erat TM each group of maize cobs stu — Each dot esents the actual measurements e ear. Horizontal and vertical axes and seven additional characters are scored as in text-fi xcept row number a nd roy erue which were ind. as indicated in the lower right-hand corner of text- ug E Dotted 8. 2, ray indicates no iita could be m A9SO'IOHdWONW 800. NI NOLLVINVA—NOSWHMOIN [£861 92 ANNALS OF THE MISSOURI BOTANICAL GARDEN > > »»»» НОРІ WHITE FLOUR MODERN ~ [Vor. HOP! BLUE FLOUR MODERN 40 PERU PLOUR MODERN GUATEMALA FUNT MODERN + * Ф v * + aww wes vv A. c ou» X + + » 9% Ф y + № wre & % + v Ne + ^ x € » Ф Ф «4. У % LOWER CALIFORNIA 1650 1700 MARSH PASS 300 1000 v à Ww Y we "nw Sew у $ wee ч аҹ <“ é ч чоч i в + % * * r ¥ » < COROICO MODERN TURNER SITE 1000 . 1100 Text-fig. 4 AS9O'IOHdWONW ЧОО NI NOLLVIWVA—NOSWAWOIN £6 [8661 є 7 v v X* ~ » v. WM у v + x چ‎ САЛЫ $ ~ > * LUSTER CAVE 900 - 1000 PAPAGO MODERN ¥ % * ¥ + 4 $ М 422972. v 4 69 à Ф é € - CHAPALOTE MODERN TULAROSA LEVEL 3 900 - 1000 r6 NHGWVO 'IVOINV.LOH INNOSSIW AHL ЧО SIVNNV 0p '10A] > Ф s% bé de ۳ xy ei. Per. $$ • * * © POINT OF PINES 1250 1300 e Y “i v Ф v vvv M $ LY ر‎ v Wey ¥ TULAROSA LEVEL 6 500.700 BASKETMAKEP 200 300 Text-fig. 5 TULAROSA LEVEL П 300 BC.- 150 BC. A9O'IOHdWONW 800 NI NOLLVIYVA—NOSYAMOIN S6 [£ $61 [Vor. 40 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN conform to the same general description. They are tapered, 15—25 cm. in length, with a medium-sized shank and bore 14—16 rows of narrow kernels. That they are markedly lighter in weight than ears of North and Central American maize of comparable size is undoubtedly another expression of their lack of heavily sclerenchymatized tissues. MEASUREMENTS OF SAMPLES The number of cobs in each of the above-listed samples varied, but an attempt has been made to examine enough cobs to form consistent pictures of variation in the populations represented. Measurements recorded for each cob include, in addition to those of four external and five internal characters mentioned earlier. diameters of cob, rachis, and pith. From these last measurements, а cob/rachis index and a rachis/pith index were computed. Since plants are affected by their environment in numerous ways, several rep- resentatives of any population must be studied to formulate a complete picture of their range of variation. This idea applies quite as much to archaeological remains as to living plants, for, as Cutler (1952) has pointed out, a single speci- men of any archaeological plant remains is of little significance; it may represent TABLE II AVERAGE VALUES OF FIVE MORPHOLOGICAL CHARACTERS FOR ТУ/ЕМТУ-ЕТУЕ EXISTING RACES OF MAIZE IN wr ur (DATA FROM WELLHAUSEN ET AL, 1 5 х (Measurements іп mm. except where otherwise indicated.) Cupule Width of % cobs Shank Kernel Name of race width lower glume 8-rowed diameter thickness Palomero Toluqueüo 3.0 3.0 0 8 2.8 АгосШо Amarillo 3.5 3:5 0 8 2.5 Chapalote 5.5 55 20 9 4.1 Nal-Tel 5.0 9:9 20 Z. 3.9 Cacahuazintle 5.0 7.0 10 5.2 Harinoso de Ocho 8.0 9.5 100 14 4.4 Olotón 7.0 8.5 20 17 6.0 Maíz Dulce 7.0 6.0 0 11 4.0 ӧпісо 4.0 5.0 0 8 3.6 Reventador 5.0 5.5 20 8 3.6 Tabloncillo 9.0 10.0 90 11 4.3 hua 10.5 9.5 0 21 3.9 Tepecintle 9.0 8.0 0 10 3.7 iteco 8.0 9.0 0 22 4.5 8.0 7.0 0 34 4.6 Zapalote Chico 7.5 7.5 20 13 3.6 Zapalote Grande 7.5 7.0 0 18 3.8 Pepitilla 5.0 0 12 3.5 Olotillo 7.5 10.5 90 10 3.9 Tuxpefio 7.5 0 20 13 3.7 Vande 8.5 8.5 0 13 3.6 Chalquefio 6.0 6.5 0 10 3.9 aya 7.3 7.0 20 9 LY Cónico Norteño 6.0 3:5 0 11 3.5 lita 9.0 10.0 50 9 4.1 ا ا Шш‏ ت 1953] NICKERSON—VARIATION IN COB MORPHOLOGY 97 an accidental deposition, or it may have been buried in an old layer or brought to a recent one by rodents, pot hunters, or a recent occupant of the site. To mini- mize these effects, arithmetic averages of the measurements of each character have been computed for each sample. Alava (1952) pointed out that such a technique provides the only safe basis for studying variation between different varieties of plants. Table I lists the average values for each of the characters of each sample studied. Ear shape contributed two columns, one for straight ears and one for tapered ears, thus giving a total of nine characters in which to compare variation. A pictorialized scatter diagram of these same results is presented in text-fig. 2. зр 1 MARSH 9ASS 2 TULAROSA LEVEL Ф 3 PAPAGO — ^ OUR ne 4 TULAROSA t 5 OWER — CALIFORNIA Е Др». i нот BLUE nour a ; мот white пош rom Ó SOUTA т NORTHERN „>. TABLONCILLO \ You = E ane „ә Е ня a£ aped tyster > ГЕ B on m? b E di s № | v : 25 pe XA „бы "Sar d A me S 5 or hints pes conico 5 ВАЗ ЕТМАКЕЕ ENO CHALQUENO w ore у О araor - d a E] REVENTADOR d v 5 TURNER TULAROSA coroico CACAMUAZINTLE © SITE LEVEL п e das "0 b м comico те oS HUACA PALOMERO AROCILLO PRIETA $ TOLUQUENO AMARILLO r : А : x ^ a a A 2d —— EUN Y А т n n А ~ 8 9 10 n LOWER GLUME WIDTH MM. LJ PERCENT OF COBS 8 - ROWED E Ф 100 . 75 74 - 5l 50.0 SHANK DIAMETER E é Ф - lé 15 . 10 9. LJ KERNEL THICKNESS » ә „АЛ 41. 36 35 - Text-fig. 6. Pictorialized scatter diagram of material from Tables I vi II —Á — ships betw variation in five different cob characters among the twenty-five races tw * Each dot represents the average measurements of five characters for all maize 8rOup; horizontal axis, width of ees er glume; vertical axis, width of Lp - diagrammed by rays as sistas d on the figure. Further explanation in [Vor. 40 98 ANNALS OF THE MISSOURI BOTANICAL GARDEN Text-figs. 3, 4, and 5 show the variation and relationships of these same nine cob characters within each sample of cobs. The method of construction, as well as the general usefulness and reliability of these diagrams, is further discussed below. An attempt was made to compare the results of this investigation with those obtained by Wellhausen e£ al (1951, 1952) for races of maize in Mexico. Several of the major characters employed in the present study were not employed by Wellhausen and his co-workers. However, their paper contained excellent diagrams of cross-sections of ears of each race drawn to scale; it was possible to make certain measurements directly from these diagrams. Lower glume width in mm. was measured at the widest point indicated on the third concentric circle of each езг diagram. Cupule width in mm. was measured as a straight-line distance across the bases of any two consecutive kernels from the points where their boundaries intercepted the rachis diameter (innermost circle). Percentage of 8-rowed ears was obtained by comparing the number of rows in each drawing with the average values listed for each race in their Table 15. Average values for shank diameter and kernel thickness were obtained directly from their Table 15. The informa- tion obtained is summarized here in Table II; it is also presented as part of text- 8. 6, a pictorialized scatter diagram which compares related data from both Tables I and II Male spikelet variation has been studied by Alava (1952) for seven of the twenty-three samples used in this investigation. А crude index has been con- structed after the manner outlined by Anderson (1936) for both Alava's data and the appropriate data set forth in text-fig. 2. Text-fig. 7 shows the relation- ships of these index values. The significance of each of the above comparisons is discussed below. Discussion Whiting (1944) and Cutler (1951) pointed out that botanists alone probably cannot solve the problem of the origin of maize, because maize is so intimately linked with human cultures. This point is emphasized by the fact that maize is completely dependent upon man for its preservation. Whatever may have been the original factors leading up to this novel situation, the fact remains that maize and man have been associated years. In a study which involves the disciplines of ethnology, archaeology, and botany, caution must be exercised lest theories proposed in one science be regarded as proven facts by workers in another. Since the present investigation has been conducted from the standpoint of botany, such conclusions as can be drawn should depend in so far as possible upon botanical evidence for their support. To this end, botanical information is discussed first, and an attempt is then made to relate this discussion to known facts of ethnology and archaeology. 1953] NICKERSON—VARIATION IN COB MORPHOLOGY 99 BOTANICAL AsPECTS OF MAIZE Сов ANALYSIS Variation among races of maize is nowhere better illustrated nor more difficult to describe than that found among female inflorescences, or cobs. Within any race, many cobs possess characters the measurements of which lie well within the range of variation of the same measurements from several other races. This exasperating fact is true for all the characters used in this study. Using any one cob character at a time, it is nearly impossible to distinguish races of maize; it is for this reason that the empirical classification of maize by kernel types proposed by Sturtevant (1899) is not of much value to modern maize breeders. Mangels- dorf and Reeves (1939) used evidence from several sources to characterize maize from different parts of the New World. Anderson (1946, 1947a), Anderson and 15Р өс DE ФЕ 2 Б oF өс < eb ot < > of o 6 № ш өз ә ә < Гем 3 № ФА + r1 i L JL — НЮ 3 6 9 12 15 18 СОВ CHARACTERS INDEX VALUES MAIZE RACE СОВ CHARACTERS TASSEL CHARACTERS А ARICA 1 1 B ASSAM 9 5 C CHAPALOTE y 9 D PERU FLOUR n 8 Е GUATEMALA FLINT 13 8 F NORTHERN FLINT 18 12 7 15 С PAPAGO Text-fig. 7. Scatter diagram of index values of seven races of pis i in which both the x ага (male їпйогезсепс ce) and the cob (female inflorescence) have been inves gated. Horizonta к ex values red e" cob study; vertical axis, index values derived as tassel study, Further explanation’ in the [Vor. 40 100 ANNALS OF THE MISSOURI BOTANICAL GARDEN Cutler (1942), Mangelsdorf and Smith (1949) and Wellhausen e£ al (1951, 1952) ave used an increasing number of subtler characters in distinguishing both modern and prehistoric races of maize. Pictorialized scatter diagrams (Anderson, 1949b) are a means by which several such variable characters can be considered at once, thereby emphasizing consistent and typical differences as well as similarities among the particular entities involved. Methods by which a pictorialized scatter diagram is developed are explained by Anderson (1949b, 1952) and Anderson and Gage (1952). Anderson (in press) has discussed the biological and mathematical criteria upon which pictorialized scatter diagrams are based. The value of such diagrams has been emphasized by Stebbins (1952), who considered the method “by far the best yet devised for making the observer aware of a pattern of variation in respect to three or more characters which are varying simultaneously." He also stated (1952) that the method "has the advantage of being repeatable and of focusing the attention of the observer upon certain essential features of variation." The data listed in Table I have been plotted as a pictorialized scatter diagram (text-fig. 2) to show how the several cob characters are varying and to what extent their variation is correlated. Measurements of depth and width of cupule were chosen as abscissa and ordinate respectively because they varied consistently within each sample (text-figs. 3, 4, 5), could be ascertained from fragmentary cobs, and were susceptible to accurate measurement. The other seven characters were then indicated by rays from each dot of the scatter diagram. Extremes of each character tending to be associated with higher values of cupule width and depth were represented by long rays, while extremes tending to be associated with lower ordinate-abscissa values were represented by no rays. Several facts concerning the maize samples studied were immediately apparent. At one extreme (upper right-hand corner of the diagram) were predominantly straight 8-rowed cobs with wide, shallow cupules, large rachis-flaps, wide lower glumes, heavy shanks, and thick or moderately thick kernels. This complex of characters was found in Northern Flint and Iroquois Sacred Flour; occurrence of endosperm as flinty or floury is dependent upon a single gene and is therefore of no significance here. These results agreed with those found by Brown and Anderson (1947) for typical Northern Flint varieties; the same complex was identified by Carter and Anderson (1945) as the Eastern complex. At another extreme (lower left-hand corner of the diagram) were cigar-shaped cobs with high row numbers narrow deep cupules, small rachis-flaps, narrow lower glumes, small shank diam- eters, and thin kernels. "These characters were associated in prehistoric maize recovered at Huaca Prieta, Peru, and Arica, Chile. It is significant that the majority of cobs were essentially complete but only about 5 cm. long. As a whole, these samples were remarkably homogeneous (text-fig. 3). Existence of a thi extreme (middle left area of the diagram) is indicatd by tapered cobs with high row numbers, extremely deep cupules, little rachis-flap development, wide lowe! — glumes, medium shank diameters, and thick or moderately thick kernels. The two _ 1953] NICKERSON—VARIATION IN COB MORPHOLOGY 101 maize samples from Asia were responsible. Cobs studied were those grown by Stonor in Asia; varieties involved were described by Stonor and Anderson (1949). The most striking features of the Asian corn cobs were their extremely deep cupules and their particular form of sclerenchymatization. The rachis tissue and lower glumes of all Asian cobs studied were quite hard, but their hardness was not identical with that of “bony” rachis tissue described by Mangelsdorf and Smith (1949) in connection with teosinte introgression. Whatever its cause, the end product was an extremely stiff, light-weight bulky cob. Studies by Lenz (1948) did not indicate that cell wall thickness had much to do with degree of sclerenchy- matization among various races of maize. A deep cupule could possibly be in- terpreted as a condition which showed little if any “Тирзасо influence" (Cutler, 1946). An inference cannot be drawn that all Asian maize varieties are alike in this respect, but for both Assam and Lushai samples, this conclusion seems justified. The nine characters employed in constructing text-fig. 2 showed a strong tendency for association at both extremes. A strong overall association between all sample averages as well as within each sample is clearly shown by comparing the average of any sample in text-fig. 2 with its appropriate population diagram in text-figs. 3, 4, or 5. Individual cobs in the upper right-hand area of each population diagram consistently showed more and longer rays than did those from other parts of the same diagram. This similarity of separation patterns in the several diagrams emphasizes the fact that those limits employed in separating sample averages, while originally chosen in a subjective taxonomic fashion (Steb- bins, 1952), were also working within each sample in such a way that morpho- logical trends associated with a particular race of maize were associated with each other. The diagram as constituted contains samples of rather diverse races of maize; closeness of position on the diagram and similarity of glyphs indicate similar morphology but do not necessarily imply close relationship. Weatherwax (1936) stated that no sound basis exists for any attempt to trace maize origin by arranging Varieties from different localities in an evolutionary sequence. Introduction of the dimension of time in the form of archaeological material, however, allows the investigator to arrange maize varieties of both present and past according to cultural succession; the result may or may not be an evolutionary sequence, but Such an arrangement is the strongest means available for documenting maize history and has therefore been employed in this investigation whenever possible. Results obtained in this investigation are compared below with those obtained in three other independently conducted investigations. A high degree of correla- tion is found in each instance, a fact which indicates a basic soundness in methods employed for cob analysis in this investigation. NE Brown e£ al (1952) have discussed relationships among three — of Hop Р maize, two of which, Норі White Flour and Норі Blue Flour, were included in the present study. These workers found that the two varieties had certain re- [Vor. 40 102 ANNALS OF THE MISSOURI BOTANICAL GARDEN semblances to Basketmaker and Eastern maize as well as to each other. Unfor- tunately, it was not possible to obtain a large enough sample of Kokoma, the variety they reported as resembling Basketmaker most closely, to warrant its inclusion here. Text-fig. 2 indicated that Hopi Blue Flour was basically similar to Basketmaker, differing in cupule width, wide lower glumes, heavy shanks, and presence of 8-rowed cobs. It should be noted that there has been no change in cupule depth and predominance of cigar-shaped ears in development of Hopi Blue Flour from Basketmaker maize. Hopi White Flour was found by Brown e£ al (1952) to contain an admixture of both Eastern and Mexican germ plasm. The Mexican complex (Carter and Anderson, 1945), known to be present on the Mesa Central of Mexico, is char- acterized by strongly tapered cobs with high row numbers, thick kernels, and small shanks. This conclusion was borne out by text-fig. 2, which showed Hopi White to have all the characters associated with the Eastern complex with the exception of row number and cob shape; tapered cobs of high row number may be attributed to Mexican influence. Cobs of these two Hopi Flour maizes differed from each other in cupule depth (text-fig. 2 indicates this to be the strongest character con- tributed by the Eastern complex), cob shape, shank diameter, rachis-flap height, and row number. In the correlation between results obtained in the present investigation and those of Wellhausen e£ al (1951, 1952) which are listed in Table II, the number of characters used was less than the number employed to separate the samples listed in Table I; yet, a pictorialized scatter diagram (text-fig. 6), using five measurements, presented essentially the same variation pattern as did text-fig. 2, in which nine measurements were employed. It should be noted that the basic pattern of maize cob variation was not altered by using measurements of a different character for the abscissa of text-fig. 6 than that employed in text-fig. 2. If a sample on text-fig. 6 has any real affinity to races with which it may be allied on other grounds, the fact should be shown by its position on the diagram. Text-fig 6 should also lend further confirmation to the genealogies postulated by Wellhausen et al for several hybrid races of maize. That these confirmations are indeed pos- sible was shown quickly and easily. The four races classified by these workers as Ancient Indigenous Races (Palomero Toluquefio, Arocillo Amarillo, Chapalote, Nal-Tel) were closely grouped; moreover they were all placed in the lower left- hand corner of the diagram and all had few to no rays. Thus their characters 0 small cupules, narrow lower glumes, high row numbers, thin kernels, and small shanks, all believed to indicate primitiveness in relation to other races of maize here considered, were strikingly indicated by text-fig. 6. Three of the four races classified as Pre-Columbian Exotics (Cacahuazintle, Harinoso de Ocho, Olotón, Maíz Dulce) formed a consistent graded series along the lower edge of the upper region of text-fig. 6. They were placed in a separate class because of their antiquity and resemblance to South American maize. i 1953] NICKERSON—VARIATION IN COB MORPHOLOGY 103 the class was somewhat artificial, the positions of three of its four constituents on the diagram in a nearly perfect ascending order indicated a basic similarity in variation pattern. From these races a group of thirteen Prehistoric Mestizos and a group of four Modern Incipient races were thought to have arisen. For many of these, Well- hausen e£ al compiled a chart showing the two putative parents of the particular race under discussion. Out of a total of eleven genealogical relationships involving other races plotted on text-fig. 6, seven were found to be borne out by diagram position, being for the most part midway in both position and number of rays. These genealogical relationships as postulated by Wellhausen e al are listed below. Group 1 is comprised of those races whose positions on text-fig. 6 offered further proof of relationship to their putative parents. Group 2 is comprised of those races whose positions on text-fig. 6 did not approximate positions between their putative parent races. GROUP 1 Cónico. Palomero Toluqueüo X Cacahuazintle Zapalote Chico : al-Te X Tepecintle Chalquefio Cénico X Тахрейо ónico Norteño. Celaya х nico ; Bolita Zapalote Chico x Tabloncillo Tuxpefio. Tepecintle X Olotillo Comiteco T X Olotén GROUP 2 Zapalote Grande Teh x Zapalote Chico andeno. Тох; x Zapalote Grande Celaya Tabloncillo х Tuxpeño Mhao Tabloncillo x miteco The fact that а majority of such genealogies can be substantiated by arranging data obtained from the cobs alone is added proof that races may not only be recog- nized by cob characters, but that their histories can also be ascertained in the same manner with a high degree of accuracy. It should also be borne in mind that data for the Mexican samples represented averages determined from 3—5 cobs of each race; the accuracy might well have been increased had more cobs been used in determining average values. Text-fig. 6 further showed that variation in Mexican maize followed the same general pattern of character association as did Variation among the samples employed in this investigation, and that prehistoric North and South American samples were close to the primitive Mexican races. - Spikelet variation in Zea Mays was studied by Alava (1952), using methods Which, while applicable to male inflorescences, were of no use in the investigation of maize cobs. Thus if any correlation between results of two independent y conducted investigations should exist, it would not only be further substantiation for validity of methods employed and results obtained, but also would indicate that cob and tassel analyses yield comparable results. Similar material from seven different races was examined in each investigation. А numerical index (Anderson, [Vor. 40 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1936; Stebbins, 1952) for both cob and tassel characters was computed for each race. ile admittedly a crude indicator of morphological characters, numerical indices are valuable for summarizing diverse data in a manner which permits direct comparisons to be made. "Text-fig. 7 shows the extent of this correlation. Arica had the smallest value in both indices, and there was a general progression toward the upper right-hand corner of the graph occupied by Northern Flint. Papago appeared somewhat aberrant as far as tassel index was concerned, but came closest to Chapalote, a related race, in both values. Peru Flour and Guatemala Flint were close, a further expression of their presumed relationship to one an- other. Guatemala Flint was closer to Northern Flint for cob-index value. Although a considerable number of units away, Asian material was closest to Arica in both cob and tassel values. Thus with the possible exception of Papago, in which variation is unique but consistent with its past history, the samples showed high correlation between tassel and cob morphology in distinguishing races of maize. Maize Сов ANALYSIS IN RELATION TO ETHNOLOGY AND ARCHAEOLOGY Knowledge of ancient peoples is in large part built upon a detailed study of their refuse heaps; rubbish being what it is, a place is assured for the botanist in archaeological analysis. Analysis of maize remains is especially helpful, since few other plants have become so closely associated with man to the extent that they are reliable indicators of his early history in many parts of the New World. SOUTH AMERICA AND ASIA Lowie (1940) pointed out that it was a common mistake to identify all early agriculture with maize. Sauer (1952) contended that seed-crop agriculture was developed after a root-crop agriculture which involved vegetative propagation only. Excavations in northern Chile (Bird, 1943), and more especially in Peru (Bennett, 1948; Bird, 1948), showed that there were indeed agricultural com- munities without maize. Layers of middens at Huaca Prieta, Peru, beginning about 4000 years ago, contained squash, beans (4 types), chili peppers, Canna, cotton, and bottle gourds (Lagenaria siceraria, Whitaker and Bird, 1949). Maize did not appear until 850 В.С. Whitaker (1948) pointed out that it is extremely difficult to account for the bi-hemispheric distribution of Lagenaria, as it is pre- sumed to be of Asiatic origin. Partly on such distributional patterns, Carter (1950) postulated pre-Columbian contacts between the Old and New Worlds, a theory also put forward by Stonor and Anderson (1949) and Sauer (1952). That it was quite possible to sail in either direction across the broad expanse of the Pacific has been amply documented by Buck (1938). Another odd fact about prehistoric plants of west-coast South America is that they are presumed to be native to Central America. Either these plants reached the coast as articles of trade among established peoples, or they were taken there by the first settlers of the area, who either came from or passed through the Cen- 1953] NICKERSON—VARIATION IN COB MORPHOLOGY 105 tral American region. It does not seem possible that maize would have been over- looked by these peoples as a crop plant had they passed through an area in which it was growing or being grown. Yet evidence on the possible antiquity of maize in central Mexico was recently reported by Barghoorn (1952). Maize pollen has apparently been recovered from sedimentary deposits under Mexico City which may antedate human occupation. Recently, however, the date of human occupa- tion has been pushed back to 9000 years in the same area (Richards, 1953). A type of maize hitherto not mentioned in this investigation is that which is generally referred to as popcorn. True pops are generally of quite primitive mor- phology (Mangelsdorf and Smith, 1949). Prehistoric Arica and Huaca Prieta maize remains may quite possibly be those of popcorns; Wissler (1945) showed that this is true for some whole maize ears excavated at Arica. In ear shape and size and in tassel characters (Alava, 1952), these prehistoric remains were com- parable to a modern pop variety collected by Parodi in Argentina. They may also be related rather closely to Asian types, since they had deep cupules, high row numbers, and some tapered cobs. Tassel analysis (Alava, 1952) also showed an ' Asian similarity. The above evidence is suggestive, but it also indicates the neces- sity for more data before the history of maize in both South America and Asia can be considered to be completely known. THE AMERICAN SOUTHWEST Maize in the American Southwest has had a long and complicated history. Man was present in the area 10,000 years ago (Wormington, 1949; Johnson, 1951) but there was little evidence that the area was under continuous occupation from that time until the advent of agriculture. Randolph (1952) stated that an ap- parently humid and subtropical climate existed there 5,000—10,000 years ago, and that the area itself could be the original home of maize. Carter (1945) likewise postulated the Southwest as a center of agricultural dispersal. All known archae- ological remains which contain maize are younger than 5,000 years, so the problem at present is to evaluate the evidence at hand. It should be kept in mind that whatever statements are made below are subject to revision in the light of further discoveries, Carter and Anderson (1945) noted three major cultural provinces in the Southwest: Hohokam, Mogollon, and Anasazi (Puebloan). At the time of their survey, little was known of the Mogollon civilization, but recent excavations at Tularosa and Cordova Caves (Martin et al, 1952) have produced evidence estab- ishing this cultural province as one of equal importance with Hohokam and Anasazi. Archaeological maize is now known for each of the three cultural Provinces. Hohokam maize was found in Ventana Cave, Arizona (Haury, 1950). It is similar to but not identical with modern Pima-Papago varieties (Carter and Anderson, 1945). Chapalote, an ancient Mexican race, is today found in north- Western Mexico, an area which was also part of the Hohokam cultural province (Amsden, 1949). Relationships between Papago and Chapalote have been dis- [Vor. 40 106 ANNALS OF THE MISSOURI BOTANICAL GARDEN cussed above. Since the Hohokam area was relatively isolated from influences of neighboring cultural patterns, the maize found there today is much the same as it was in ancient times. It is reasonable to assume that prehistoric maize in this cultural province was similar but not identical to that of Basketmaker and Mogollon cultures, and that modern Papago and Chapalote were derived from it. It was shown (text-fig. 2) that maize remains recovered from Tularosa Cave Level 11 were quite similar to both Chapalote and Basketmaker. Similarity between Chapa- lote and Basketmaker П maize from Cottonwood Cave (Hurst, 1948; Hurst and Anderson, 1949) was pointed out by Wellhausen e£ al (1951, 1952). Amsden (1949) noted the similarity of modern Papago to Basketmaker, as did Carter and Anderson (1945). The present investigation showed that Basketmaker resembled Chapalote much closer than it did Papago, but the important point is that a three cultural areas, Hohokam, Anasazi, and Mogollon, had at an early date maize which was variable but essentially similar. Maize of the Hohokam area remained _ relatively unchanged; that of the other two areas was strongly influenced by maize from more remote places. Text-fig. 5 illustrates the variable nature of early southwestern samples. In contrast to maize of the Southwest, remains from Arica, Chile, and Huaca Prieta, Peru, are very homogeneous; furthermore, maize of these two areas (American Southwest and South American West Coast) were con- temporary. Explanation of the variability in the one location and homogeneity in the other at the same time is but one of the many problems of maize history. Basketmaker maize was widespread, having been reported from several sites in the Anasazi area. Some of the better known sites are Cottonwood Cave, western Colorado (Hurst, 1948; Hurst and Anderson, 1949), Mummy Cave, Сайоп del Muerto, Arizona (Anderson and Blanchard, 1942), and Painted Cave, northeastern Arizona (Haury, 1945). Basketmaker-like maize was reported outside the Anasazi area by Brown and Anderson (1947) in prehistoric remains of rock shelters and caves from the Ozarks to southern Ohio. Two combinations of maize characters, one peculiar to the Mexican Mesa Cen- tral and the other to the eastern United States, were superimposed in varying amounts on southwestern maize. The spread of each influence may be traced by analysis of successive samples; in such work, the importance of dated maize remains can hardly be over-emphasized. Mexican influence is characterized by strongly tapered cobs with high row numbers (Carter and Anderson, 1945), thick kernels, and small shanks. This type entered the Southwest from the Northeast rather than from Mexico directly; its greatest influence has been found in remains of the Fremont Basketmakers of Yampa Canyon, Utah, dated at 400-800 A.D. (Burgh and Scoggin, 1948). At Luster Cave, in extreme eastern Utah, maize was subjected to strong Eastern influence prior to 1000 A.D. Text-fig. 2 showed this sample to be closely related to Hopi White Flour among ethnological races examined in this study. Нор. | White Flour differed, however, in being more extreme for certain Mexican (tapered Ad 1953] NICKERSON—VARIATION IN COB MORPHOLOGY 107 ears, high row numbers) and Eastern characters (large shanks, wide lower glumes). Maize from the Davis Site (Newell and Krieger, 1949) was analyzed by Jones (1949) and found to be entirely Eastern. These maize remains were dated at about 700 A.D. by Jones and at about 400 A.D. by Johnson (1951). The question of origin of Eastern influence has long been a puzzling one. Anderson (1947a) and Brown and Anderson (1947) presented evidence that Eastern maize is related to Guatemalan maize. Jones (1949) reported that he and Krieger independently reached the conclusion that both maize and pottery types recovered at the Davis Site bore a resemblance to those of Guatemala. Carter (1946) suggested that maize could have been carried up the east coast of Mexico or across the Gulf to southern United States. Regardless of the method employed, this movement must have taken place in time for the influence to have been carried to the Southwest by 1200 A.D., a date which Carter and Anderson (1945) recognized as closely approximating its first appearance in that region. Judson (1951) called attention to the fact that a great drought occurred in the Southwest in 1276-1299 A.D., and that many Indian tribal migrations took place around that time. The direction of movement of maize into the Southwest was from east to west; Carter and Anderson (1945) noted that among the present-day Puebloan tribes, those of eastern pueblos have more eastern-like maize than do those of western pueblos. It is generally accepted that there is influence of Plains cultures in the southwestern area. This influence also is stronger in eastern pueblos than in western ones, and is another indication of how well maize history is cor- related with the history of the peoples who grew it. THE AMERICAN SOUTHEAST AND EAST Wherever the Eastern complex came from, it was probably carried up the river valleys of the Mississippi and its tributaries by Indian tribal migrations. A date of 900 A.D. was assigned to the earliest known Burial Mound I culture by Ford and Willey (1941), who also identified the beginnings of horticulture in eastern United States at this time. Burial Mound I peoples were in turn supplanted by Burial Mound П and later (1200-1400 A.D.) by Temple Mound I and Temple Mound II cultures. Ford and Willey derived the Iroquois culture of New York from a welding of these four intrusions onto an archaic hunting-gathering popula- tion; they considered that this and other allied upper Mississippian cultures reached a peak after 1500 A.D., and lasted until historic times. Although other evidence has been presented in support of the idea that maize was a late arrival юса woodland cultures (Linton, 1924; Kroeber, 1939), from a botanical point of view the postulated time of arrival of maize in this area appears too recent. This impression is strengthened by consideration of the length of time the same type of maize was present in the Southwest. Recent radiocarbon dating (Johnson, 1951 and other eastern cultures were of greater antiquity ) indicated that Mississippi valley than had previously been [Vor. 40 108 ANNALS OF THE MISSOURI BOTANICAL GARDEN thought. Johnson (1951) reported that Hopewell sites from Ohio and Illinois were about 1000 years older than had previously been estimated. The whole chronology of the Southeast has become unsettled because of radiocarbon dating results, but the presence of cultures in this area at earlier dates further strengthens the idea that maize is of greater antiquity than had been thought possible, and opens a way to minimize a long-standing inconsistency between archaeological and botanical evidence. Since there was, as Ford and Willey (1941) and Waring and Holder (1945) pointed out, a suggestion of strong Mexican influence in Temple Mound I and П cultures, the idea would be greatly substantiated should remains of Mexican maize ultimately be found in excavations. Botanical evidence exists that maize of the Southeast is related to a Mexican Gulf Coast race (Wellhausen e£ al, 1951, 1952), and hybridization of this Mexican maize with eastern flint maize already present in the Southeast probably resulted in the variable forms called Southern Dents by Brown and Anderson (1948). Northern Flint and Southern Dent maizes were subsequently brought together to form modern hybrid corn-belt maize (Anderson and Brown, 1952a, 1952b). A significantly large collection of Southern Dents was not available for inclusion in the present study, but the few available cobs which were studied showed a rather close resemblance to Northern Flints in cob characters. Evidence that many Southern Dents were intermixed with Northern Flints was presented by Brown (1949), who studied chromosome knob numbers in United States maize. He found Northern Flints to have the lowest knob numbers, Southern Dents the highest, and corn-belt forms intermediate between these extremes. Longley (1938) had previously surveyed Indian maize varieties from the United States and northern Mexico. He reported the same general rise in knob number among southeastern varieties. Не also found that maize of Arizona and New Mexico was high in knob number. Carter (1949) used knob numbers on chromosomes of Indian maize to indicate tribal affinities and differences. The fact that such an investigation yielded results substantially the same as those of different approaches is another example of how closely maize mirrors the history of those with whom it is associated. Ап important point regarding use of Northern Flint and Southern Dent maizes in corn-belt maize production is that the hybrid vigor manifest in this cross is based upon small differences between two already-intermixed races. This situation may possibly indicate that improvement of maize by hybridization techniques has barely begun. Since identification and classification of races are becoming à necessary part of maize breeding, a knowledge of cob variation should be of prac- tical value in the development of new strains of hybrid maize. The principles of cob analysis set forth above, even though they employed small morphological differences, may be considered fundamentally sound, because results obtained from their analysis were in agreement with ethnological and archaeological as well as botanical data of other workers. 1953] NICKERSON—VARIATION IN COB MORPHOLOGY 109 SUMMARY Morphological characters present in the female inflorescence, or cob, of Zea Mays L. have been measured on over 500 cobs representing both modern and archaeological varieties. External characters measured included row number, shank diameter, cob diameter, and over-all cob shape. Internal characters measured in- cluded cupule width, cupule depth, height of rachis-flaps, kernel thickness, lower glume width, rachis diameter, and pith diameter. Analysis of pictorialized scatter diagrams of averaged measurements showed a high degree of association of these characters. These results agreed closely with those of previous investigations employing other methods. The results of archaeo- logical maize analysis were in harmony with previous conclusions based on purely archaeological data. Such agreements indicate the validity of cob analysis for characterizing variation in races of maize. LITERATURE CITED Alava, Reino O. (1952). Spikele variation in Zea Mays L. Ann. Mo. Bot. Gard. —96. Am > en, a чуч pui ү ку 28 ehistoric southwesterners Кон basketmaker to етты 163 рр. uthw. Anderson, bani. (1956). x m жнын in American Tradescantias. Ann. Mo. Bot. Gard. 23:511- ‚ (1943). A variety of maize from the ча Loa. Ibid. 30:467—475. ‚ (1944a). Maiz Asiae Ibid. 31:301-315 , (1944b). Homolog the ear and tas ats Zea Mays. Ibid. 31:325—344. (1944c). Two pt saria of prehistoric corn tassels from southern Utah. Ibid. 31:345- > 354. — — —— , (1946). Maize in Mexico—a preliminary survey. Ibid. 33: و بر‎ — — (1947a). Field suiii rà E die a aize. Ibid. 34:433—4 —————, (1947b). Corn before Columbus. 24 pp. Pioneer Hi-Bred я Co; Des Moines. oft —— ——, (1949b). Introgressive hybridization. 109 pp. John Wiley & Sons, New York — — ——, (1951). The sacred plume. 24 pp. Pioneer Bre r , Des аем 19323 nts, man and life. 245 pp. Little, Bro ., Boston. » (In € s). Efficient and inefficient methods of measuring specific differen 3 O. Bla aes (1942). Prehistoric maize from Cafion del Монев Ка: Jour. Bot. 29: 832-835. , and William L. Babe (1948). A morphological analysis of row number i Mo. Bot. Gard. di п maize. Ann. ae MEE ini. y of corn belt maize and its genetic significance. Heterosis. РР. 124-148. 1 Y. Gowen, Ed. Iowa State Coll. Press, Ames. "o. The history of the common але varieties of the United States corn bele Авт H EER RE ugh c. “Cuter (en. Races of Zea Mays: Ann. n dm Gard. 29 , ор ст Gage MT Introgressive hybridizati I. Their recognition and classification. on in Phlox bifida. Amer. Jour. Bot. 39:399—4 SEMIS and D po. кона (1944). А method for recording and analyzing variation in в internode pattern t. Gard. 31:241-247. "PE em BSS. (19 52). PME "No нт er e U.S. Thé а em mes d С. (1948). А resp raisal of Peruvian carin А Атег. Агс Eon Antiquity 13). 128 pp. í Bird, ns i 1943). oer! in оя Chile. Amer. Mus. Nat. Hist., Anthrop. Papers а 94): America’s oldest farmers. Nat. Hist. 57:296— ^ Bonnett, "dd T. (1940). Dept of the staminate je td inflorescences of sweet corn. Jour. Agr. Res. 60:2 ; ТГ» S MB. Ear 2 ыз development in maize. Апп. Мо. Bot. Gard. 35:269-287. [Vor. 40 110 ANNALS OF THE MISSOURI BOTANICAL GARDEN Brown, William L. (1949). Sene and distribution of chromosome knobs in United States maize. Genetics 34:524—53 and Edgar AER т. 1947). The Northern Flint o Ann. Mo. Bot. Gard. 34:1-28, — — ——, (1948). im Southern Dent corns Md e 255—268. А E: G. ARA, and R Lo Je. (1952). Ви on three varieties of Hopi maize. Amer. Jour. Bot. p 597—6 Buck, Peter Н. (1938). Vikings of the sunrise. 33 tokes Co., New York. Burgh, weis F., and Chas. R. Scoggin (1948). The d of = Park, Dut National Mon ment. Univ. Colo. Studies, Ser. in Anthrop. 102 p s Gate F. (1945). Plant geography and СҢ Aie in eg American Southwest. Viking und Pub 14 ork. nthrop., No. 5. ; (1946). Origins of CSS Di. pee ге. Ап гор. 48 ‚ (1949). Чуку: implications of chromosome A of Nock kes Indian maize. Southwest Jour. Anthrop. 5:199—207. ‚ (1950). Plant evidente for early contacts with America. Ibid. 6:161—18 ; and Edgar mno dea А pre ED ci survey of maize in the usas United Ann. Mo. 297— Cutler E а (1946), Ёле of maize in South America. Leafl. Bot. Mus. Harvard Univ. "8 (т The geographic origin of maize. Chron. Bot. 12:167—169. ( А preliminary соге = аван remains of Tularosa Cave. In Fieldiana: Anthro- pology ( chess Nat. Hist. Mus.) 4 Esau, АЯ (1943). Ontogeny of ха е bundle in Zea Mays. Hilgardia 15:327-356. Ford, a ; and Gordon Willey 1941). An аена ир е of the prehistory of the eastern United States. Amer. Anthrop. 43:325— neta т. (1939). Über die Organstellung bei M aiskolben. ас Jour. Bot. 10:113-140. Hau mil W. (1945). Painted Cave, northeastern Arizona. 7 pp. Amerind Foundation, Inc. jos n, Ariz. x 1950). The stratigraphy and еее of Ventana Cave, Arizona. Univ. of Ariz. Tucson; Univ. of New Mex Albuquer Hurst, C. T. (1948). The Cotto ied Cum expedition, 1947— cave and pueblo site. South- wes — — ———, and Edgar ‘Anderson (1949). A corn cache from western Colorado. Amer. Antiquity 14:1 161- Johnson, F. O. os). Radiocarbon dating. Mem. Soc. Amer. Arch., No. 8. (Suppl. Amer. An- tiquity 17). Jones, ier H. 19), Maize from the Davis Site: its nature and Run ceqpoaterion, In: Newell and r, The George C. Davis S MN scias County, Texas. . Soc. Amer. Arch., No. 5 e Т Amer. Antiquity 14) :239 tole Sheldon А 19 p Arroyos. s te Amer. 1879:7 Kiesselbach, . (1949). The structure and undue. of corn. Univ. Neb. Coll. Agr. Bull. А уз Kroeber, А. Je (1939). Cultural and natural areas of native North America. Univ. Calif. Publ. in Amer. Arch. and Ethnol., No. 3 Lenz, m Wayne Rs 8). d histology of the female inflorescence of Zea Mays L. Ann. t. Gard. 35:353—376. Yide, Ralph (1924). beg significance of certain traits in North American maize cultures. Amer. nthrop Lister, Robert H., and He rbert ыы ох (1952). Archaeology of the Glade Park area—a progress : S Mon of maize from North American Indians. Jour. Agric. Res. wie; К obert H. (1940). American Се" history. Amer. Anthrop. 42:409—428 m . Gard 3 en 196. The role of pod corn іп the or rigin and evolution с maize, Ann. ar 1377. — de noe The origin of Indian corn and its relatives. Texas Agr. Exp- Sta. Bull. 574. 315 ‚ and C. Earle Sith; Jr. (1949). мы archaeological evidence on evolution in maize . Bot. Mus . Harva Leal Univ. 13:213—24 Mattia, Раш $ ; John B Rinaldo, Elaine ME uS Hugh C. Cutler, and Roger Grange, Jr. os Mogollon cultural саол and change. Fieldiana: Anthr opology (Chicago Nat. Hist. Mus.) 40:1— — H s and Alex D. Krieger ( 1949). The Geor А rge C. Davis ах (dero County, Texas. m. Soc. Amer. Arch., No. 5. (S uppl. Amer. Antiquity 14). 256 Ў 1953] ; NICKERSON—VARIATION IN COB MORPHOLOGY 111 Nickerson, Norton H. (In press). Morphological analysis of the maize Prat, nt (1948). General features of the epidermis in Zea Mays. pet "мо. Bot. Gard. 35:341- Randolph, І. Е. (1952). New evidence on the origin of maize. Amer. Nat. 86:193—202, Richards, Annette H. (1953). The myst ed of the first Midian. Nat. Hist. 62:168—174. Sauer, Sek = мне ааа origins and dispersals. Bowman Memorial Lectures. Ser. II. New Yor fub. G. are erg kx (1952). “The кыр of cultivated plants and weeds, Evolution 6:445- Stonor, С. К., апа га Anderson (1949). Maize among the hill peoples of Assam. Ann. Mo. Bot. Gard. 36:355—4 ошл» E. L. (18 WI Varieties of corn, U. S. D r. Off. Exp. Sta, Bull, 57:1 Р and Preston sen СЭ А уча, poiius complex in de: ive dudo nited States. Amer. Ant 7:1— Vidua. Paul (1918). The evolution г maize. Bull. €— Bot. Club 45:309-342. — ———2, (1920). А misconception as to the structure of the of maize. Ibid. 47:359-362. ‚ (1936). The | ра of the maize poe and maize и in ancient America. Univ. New Mex., pte p. Ser. I, No. 5:11-18. Wellhausen, Е. J., L m rts, and E. нгаб X. (1951). Razas y^ maiz en México. Ofic. tud. E 299ге. 9 с" Agric, y iut. Foll. Tech. 5. 237 pp. Mexico ———, (1 952). Races of maize in Mexico. т М, Harvard Univ. Whicker, "t. W. (1948). Lees a pre-Columbian cultivated plant in the Americas. South- ern Jour. Anthrop. 4:49—6 id Junius Bird —— Identification and (à: of the cucurbit materials from a Prieta, Peru. us. t. 1426. 15 New York, Whine Alfred F. ur hs ОНЫН a corn: n evaluation А p and theory. Amer. Anthrop. Wiser, Cark, Саму Corn and early American civilization. Nat. Hist. 54:5 Worm fà: M. (1949). Ancient man in North America. Denver Mus. КС Не, Рор. $ег. vus i" THE CYTOLOGY, MORPHOLOGY, AND SYSTEMATIC RELATIONSHIPS OF DELPHINIUM x; BELLADONNA HORT. EX BERGM.* MARILYN AMY GAGE I INTRODUCTION Among the cultivated perennial Delphiniums the problem of the origin of the forms variously termed "garland larkspurs" (Bailey, 1930) or Belladonna types has been raised more than once. The group as it is known at present includes not only Delphinium X Belladonna Hort. ex Bergm., but at least the 45 variants listed by the Royal Horticultural Society (1949), and probably certain other hybrids of unknown origin some of which are no longer in cultivation but were recorded in the literature of the last century. Two main theories have been proposed regarding the nature of the assemblage. (1) L. H. Bailey, following the example of Huth (1895), the last monographer of the genus, considered these plants to be forms of D. cheilanthum Fisch. var. formosum; his grounds are wholly morphological (2) Lawrence (1936), on the other hand, has suggested that D. Belladomna originated in gardens as a triploid hybrid between a tetraploid species and a diploid species, and that subsequent doubling of the chromosome number resulted in the fertile line now in cultiva- tion. His hypothesis is in line with the known facts, for until early in the present century Belladonna was quite sterile and was propagated wholly vegetatively; moreover, the present fertile strain is a hexaploid. In studying the origin of cultigens the historical method alone is obviously in- adequate. Students of cultivated plants are well acquainted with the hazards of attempting to place even well-known species into conventional taxonomic cate- gories. Even with such a relatively recent variety as D. Belladonna, caution is necessary in considering historical references, for it often happens that opinions given in early descriptions are contradictory or suggest some parentage which, in the light of present knowledge of cytology and genetics, is far from the truth. One can accept, with caution, species listed in cultivation at a certain period, but there must always be the reservation that morphologically similar species are sub- ject to confusion, and that any of these species may, and probably do, differ from their wild relatives as a result of horticultural “amelioration.” Morphological comparisons with known species have often been used in estab- lishing the possible parentage of plants of unknown origin. This method is most useful in groups where there are clear-cut characters whose extreme and inter- mediate expressions are easily recognized. The genus Delphinium is, however, notoriously variable, and specific distinctions are based to a very great extent upon quantitative differences, such as width of leaf segments, degree of dissection of чел ap i ; haw School of Botany «t al tt Ter erg ا‎ варце. ко of the requirements for the degree of Doctor of Philosophy. During the last year the study was carried out under a pre- doctoral fellowship of the National Science Foundation. (113) [Vor. 40 114 ANNALS OF THE MISSOURI BOTANICAL GARDEN the leaves, extent and type of axial branching, degree of incision of the "bee", length of spur, and the like. Every one of these characters is subject not only to genetical, but also, to a high degree, to environmental variability. Therefore, before employing such characters, it is important to understand to what extent each character may be modified by the environment. A third method of general use in establishing relationships between species, that of cytological analysis, is often more reliable. А number of investigators have shown that degree of relationship between species may often be estimated by studying the morphology and especially the pairing affinities of the chromosomes (see Stebbins, 1950, for a review of the literature). However, where the chromo- somes of the parental genomes are not strongly differentiated there must often remain doubt as to the sources of the chromosomes which are pairing in the hybrid. Especially in polyploids above the tetraploid level such problems are intensified by the presence in the same individual of several genomes, so that autopolyploidy and allopolyploidy may coexist. The relationships of D. Belladonna must therefore be considered from a com- bination of several standpoints, and the solution must be based upon that hypothesis which best makes use of the information obtained by each method. Of the materials employed in this study, families of various species and hybrids were grown in the experimental greenhouses of the Missouri Botanical Garden and of Washington University, in St. Louis. Herbarium specimens were made available by the Bailey Hortorium of Cornell University and from the collections of the Missouri Botanical Garden. I wish to acknowledge my indebtedness to Dr. G. A. L. Mehl quist, of the University of Connecticut, formerly Research Horticulturist at the Missouri Botanical Garden, who suggested this problem and who stimulated my interest in it by his own enthusiasm. I am also indebted to Dr. Edgar Anderson and other members of the staff of the Missouri Botanical Garden who have aided me in many ways. IL. TAXONOMY AND MORPHOLOGY A. NATURAL SPECIES History of the Genus.— Since the establishment of the genus in its modern form Бу Tournefort (1700), Delpbinium has received various treatments. In his 'Species Plantarum' Linnaeus listed the six species, D. consolida, D. ajacis, D. peregrinum, D. grandiflorum, D. elatum, and D. Staphisagria. By the close of the eighteenth century only six more species had been added, but in the century and a half since then, the number of species described has increased to more than 300. A. P. DeCandolle was the author of 23 species, and in his ‘Prodromus (1824) he grouped 53 species into the four sections, CONSOLIDA, DELPHINELLUM, DELPHINASTRUM, and STAPHISAGRIA. Prantl, in Engler and Prantl's *Die Natürlichen Pflanzenfamilien’ (1888) merged DeCandolle's srAPHIsAGRIA and DELPHINASTRUM into one section, STAPHISAGRIA; 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. 115 and later, modifications were made by others—notably Asa Gray (1887), Robin- son and Fernald (1908), and Huth (1895). The most recent treatment of the American species is that of Ewan (1945). Huth published in 1895 his "Monographie der Gattung Delphinium,” in which he attempted a natural classification of the 200 odd species treated. Не recognizes two subgenera: CONSOLIDA, in which there is a single carpel, the petals are fused into one, and seeds are scaled and three-cornered; and EUDELPHINIUM, in which there is more than one carpel and the four petals are free. Within the second sub- genus he distinguishes three sections: ELATOPsis—petals deep violet or black, lower petals bifid and bearded. DIEDROPETALA—petals light, the same color as the sepals or dirty yellow, deeply ifid, lobes acute. KOLOBOPETALA—color of petals as in DIEDROPETALA; limb of lower petals round or rectangular, entire or bilobed, lobes round or truncate at tip. Of the species considered here, D. elatum L. belongs to the series КАСЕМОЗА of section ELATOPSIS, and D. grandiflorum L., D. cheilanthum Fisch. ex DC., and D. tatsienense Franch. are included in the series CHEILANTHOIDEA of section KOLOBOPETALA. While D. Belladonna Hort. was not known to Huth, he in- cluded as varieties of D. cheilanthum several cultigens which are certainly very similar to it. Further consideration of the taxonomic categories will be restricted to D. cheilanthum, since it alone has been confused with D. Belladonna. Тһе other species (D. elatum, D. grandiflorum, and D. tatsienense) are taxonomically well- known, and while they include many natural and horticultural variants, there is no problem in determining the affinity of such variants. Delphinium cheilanthum Fisch. ex DC.— Apart from the question of any direct relationship between D. Belladonna and D. cheilanthum, a problem which will be discussed later, the two species are con- used in herbaria and in gardens, where identifications must be made largely upon a morphological basis. The magnitude of the problem is amply brought out by L. H. Bailey in ‘The Garden of Larkspurs’ (1939), in which he attempts to trace the history of D. Belladonna as a derivative of D. cheilanthum. He bases his con- clusions upon a comparison of certain historic plates and descriptions of D. cheil- anthum and of certain horticultural varieties (D. formosum Hort. and D. Hender- soni Hort.). The last two will be taken up in the next section, and I shall discuss here only D. cheilanthum in order to clarify its position, since it is apparently infrequent in herbaria in this country. Delphinium cheilanthum Fisch. ex DC. is based upon a specimen sent to A. P. De Candolle (1818) by Fischer from the region of Doroninsk in Dahuria (in тч терт 8А „Йи: hat cheilanthum does not appear as a native plant in the great А ыгы is ek I "aee эе ана Сы шей cited by Huth mostly are plants considered by him to be botanical varieties of D. cheilanthum.” [Vor. 40 116 ANNALS OF THE MISSOURI BOTANICAL GARDEN trans-Baikal Siberia). In 1819 von Schrank figured and described the same plant (under the name of D. cheilanthes) in ‘Plantae Rariores Horti Academici Mona- censis’ (pl. 7). Earlier, іп 1769, J. С. Gmelin had described and illustrated in his ‘Flora Sibirica’ a plant which, according to De Candolle, Fischer had indicated was the same as his D. cheilanthum. In 1820 a plant grown near London from seed sent by Fischer was figured in the ‘Botanical Register.’ In addition to the sources cited by Bailey, a number of accounts of the species given by collectors in Siberia and northern China (Ledebour, 1842; Regel, 1861; Trautvetter, 1847, 1877а, 1877b; Turczaninow, 1842; Glehn, 1876; Brühl, 1896; and Komarov and Schischkin, 1937) delimit D. cheilanthum Fisch. as a species or species complex, with well-defined characters and a distinct geographic distribu- tion. From these accounts, and the Huth monograph, the species may be character- ized as follows: DELPHINIUM CHEILANTHUM Fisch., in DC. Prodr. 1:53. 1824.—Stem tall, simple or branched; leaves glabrous or pubescent (especially on the under surfaces); 5-parted, parts oblong to narrowly lanceolate-acuminate; lower bracts many- parted. Inflorescence racemose to subcorymbose, peduncles bibracteolate; upper bractlets equal to spur. Flowers blue, rarely white; spur straight or slightly curved; sepals ovate, pubescent or puberulent externally; upper petals glabrous, pale yellow or blue, lower petals large, with ovate or subrotund limb, entire or rarely emarginate-bilobed, with yellow pubescence above. Carpels 3, glabrous or pubescent, tips curved at maturity, to 20—25 mm. long; seeds 3-angled, with winged margins, faces scarcely squamate. Flowers in June and July (pl. 9). A number of varieties have been described, but undoubtedly any future monographer must rework the alliance. The exact limits of the species are still somewhat in question, since Huth describes Delphinium Middendorffii Trautv. as a distinct species, although in his natural key to the genus he lists it as а variety of D. cbeilantbum, distinguished by having the apices of the sepals lanceolate. Moreover, he notes: “D. Middendorffii scheint eine verkümmerte Form des Dd- phinium cbeilantbum mit verlängerten Kelchblattern zu sein." By 1877 Traut- vetter, who originally described D. Middendorffii (1847), conceded that it was but | another variety of cheilanthum, and added (translation) : with бы, Tli tects рона konali «low wem (oet Middendorfi) eid rome times a taller stem; and this latter form has flowers either larger (var. typica) or smaller | Ая parviflora). Indeed, sometimes var. Middendorffi has larger, and sometim imes smaller flowers, so that sometimes it approaches var. Руріса and sometimes var . parviflora. An American species, D. chamissonis Pritz. (from the Bering Sea region and the Yukon valley) is also apparently a close ally of D. cheilanthum. Comparing D. chamissonis and D. Middendorffii, Ewan (1945) says: “All of the more than 4 score of collections examined show so much variation in spur length and, indeed, flower size as a whole, as to render it difficult to draw any distinction between two species from these characters." 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. li7 B. HYBRIDS OF THE BELLADONNA TYPE The history of the cultivated Delphiniums is extremely complex (see Wilde, 1931; Bailey, 1939). As early as 1778 D. elatum L. and another form listed as D. azureum (not D. azureum Michx., which was not described until 1803) were offered in the trade, and at least by 1824 D. grandiflorum L. (two varieties) was offered. Other species were rapidly introduced and by the middle of the century hybridization was being carried on by a number of French growers, chief among whom was Victor Lemoine, originator of most of the first varieties of perennial garden Delphiniums. English plantsmen were somewhat later in the field, but particularly since the 1870’s, when the first varieties of James Kelway were intro- duced, a large part of the development of new hybrids has been carried out in Britain. As nearly as can be ascertained, most of the present forms were derived from relatively few species. D. grandiflorum and D. elatum have chiefly been involved, although hybrids with other species, including D. tatsienense Franch. (Lemoine, 1914), D. cardinale Hook. (Wilde, 1931), and D. mudicaule Torr. and Gray (Lawrence, 1936) have enjoyed some popularity. Horticultural literature has frequently not been reliable regarding species in cultivation, and allies of D. grandiflorum or D. elatum could have been confused with these species and so have been employed in crosses at a time antedating any existing records of their use. As has been mentioned above, D. Belladonna first appeared some time after the middle of the last century; the earliest known mention of its name is in 'Flori- cultural Cabinet and Florist’s Magazine’ for 1857, where it is listed as а desirable form with large pale lavender-blue flowers. In the 1865 catalogue of James Backhouse & Son, of York, England, Belladonna is listed and described as "a lovely turquoise, perfectly hardy." There are sporadic references to the plant in various horticultural publications before 1880, in which year the English firm of Kelway introduced it to a wide market. For a long time Belladonna was reported to be sterile. However, in 1902 or 1903 a plant grown by G. Gibson, of Leeming Bar, yielded three pods, whose seeds produced five plants, two of which became the named varieties “Mrs. С. Gibson," and "Grandiflora" (Phillips, 1949); and in 1905 Sutton & Sons, of Reading, England, obtained seed from two plants. Thus it appears that a chromosome-doubling occurred at least twice within a few year's time. The fertile strain has always been true-breeding, in the sense that there has been no recovery of anything which resembles any diploid or tetraploid species Which might be assumed to have been the original parents (although it will be demonstrated later that weak but measurable associations of characters present in such species are still to be found). It it true, nonetheless, that heterozygosity in these original amphidiploid forms must have been fairly high, judging from the large number of Belladonna varieties introduced within the next few years. It is of interest that the ‘Journal of the Royal Horticultural Society’ (1907) says of Sutton's first seedlings: "Some . . . resembled Delphinium Belladonna very closely, but some more nearly approached Delphinium formosum, and one bore [Vor. 40 118 ANNALS OF THE MISSOURI BOTANICAL GARDEN flowers of a very beautiful deep blue tint." D. formosum referred to here is the horticultural form and not the species of Boissier and Huet. Like Belladonna, it was a garden hybrid of unknown origin, and, also like Belladonna, it is said to be a hexaploid (Propach, 1939, 1940; Mehlquist, unpublished). It was first offered in 1855, and according to Van Houtte (1857) it was already fertile. This, coupled with its superficial resemblance to the diploid species D. cheilanthum, caused Huth to list it as a variety of that species in his monograph, although it has never been reported in nature (pl 8). It is suggested that formosum may have originated from an early and unrecorded doubling of the chromosome complement of Bella- he We Bee Se se ми. at at t-fig. 1. Leaf types im Delphinium species and hybrids (X и). D. чл аы var. chinensis Blue Butterfly. D. — var. chinensis 20 ег х м ^ D. tatsiene D. cbeilantbum ‚ from toners псн Ьу Каго in Dahur D. бейин. from plant grown from seed at the Macer Botanical Garden. 6-9. D. elatum рч range a vs Variation. 10-15. Belladonna types: 10, Lamartine; 11, Smith's Belladonna 16-20. Experimentally produced triploids: 16, 50-15-3; 17, 50-40-2; 18, 48-27-4; 19, 48-27-63 1953] GAGE—DELPHINIUM X BELLADONNA HORT. 119 donna, or at any rate that it originated in the same manner as Belladonna. Cer- tainly, there are no consistent differences between the two. The following is a general description of D. Belladonna: DELPHINIUM X BELLADONNA Hort. ex Bergm.?—Racemose open-flowered perennial with simple or branched stem up to 1.5-2.0 m. tall; herbage pubescent or glabrous; leaves palmate or deeply 3-parted and strongly ribbed, the parts again divided or lobed, the degree of dissection of the parts gradually increasing from rosette leaves to bracts, the main division of the stem leaves narrow-cuneate to narrow-oblong, 0.2—2.0 cm. broad at base, the petioles nearly or quite as long as the blade. Inflorescences of relatively short and open (5- to 20-flowered) racemes, the peduncles with parted bracts at base and the pedicels usually long, with basal simple bracts and paired short bracteoles immediately subtending the flowers. Flowers large, single, occasionally semi-double, mostly light to dark blue and somewhat declined; sepals thinly pubescent outside, blunt; spur about equalling the sepals and straight or somewhat curved at the end; petals mostly white or light-colored, bearded with yellow, making a large "bee" which fills the throat of the flower. Follicles 3, pubescent or glabrous, up to 2.0 cm. long, the apex curved; seeds 3-angled with somewhat winged margins, not squamate. Flowering period prolonged by the production of successive shoots during the growing season (pl. 8). Another variety of interest, Lamartine, originated with the French house of Lemoine & Son in 1903. Like Belladonna, it was at first sterile, and the present fertile strain had its origin in 1916-17 when doubling of the chromosome number apparently occurred among some plants growing at the Royal Moerheim Nurseries in Holland (Lawrence, 1936). The parentage of this variety is also obscure. In answer to my inquiry, the present head of the firm replied that they had no record of its antecedents, but that experimental work had suggested that the parents might have been an elatum type (sic, Pacific Giant) and D. grandiflorum. In habit, Lamartine is quite similar to Belladonna, being of the same height, with a loose, tapering spike, and fairly numerous lateral branch spikes. The leaves are similarly cut (text-fig. 1). The flowers are long-spurred, with sepals ultramarine suffused with purple, and the petals are dark-edged, shading to white, with a yellow beard. Like Belladonna, it has an extended blooming period. There are also grown at present two triploid varieties, Moerbeimi and Capri, which, while not directly related to Belladonna, have a close morphological simi- larity to that variety. The most complete and authoritative account of triploids is that given by B. Ruys of the Royal Moerheim Nurseries (1911): *D. Belladonna Hort. is equivalent to D. Belladonna Bergm., listed in ‘Index кин. Me mans, who published his binomial in 1925, is among those who believe the plant to have originat from a cross between elatum and cheilanthum. Не lists no sources for his opinion. [Уот. 40 120 ANNALS OF THE MISSOURI BOTANICAL GARDEN all the light-colored novelties which I could obtain, with a view to crossing them with large- ligh str itution . .. ew years ago] I found in a batch of seedlings one plant with five spikes. Of these, two spikes bore pure white flowers, two bore blue flowers, and some parti-colored, half-blue, half-white flowers. Next year, when the five divided plants flowered, I noticed that two had only white flowers, two only blue flowers, and one plant had some flowers white and others blue, whilst still others were half white and half blue." Both the white and the blue variety were propagated and marketed, the former under the name of Moerbeimi, and the latter as Capri. Aside from the color, there is no single definitive character to separate these and Belladonna, although, as mentioned above, these varieties are triploid. In addition to the present-day varieties of the Belladonna type, a study of the literature reveals that other hybrids of similar appearance were known in the last century, though they have now passed from cultivation. Because of their bearing upon the possible constitution of D. Belladonna, some of these early hybrids are described below: The earliest recorded Delpbinium hybrid was Barlowii, figured by Lindley in the “Botanical Register’ of 1837. The plant, from a nursery in Tooting, had been received from an establishment in Manchester, and was believed to be the result of a chance cross between D. grandiflorum L. and D. elatum L. It was reported to be a continuous bloomer. ‘Paxton’s Magazine’ described it the following year: “Plant perennial, growing usually from 4—5 feet high. Leaves with 5 principal divisions, deeply lobed and jagged; serrations acute; deep green on surface, light green beneath, smooth. Flowers semi-double, of a most intense blue color, produced very numerously in terminal spikes. Sepals of the calyx greenish externally.” Neither the description nor the Lindley plate, which shows a rather dense spike of semi-double blooms and, in the background, part of a leaf in outline, is com- plete enough to indicate the affinities of this plant. Moreover, except that the flowers are semi-double in each case, the Lindley plate is quite different from the Paxton plate, which indicates that the plant had a brownish bee, and leaves more deeply incised than those in the earlier plate. Since both plates were drawn from living plants, one is confronted with the alternatives that Barlowii was a highly variable variety, or that the two artists had available two quite different varieties. The plant in the earlier illustration might easily belong to the group of hybrids which includes Belladonna; the later plate appears, rather, to be merely that of an elatum variety. Herbarium specimens of plants sold as Barlowii within the last 20 years or so are quite different from either of these, being single dark-blue types, and not different morphologically from D. Belladonna. The next form of interest, D. Hendersonii Hort. (pl. 7) is of extremely dubious background. It was first mentioned by Moore and Ayres in the ‘Gardeners’ Magazine of Botany’ (1850) and was described under the name D. cheilanthum var. Hendersonii A. Henfr.: 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. 121 “Hardy herbaceous perennial with large showy rather distant flowers. Leaves five- parted, lobes oblong or acuminated, trifid or obscurely bi-trifid, 4 in diameter, on long cemes inches el broadly linear, acuminated, simple. Racem axillary and terminal, lax; the pedicels larger than the Flowers large and showy, ns a little deeper; two lower petals with a roundish ovate limb, whitish in the middle, bearing a yellow beard, slightly irregular on * the margins, with a few ciliary hairs at the apex. Ovaries 3, glabrous, green, veined with h lines. Wood, from seedlings purchased from M. Chauviére of Paris var. Cbauvieri. Lemaire regarded it as merely a variant of the species, whose dis- tribution and habitat he considered briefly. ite in opposition to Lemaire, Harrison (1853), describing the same plant, stated: “It was raised by M. Chauviére, nurseryman of Paris, from D. chinense, impregnated by D. elatum-splendens.” This author also noted that Hendersonii was a profuse bloomer, of bushy habit, flowering over a long period of time. Hybrids of known parentage.— During the past eight years there have been produced at the Missouri Botanical Garden a number of hybrid Delphiniums of particular interest. Several hundre pollinations were made between D. elatum and different strains of D. cheilanthum and D. grandiflorum var. chinense, in an effort to find out which, if either, of these hybrids more closely resembled the Belladonna assemblage. Up to this time there have been no successful pollinations of D. elatum by D. cheilanthum. How- ever, the other cross (that between elatum and grandiflorum) has been successful four times, yielding two families of 3 plants each, one family of 2 plants, and one family of 13 plants. 1. 47-98-3 X D. grandiflorum var. chinense “Blue Butterfly”.—Hybrid Nos. 48-27-1 through 13. (This and the following cross were made by Dr. С. A. L. Mehlquist.) The seed parent was one of a line of white semi-double elatum forms developed by Dr. G. A. L. Mehlquist. The group is exceptional in being the only line of Whites in which there has been an approach to homozygosity without concomitant Weakening and loss of the line. 47-98-3 had dense spikes borne on the rather short main axes, and leaves of medium size, pubescent, and palmately divided, with relatively narrow segments ( pl 10). The pollen parent, on the other hand, was a low-growing plant with a branched, very open inflorescence of fairly large rich deep blue single flowers in few-flowered panicles. The leaves were relatively small, dull green, glabrous, and finely dissected. This form, too, was true-breeding, and the variety has been grown at least since 1910 when it was among the perennials planted at Wisley (Jour. Roy. Hort. Soc., 1910) (pl 9 and text-fig. 1). The Progeny of this cross included 13 individuals which were generally of intermediate bit and varied very little among themselves. The following description serves to characterize them and point out their chief differences from the parents. [Vor. 40 122 ANNALS OF THE MISSOURI BOTANICAL GARDEN SUMMER SKIES X 47031-1 50-15-4 X SMITH'S BELLADONNA ا ~. - SMITH. X se к X SELF 51- 213-2 X SELF изз X SELF 51-217 51-216 51-218 52-224 48-2-3 X BLUE BUTTERFLY 41-98-3 X BLUE BUTTERFLY 48-6-3 X SMITH'S BELLADONNA 48-27-5 X SMITH'S BELLADONNA SMITH X 50-20-1 X SELF SMITH X 51-209-1 X SELF 51-201-1,2 T 202- Y TO 14 51-223 51-222 SMITH х 51-202-5 X SELF 51-202-1 X 50-20-1 (51-220 51-219 51-221 Text-fig. 2. Pedigrees of major lines of triploids and hexaploids. 48-27- Series.—Racemose open-flowered perennials, with simple or usually branched stem up to 1.0—1.25 meters tall; herbage pubescent. Leaves palmate and deeply 5- or 3-parted and strongly ribbed, the parts again divided or lobed, the degree of dissection increasing progressively from the rosette to the bracts, the main divisions chiefly narrow-cuneate, up to about 1.0 cm. broad, the petioles 45 long as or slightly longer than the blade. Inflorescences not spiciform, the terminal one a 15- to 20-flowered raceme, the lateral of fewer flowers, the peduncles with 3-partite bracts at base and the pedicels with basal simple bracts, and pair bracteoles 0.8—1.0 cm. long subtending the flower. Flowers medium-sized, single, violet and cobalt, on long pedicels and somewhat declined; sepals thinly pubescent outside, blunt, the spur slightly longer than the limb and usually somewhat curved at the end; petals the same color as the sepals, bearded with short golden yellow hairs. Follicles 3, pubescent, not developing into fruits, apex curved. Flowering period prolonged by the production of successive shoots during the growing season (pl. 11 and text-fig. 1). 2. 48-2-3 X Blue Butterfly.—Hybrid Nos. 48-6-1 through 48-6-3. 48-2-3, the seed parent of this group, was a selection from Vetterle and Reinelt's 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. 123 Galahad Series of their Pacific Giant Hybrids. The plant, a white semi-double, was not available to me. The three hybrid individuals in this group were quite similar to the hybrids described above, the chief differences being the regular production of one or several additional petals of either sort, and the generally larger, less open flowers. 3. Summer Skies X D. grandiflorum (47031-1).—Hybrid Nos. 50-15-1,-3,-4. The seed parent of this cross is a selected seedling from the Pacific Giant series, Summer Skies, an elatum type with semi-double flowers with sky-blue sepals and white petals in tapering, somewhat dense spikes (pl. 12). It has been found to be true-breeding for color (Mehlquist, unpublished). The pollen parent was from seed obtained from the Royal Botanic Gardens at Kew as D. tatsienense. How- ever, determination of the plant, using Huth's key, showed that it must have been, rather, a corymbose variety of D. grandiflorum. Numerous attempts were made to obtain seed, but 47031-1 was completely seed-sterile. The three hybrid individuals of this cross differ in a number of particulars from those of the other crosses, being taller, with a strong main axis which bears lateral spikes in the manner of D. elatum, but these more profuse and more strongly developed than in that species. Leaves of 50-15-3 are much like those of e 48-27- series, whereas those of the other two plants are considerably larger and have the ultimate divisions rounded rather than acute. The flowers are again mixed blue and violet (although the blue of these plants is somewhat more brilliant than in the other crosses) and are more or less semi-double, with spurs about № longer than the standard of the sepal (pl. 12, and text-fig. 1). 4. 47-98-3 X 47031-I.—Hybrid Nos. 50-40-1 and 2. The seed parent was the white elatum used in producing the triploids of line 48-27; the pollen parent was also used to produce the triploids of line 50-15. The two individuals of this cross resembled those of 48-27, except that the leaves were more finely dissected, and the flower was the same brilliant cobalt noted in the 50-15 individuals (text-fig. 1). 5. Derived bexaploid bybrids.— Since the fertile strain of D. Belladonna apparently originated from spikes Whose chromosome numbers had been fortuitously doubled, it was hoped that it might be possible to produce doubling of the chromosomes of the hybrids described above. With this in view the developing rosettes were treated each year for three Years with aqueous colchicine in various concentrations for varying lengths of time. In at least one case, doubling of the somatic chromosome number of the Spike was achieved, judging not only from morphological features but from greatly increased amounts of "viable" pollen produced (Table I), but the 14 seeds ob- tained did not germinate. The other method employed, that of repeated selfing сыны hether delayed planting or some undetermi genetic factors may have been wn whether delayed planting | е ` anger ын ш Gi de Н Lac ТЫР hans ры may not germinate at all after six months, even in cold storage. [Vor. 40 124 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE I POLLEN SIZE AND VIABILITY IN SPECIES AND HYBRIDS OF DELPHINIUM: LIVING PLANTS. s Pollen diameter in д Plant % bius ble | S | Range Median Diploids D. grandiflorum Blue Butterfly 97.0 18.8—37.6 233 47031-1 65.0 18.8—28.2 23.5 Tom Thumb 91.0 23.5—37.5 28.2 D. cheilanthum 98.0 23.5—32.9 28.2 Tetraploids 47-98-3 64.6 23.5—37.5 30.6 Summer Skies 32.0 23.5—37.6 30.6 Triploids 48-6-1 p Sr d 28.2—37.5 32.9 48-6-2 7.0 23.5—40.0 32.9 48-27-1 16.0 25.9—40.0 30.6 48-27-3 14.0 23.5—37.6 324.9 48-27 21.2—47.0 33,3 48-27-5 21.0 23.5—42, 32:9 48-27-6 E 23.5—37.6 35.3 48-27-7 9.0 23.5—36.5 Е 48-27-8 4.0 23.5—42.3 2 48-27-10 16.3 21.2—51.7 37.6 48-27-11 5.0 23.5—37 0. 48-27-12 23 23.5—47.0 32.9 50-15-1 1:5 50-15-3 0.0 50-15-4 2,5 5 - treated) 2.0 21.2—37.6 30.6 50-40-2 (treated) 52.0 25.9-37.6 30.6 Moerheimi 0.5 те = ЫЕ Hexaploids OERE E E EE Smith's Belladonna 41.0—95.0 23.5—42.5 32.9 Cliveden Beauty -1 68.2 £ Б 37.6 -5 1 80.5 25.9—37.6 32:9 Lamartine 82.5 23.5—32.9 30.6 ел of the triploids, proved even less satisfactory, only two inviable seeds being produced from upwards of a thousand individual pollinations made in several seasons. However, pollinations of the triploids by Belladonna proved more suc- cessful: eventually ten plants, representing five crosses, were obtained, and of these, five plants (representing three crosses) were used in establishing new hexa- ploid lines, both by selfing and by backcrossing to Belladonna. A detailed descrip- . tion of the cytology of these hybrids follows (chapter IV), but it is to be no that in morphological characters and cytological details they do not deviate from 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. 125 the Belladonna category except in being more variable; and in some cases fertility is already as high as in Belladonna. C. DISCUSSION Two major points are to be brought out here: First, while it has been demon- strated that there is indeed a resemblance between the Asiatic diploid species D. cheilanthum Fisch. ex DC. and the hexaploid cultigen, D. Belladonna, there are also important differences. The botanical species and its allies are restricted to Siberia, Mongolia, and North China, and to the islands of the Bering Sea and the Yukon, and it has not often been planted in gardens. Moreover, when one comes to compare the two entities, the first impression of similarity is modified by an appreciation of their subtle differences. Thus, the leaf of D. cheilanthum may often be incised so that the major divisions are of about the same width and degree of segmentation as in the Belladonnas, yet the latter will invariably have the ultimate divisions more acuminate, the segments often intricate, and the main lateral veins divergent in their departure, features rarely present in D. cheilanthum, and certainly never present in combination in that species. The flowers of D. cheilanthum are small but always have an extremely conspicuous bee; while the bee of Belladonna may be of the same absolute size, it is always smaller in propor- tion to the size of the flower. Lastly (and it is realized that the botanist will not often have access to the quantity of material necessary to make this distinction), if one has on hand a number of plants of both sorts, it is a relatively simple matter to separate the Belladonnas from D. cheilanthum, since the latter holds together by virtue of concordant variation, and the former is less cohesive on account of discordant variation (Anderson, 1951). The second point brought out in this section is that Belladonna types have originated a number of times in the past, and always they have first been noted in gardens. From the evidence, it seems clear that Hendersoni, Lamartine, Moer- beimi, and the others, while not directly related, are quite similar in leaf shape, flowers, branching habit, and prolonged blooming periods. Several of them re- produce by seeds; others have been completely sterile. In no case has the exact origin of the plant been unquestioned. However, all of them have been found to be very much like certain hybrids produced by crossing Del phinium elatum and D. grandiflorum varieties. In the absence of direct evidence one cannot 1mme- diately exclude D. cheilanthum as one of the parents, but evidence against such a supposition will be given in the following pages. Ш. CHARACTER ASSOCIATION AS AN AID IN EsTABLISHING RELATIONSHIPS _ At least since the time of Gmelin, botanists faced with the necessity of classify- ing species of Delphinium have realized the difficulty of finding reliable specific characters. This is the result not so much of lack of genetic differentiation within the genus, as of the presence of numerous characters of a quantitative nature, [Vor. 40 126 ANNALS OF THE MISSOURI BOTANICAL GARDEN whose genetic basis may be modified or simulated by environmental factors. On account of this "inability to use one or two differentiae to distinguish species or subspecies," Ewan (1945) has based his treatment of the North American species on "maximum correlation of characters existing in combinations." A species characterized in such terms could be visualized, then, as a plexus of "typical" individuals, possessing the most pronounced correlation of the characters of the species, and a large number of individuals possessing these characters in lesser degree, but with no sharp discontinuities within the group. Such discontinuities, on the other hand, would be expected between species; and populations of hybri origin might be expected to be largely coherent, but with some segregation if the parental genomes have sufficient segments in common that occasional heterogenetic pairing occurs (chap. IV). The study of hybrid ancestry in such a group as this would not long ago have been considered a very difficult one from which to arrive at any very definite conclusions. However, the student of variation has been given a valuable tool in the graphic methods developed by Anderson (Anderson, 1948, '51, '52; Anderson and Gage, 1952), which permit the study of complex patterns of variation on the two-dimensional level of the scatter diagram and ideograph. Аз Stebbins (1952) has recently pointed out, the method of extrapolated correlates involves more than the mere random choice of characters; rather, the choice of a measure is determined by how well it expresses a feature of the plant, and how subject that feature is to environmental variation. While the methods are thus a compromise between pure statistical analysis and subjective intuition, their success in a number of cases indicates their great usefulness. Before employing these methods in Delphinium, it was necessary to study | thoroughly within-a-plant variation. Good material for such a study was avail- able in pressed material, and in living plants of several vegetatively propagated individuals, chiefly hybrids of the 48-27 and 50-40 series and Smith’s Belladonna. Preliminary measurements of a number of characters demonstrated what was already suspected, that only a few features of the leaf could be used. (Leaf variation in Delphinium has been the subject of various investigations: Brown, 1944; Lewis, 1947; Ashby, 1948.) Ву trial and error it was eventually possible to score a number of characters which exhibited low variation even on divisions of a plant grown in different years. In addition to the few leaf characters, those characters most effective for measurement were those of the flowers and inflore- - cence. (Certain definitive characters, such as internode pattern, mode of branch- ing, etc. could not be employed because of the scanty amount of herbarium material) With these measurable characters as guides, the three natural specie (D. elatum, D. grandiflorum, and D. cheilanthum) and Belladonna were studi and those features which could be used alone or in combinations as differential characters were selected for use in separating the units on а multiple-charactet basis. The characters used were the following: 1953] GAGE—DELPHINIUM X BELLADONNA HORT. 127 W Text-fig. 3. Floral and leaf characters used in constructing scatter diagrams. Leaf.— Length of median segment/length of C (text-fig. 3). à; аге about equally dependent on absolute size of leaf, their ratio expresses fairly well the depth of cutting of the segments. Width of median segment.— This was found to be the best measure expressing what may degree of dissection of the segments. Inflore М TECE Number of flowers. The absolute number of flowers on the main axis moe ari Density. This was computed as the length of the inflorescence di noe (measured from the attachment of the lowest flower on the main axis) divided Ы; the number of flowers. Margin of Бее. Scored in three grades: bifid, notched, and entire or emarginate. Length of standard divided by length of limb of the bee (text-fig. 3). Since both measures be termed the [Vor. 40 128 ANNALS OF THE MISSOURI BOTANICAL GARDEN In laying out the scatter diagram (text-fig. 4), the ratios used as ordinate and abscissa were selected because, varying widely among the plants measured and expressed as individual grades, they permit a wide scattering of the points on the diagram. Moreover, the use of measures of one floral character and one leaf char- acter lessens the possibility that they might be merely different manifestations of the same factor. The scoring of the other characters was then adjusted so that each could be expressed in three grades, to which ray lengths and positions were assigned (further details accompany text-fig. 5). The "population" represented by the scatter diagram consists of herbarium specimens from the Missouri Botanical Garden and from the large collection of cultivated Delphiniums at the Bailey Hortorium of Cornell University. Extreme types such as double varieties and dwarf forms were excluded. АП other measur- able sheets were included in making the diagrams. Since pollen measurements made from herbarium specimens were found to be reliable indicators of approximate polyploid level (chap. IV), an additional measure, determined by pollen grain size, is expressed by the size of the dots on the scatter- diagram, the smallest dots representing diploids, the intermediate dots, tetraploids, and the large dots, hexaploids. A survey of the individuals thus diagrammed shows two distinct entities, D. grandiflorum in the lower left-hand corner, and D. elatum in the upper right- hand corner. Occupying the area between these species is the Belladonna complex, more variable than either but generally intermediate between the diploid and the tetraploid species. The position of D. cheilanthum just off the spindle but adjacent to Belladonna points out its notorious superficial similarity to Belladonna, but its actual physical distinctness is shown first by its diploid nature, and second by its almost "all or none" character with respect to the rays. It is seen to be not ай intermediate between grandiflorum and elatum, but, rather, it possesses some char- acters in common with grandiflorum and some with elatum. (The plants of cheilanthum scored included two collections of two plants each, as well as tw? plants growing in the greenhouse at the Missouri Botanical Garden, and so shows more grouping of the dots than would a random selection.) A graphic analysis of another sort, the internode diagram (text-fig. 6); points out clearly the differences in habit of the entities studied. Both D. elatum and D. cheilanthum have a relatively large number of nodes before the axis begins ® branch; these increase somewhat in length from the highly shortened internodes of the rosette (not illustrated) but do not increase in a regular pattern and afè generally not more than 7—8 cm. long. For D. elatum this was true not only of the cultigens, but of the wild species as well, where even dwarf alpine plants have a high node number, only the length of the internodes being shortened. In branch- ing pattern these species may also be compared. On the whole (the exception being some garden types which are bred for a diffuse habit) side branches are few and much shorter than the main raceme which they subtend, generally decreasing in length from the lowest to the uppermost. The most evident distinction betwee” 1953] GAGE—DELPHINIUM X BELLADONNA HORT. 129 у y з * | 3 d al Y ё ade Y n ees % ol o 5 bb 18 74 6 Y Уу 6 "T ee iv vf x w| у € 3 4. YOE BED 11.9.49 €- € $9 Text-fig. 4. Scatter diagram showing character correlations in Delphinium grandiflorum (small dots, rayless), D. cheilanthum (small dots, rayed), D. elatum (intermediate dots), and D. X Bella- donna Hort. (large dots). 18 Bee 36 @entire - emarginate M notch 2 bifid z 30 Width of median segment T ei- z” 3-7 9 2 $e v ш 2 B За Density of inflorescence = 1 @10-70 д” 649-40 = 0 ф39-10 £N 9-1 ә 48 Number of flowers Ew ei4 or 9 - 95-25 и. №26 ог more 12 10 зи оса LENGTH MEDIAN SEGMENT/LENGTH C lati ext-fig. 5. Scatter diagram showing character correlations in members of the 4 triploid popu- ations of this study. Distribution of D. Belladonna in outline. these species, on the basis of habit, is that the inflorescences of D. elatum are rela- ing successively shorter through the inflorescence. may branch very little, characteristically the plant is much-branched, the branches Sometimes being placed at such an angle and being of such a length as to have the superficial appearance of dichotomies. (This is especially true in the closely [Vor. 40 130 ANNALS OF THE MISSOURI BOTANICAL GARDEN related species, D. tatsienense.) The branches may again branch in a similar manner (not figured). The maximum internode length in these relatively dwarf plants ranges from 5 to 18 cm., the higher numbers being quite common. Variability in habit is characteristic of hybrid populations, as Anderson has repeatedly pointed out (see, for example, Anderson and Schregardus, 1944; An- derson and Gage, 1952). In allopolyploids such variability should be small if the variety is of the type termed by Stebbins a true allopolyploid (1947, 1950). If however, it is a segmental allopolyploid, more frequent pairing between members of the different genomes will result in a higher amount of segregation. Belladonna does, in fact, vary somewhat in habit, although the general tendency is for these plants to possess a more or less regular internode pattern, with branching common below the main inflorescence. In extreme cases the node number may be low, as in D. grandiflorum, and the aspect of pseudodichotomies may be found in the branching pattern; or they may be tall plants with short branches and internode lengths irregular below the inflorescence. On the whole, the direction of variation in this respect tends to be more toward D. grandiflorum than toward D. elatum. TABLE II POLLEN SIZE AND VIABILITY IN SPECIES AND HYBRIDS OF DELPHINIUM: HERBARIUM SPECIMENS.* UN % Viable Pollen diameter in ш pollen Range Median Diploids = naciais, Vilmorin-Andrieux 98 20.0—25.0 22.5 Blue Butterfly 96 17.5—25.0 22.5 D. сБейат ит, coll. Karo 92 20.0—25.0 20.0 Tetraploids — оо D. elatum, Bot. Gard., Cambridge 98 22.5—25.0 25.0 D. elatum, King of the Blues 60 22.5—25.0 25.0 Hexaploids D. formosum Hort., N 7186 60 25.0—32.5 30.0 D. formosum Hort., Helms. 17/6/25 25 25.0—32.5 27.5 D. formosum Hort., Dreer, July 1924 50 22.5—32.5 30.0 ormosum Hort., For 80 22.5-32.5 30.0 D. Barlowii Hort.t 77 22.5—35.0 30.0 D. Barlowii, Aug. 1t 31 22.5-35 30.0 Lamartine, Aug. 20 79 25.0—32.5 30.0 о о "Specimens listed are in the herbarium of the Bailey Hortorium, Cornell University. : e confusion as to the identity of D. Barlowii has been discussed on page 120. The sper mens cited here are morphologically of the Belladonna sort. 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. 131 ELATUM- HORTICUL TURAL 0.909.999 7.0500 оо оро РҮҮ PM ELATUM- ALPINE [ BELLADONNA CHEILANTHUM А А МАМА, GRANDIF ORUM TRIPLOID- 48-27-3 Text-fig. 6. Internode diagrams to scale of diploid, tetraploid, and — Рон Ordinate, length of internode; abscissa, number of internode from rosett pright. unes р" le branches and are drawn to a scale 16 that employed for internodes. The black dots represent sing ower Text-fig. 5 is a scatter diagram for such of the triploids as were adequately represented by living or herbarium material, and an internode diagram of a typical triploid is shown in text-fig. 6. From the superimposed silhouette of the Bella- donna distribution (fig. 5) all these plants are seen to be within the limits of that group, ог to be intermediate between D. Belladonna and D. elatum. The same observation is borne out by the internode diagram, which shows the internode pattern of Belladonna, but with the high node number more characteristic of D. elatum, Summary.—Through the medium of the scatter diagram it has been demon- strated that Belladonna may be visualized as a more-or-less variable group, inter- mediate between the diploid D. grandiflorum (vel aff.) and the tetraploid D. elatum, and having as the source of its variation the segregation of characters derived from these species. Further, the experimentally produced hybrids between D. elatum and D. grandiflorum have been shown to be very similar to Belladonna, deviating from it only in characters for which their D. elatum parents were ex- treme. The scatter diagram also amplifies the position of D. cheilanthum: it is instantly differentiated from D. Belladonna by its diploid constitution, and by Constant morphological characters. The same conclusion is supported by the internode diagrams: amount of variability in D. Belladonna (extreme types were diagra there is a greater mmed, but are [Vor. 40 132 ANNALS OF THE MISSOURI BOTANICAL GARDEN not included in text-fig. 6) than in the three natural species, but on the whole it is intermediate between D. grandiflorum and D. elatum. In this respect D. cheilantbum is closer to D. elatum than to any other entity, differing chiefly in the development of its inflorescence. On the basis of character association, then, it is to be concluded that D. grandi- florum and D. elatum fulfill the morphological requirements for the parents of Belladonna; but that D. cheilanthum could not have been the diploid parent, given D. elatum аз the tetraploid parent. One cannot thereby rule out the pos- sibility that a tetraploid species other than D. elatum was involved, but no other known tetraploid species possesses the required combination of characters, nor is any other tetraploid so commonly grown. ГУ. CYTOLOGY or DELPHINIUM HISTORY The chromosome complement of Delphinium has been studied by a number of workers (Hocquette, 1922; Tjebbes, 1927; Langlet, 1927, 1932; Tischler, 1927; Beckman, 1928; Lewitsky, 1931; Lawrence, 1936; Propach, 1939, 1940; Gregory, 1941; Mehlquist её al, 1943, and unpublished; Lewis et al, 1947; and Lewis and Epling, 1951), and it has now been generally established that the basic complement of the genus consists of one long chromosome with a median centromere, one long chromosome with a sub-median centromere, five medium-length chromosomes with sub-terminal centromeres, and one short chromosome with a sub-terminal centro- mere. Langlet (1932) reported that satellites occurred in the genus, but his results were not conclusive regarding their number and location. Lewitsky (1931) published some data regarding chromosome differentiation within the genus, ай furnished ideograms for a number of species. Noting that in certain instance the four “homologous” chromosomes are not identical, he postulated amphidiploid origin for two tetraploid species, D. azureum* and D. elatum. Moreover, he pointed out that two species belonging to two distinct sections of the genus D. cardiopetalum and D. Staphisagria, have distinct types of ideograms from the other species he investigated. Моге recently, Lewis, Epling, Mehlquist and Wyckof (1951), reported that the chromosomes of the California species of Delphinium (which according to Ewan (1945) include representatives of several of the old World lines as well as a majority of endemics of uncertain affinities) 2 morphologically quite similar. They figure a composite ideogram from species included in Ewan’s SPICIFORM series, which shows one satellited chromosome (their E, which they report is otherwise very close in length to C, D, and F). That situation is not 80 simple as this and that greater possibilities exist for identifying different chromosomes in certain cases have been made clear from the present study in which, apparently for the first time in Delphinium, Feulgen and ace lacmoid squashes have been employed for somatic material. * D. azurem of Lewitsky was probably D. elatum; see footnote p. 164 1953] GAGE—DELPHINIUM Ж BELLADONNA HORT. 133 Numbers of chromosomes published so far in Delphinium are listed in the Ap- pendix. Diploid species are by far the most numerous, tetraploids are occasional, and hexaploids are known only in cultivated forms. Meiotic studies of Delpbinium have been largely confined to the garden forms, the major exception being the work reported by Lewis, Epling, Mehlquist, and Wyckoff on the California species, where both diploids and tetraploids were analyzed. For the diploid species, metaphase bivalents fall into several easily recognized types, since terminalization regularly occurs and chiasma frequency for a given pair shows little variation, the large chromosomes having usually two chiasmata at metaphase (there may be three or rarely four at diplotene and diakinesis) and the others having regularly one chiasma only, and that in the longer arm (Mehlquist e£ al, 1943; Lewis e£ al, 1951). In the tetraploids, the type of pairing varies, depending upon the origin of the tetraploid condition. Thus, in the California tetraploids, which are designated as races of the diploid species, D. Hanseni, D. gypsophilum, and D. variegatum, quadrivalents are formed by about 70 per cent of the long chromosomes with the submedian centromere, and by about 45 per cent of the long chromosomes with the subterminal centromere. (Likelihood of multivalent formation by the other chromosomes is precluded by their known low chiasma frequency.) Anaphase separation is regular, and the plants are fully fertile. However, it was demonstrated that the chromosomes are not completely homologous, since anaphase bridges indicative of inversions were common. Of other tetraploids studied, Lawrence (1936) recorded one to four univalents and an undetermined number of multivalents in Delphinium Ruysii, a garden hybrid possessing one D. elatum (n — 16) and two D. mudicaule (n — 8) genomes. А similar condition was found to exist in a group of tetraploid hybrids Produced by Mehlquist (unpublished) from crossing D. elatum with artificially doubled D. cardinale, in which meiotic pairing was found to be primarily by bivalents and univalents, although segregation in succeeding generations indicated that some multivalent pairing occasionally occurred. Particularly interesting in so far as this study is concerned are the varieties bred by К. Foerster of Bornim, Germany, and studied by Propach (1939, 1940). According to Propach, Foerster produced his hybrids by intercrossing D. Bella- donna varieties and D. grandiflorum var. chinensis, obtaining in the first generation fully fertile tetraploids. Later, some of these hybrids were again crossed with D. grandiflorum to form sterile triploids (Appendix). Unfortunately, full cyto- logical details were not given. Propach interpreted his results as indicating that there is little genomic differentiation in Delphinium and that autopolyploidy is involved.5 5 i . i ieties in all cases, d О (909) dee n Fr аши Тури and кире dtum X reno va chinense hybrids. Further, the accom i lates confirm his statements; нане thorough е companying р nfirm а cytological study of these forms is desirable before Propach's conclusions can be accepte f [Vor. 40 134 ANNALS OF THE MISSOURI BOTANICAL GARDEN The first object of this study was to discover whether any morphological differentiation of the chromosomes could be made among species of Delphinium, so that, if possible, the component genomes of the polyploid species might be identified. The second phase of the cytological work has dealt with the meiotic behavior of the chromosomes in the species and hybrids. Data obtained from chromosome studies have been supplemented by pollen studies which, to some ex- tent, serve to indicate the degree of fertility of a plant; and pollen measurements have been used, especially of herbarium specimens, to give information (which could not be obtained by other means) on the probable polyploid level of these plants. METHODS 1. Somatic Tissues—Chromosomes: Although the paraffin method has been the one used by previous workers, it was seldom employed here. This was not only on account of the length of time required to prepare the mounts, but also because root tips from sectioned material did not show the structural features evident in acetic squashes, nor could counts of chromosome number be made so readily. For squash techniques, it was necessary to adapt the method to the particular requirements of the material. Pre-treatment with a spindle inhibitor was found to be essential since the chromosomes of meta- phase plates were otherwise obtained in face-view only with the greatest difficulty; further, aceto-carmine did not give the degree of differentiation obtained with aceto-lacmoid. The schedule follows: а. Actively growing root-tips were removed and treated with a saturated solution of paradichlorobenzene (Meyer, 1945) for 3 to 4 hours. b. Tips were transferred to 7:3 acetic alcohol (Darlington & La Cour, 1947) for 1 to 5 days. Preparations from material preserved longer than this are not clear enough for studies of details of chromosome morphology, but are adequate for counts. Tips were hydrolyzed in a mixture of approximately 2 drops 0.1 normal НСІ and 5—6 drops acetic-lacmoid stain (Darlington & La Cour, 1947), heating on a glass until vapor could be seen, and then letting the tips remain in the mixture for 10 to 20 minutes, or until they became soft. (Sometimes it was necessary ® increase the proportion of HCl in the staining mixture in order to soften the tips sufficiently, but generally as little acid as possible was used since an excess decrease staining.) d. Root tips were rinsed in 45 per cent acetic acid for 5 minutes to about a? hour. e. The deeply stained portion of a tip was transferred to a clean slide, and stain and cover-glass were added. The mount was heated and squashed and then sealed. After the cells were dispersed by gentle tapping, considerable pressurt could be applied in order to spread out their contents. The preparations will keep for about a week if stored in a cool place, but they are best within a few hours after being made. 1953] GAGE—DELPHINIUM X BELLADONNA HORT. 135 2. Somatic Tissues—Nucleoli: . The Feulgen-fast green squash technique (Darlington & La Cour, 1947) was used. Hydrolysis time was about 20 to 40 minutes. 3. Meiotic Chromosomes: The exact method used depended upon the age and condition of the material. In general, the following schedule was employed: Anthers were killed and fixed in acetic alcohol for several hours to a week or more. (Best results were obtained if mounts were made in about one to four b. A whole anther was transferred to a slide, flattened with the needle, and stained. 1. Fresh material: acetic lacmoid gave the best results. | 2. Older material: a mixture of aceto-lacmoid and aceto-carmine was employed, the relative amount of carmine being increased with increasing age of the material. 3. If the material was not stored for more than two weeks, it could be transferred to 70 per cent ethanol, and thereafter stained with carmine, in which case mordanting was found to be necessary. с. A cover slip was added and the slide was squashed gently and sealed. After the slide had been sealed it could be heated and again squashed if necessary. 4. Pollen Studies: Preparations were made with either aceto-carmine or acid fuchsin, at least three slides being made for each plant of which the pollen was available. The anthers were taken on different days and from different flowers so that the effect of environmental variation could be observed. A total of 200 pollen grains was counted on each slide, only the deeply staining, normally shaped grains being considered "viable." Where possible, 100 g grains of each individual were measured, but in the triploids the number is understandably considerably less. Pollen preparations from herbarium specimens were made employing the fast- green glycerine jelly method of Wodehouse (1935). 5. Illustrations: Camera-lucida drawings were made with a Spencer Camera Lucida on a Spen- сег microscope with 15 X ocular and 95 X М.А. 1.40 apochromatic oil-immersion objective, giving a magnification of 2180 X at table height. Reductions are given with each figure. The photomicrographs were taken with a 1 X Bausch & Lomb fixed bellows camera; magnifications accompany the plates and text-figures. DETERMINATION OF THE LEVEL OF PLOIDY While the determination of polyploid level in the living plants wae always made by chromosome counts from root-tip preparations, the impossibility of applying direct methods to plants represented only by herbarium material made necessary the use of the indirect method of pollen measurement. In studying the Percentage of "viable" pollen produced by hybrid individuals, it had previously [Уот. 40 136 ANNALS OF THE MISSOURI BOTANICAL GARDEN been noted that diameter of the pollen grain was a good indicator of the polyploid level of the plant (Table I). For the plants measured there was good separation of diploids and tetraploids on this basis. The hexaploids and triploids generally produced the largest pollen, although there was some overlapping between these and D. elatum. It is to be noted that pollen production by the triploids is very low (except in the colchicine-treated spike of 50-40-2, which is presumed to have been doubled); the similarity in size of their few pollen grains to that of the hexaploids is ап indication that the pollen from these plants is unreduced, and hence triploid. For pollen grains of the herbarium specimens, size of pollen grain was again found to be a differential character for the polyploid level, although the actual sizes obtained were somewhat smaller in all classes, as might be expected on the basis of the difference in the suspending medium of the stain and from long drying of the material (Table II). CYTOLOGY OF NATURAL SPECIES, DELPHINIUM BELLADONNA, AND THE TRIPLOIDS Morphology of the Somatic Chromosomes.— n view of the general similarities in size of the chromosomes already noted as characteristic of the genus, ideograms are presented here for two species only: D. grandiflorum var. chinensis and D. cheilanthum. ‘The A and B chromosomes of these species probably are to be compared with similarly lettered chromosomes in the ideograms of Lewitsky (1931) and Lewis e£ al (1951), but because of the similar sizes of the smaller chromosomes it is not to be assumed that these are com- parable to similarly lettered chromosomes in the other ideograms. The major distinguishing features of the genomes figured here (pl. 13) have been the satellite-bearing chromosomes. The presence of nucleolar-organizing satellited chromosomes in the complement of many plants has been often noted since the classic papers of Heitz (1931), Navashin (1934), and McClintoc (1934). Later, the discovery of polyploid species in which the satellite number was comparably increased led certain workers to assume that often, if not always apparently diploid species which possessed several satellites in the gametic comple- ments are actually of polyploid derivation. This was the view held by Gates who published a comprehensive review on this subject in 1942. However, since many polyploids never have more than two satellites, and since, on the other hand, in such plants as Leontodon leysseri three of the four chromosomes of the haploid complement bear satellites (Elliot, 1950), inferences as to polyploid level based on satellite or nucleolar number alone must be regarded as unsound. Another problem related to the existence of satellites must be considered № order to appreciate the variations in satellite number observed in the Delphinium hybrids, and that is differential amphiplasty. This phenomenon was first noted by Navashin (1927, quoted by him in 1934), who applied the term to the apparent loss of satellites which sometimes occurs in hybrids. The following year, Mc- Clintock, in her paper on the nucleolar organizing chromosome of Zea Mays 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. 137 discussed in detail Navashin’s work, interpreting the loss of satellites in hybrid forms as being due to the inability of the chromosome to "organize" a nucleolus in the presence of another more "active" chromosome. Amphiplasty has recently been reported in the genus Leontodon (Elliot, 1950), and had been occasionally reported in other genera (Meurman and Therman, 1939; Levan, 1937). While our present understanding of differential amphiplasty is by no means perfect, the cases cited above seem to show that under certain conditions such as obtain in some hybrid cells, the apparent morphology of nucleolar organizing chromosomes may be modified. Since the presence of a satellite at metaphase is usually directly related to the formation of a nucleolus in that region in the preceding telophase, then the causative factor is apparently related to nucleolar metabolism. Nevertheless, the actual physiological differences between individual nucleolar "organizers," which are the essence of the problem, are not known at all, and it is therefore the morphological aspect of the phenomenon which concerns us ere. Diploids.—In pl. 13 is given the ideogram of D. grandiflorum L. var. chinensis Fisch. hort. var. Blue Butterfly, pollen parent of the 48-27 and 48-6 series of triploids. The actual length of metaphase chromosomes ranges from 3.1 to 10.1 p. (These measurements are based on chromosomes not artificially contracted by PDB, although the chromosomes measured were subjected to minimum treatment in order that the spindle would not interfere with their being well-flattened.) Of the shorter, subterminally constricted chromosomes, chromosome H is readily identified by its length as well as by the satellite; of the rest, chromosome C may be identified by its satellite and sometimes by the secondary constriction of the long arm. Chromosomes D through G are individually distinguishable only with difficulty; the lengths of their short arms, on the whole, were found to be more diagnostic than the long arms, which may be variously extended, being slower to complete their coiling cycle than the short arms. А difference in the activity of the satellites was noted here. While usually both C's were satellited, no cells were ever observed in which at least one C did not bear a satellite; the satellite of the Н chromosome, on the other hand, was commonly present in unicate, and sometimes was even missing altogether. This Was generally true for all plants of D. grandiflorum and was statistically deter- mined for two individuals (Table II). Related plants, including 47031-1 from the Royal Botanic Gardens at Kew, D. grandiflorum L. var. chinense Fisch. from Basel, and numerous plants of horti- Cultural varieties such as Tom Thumb and White Butterfly were examined and found to have a similarly differentiated complement (pl. 13). 47031-1 was found to possess a telocentric fragment, about the length of the long arm of the H “Ju, the conres of numerous observations, in one plan Оп one of the oi chromosomes, but as it was not present in an omitted from the ideog this ies a third satellite was noted die ui er case it has been [Vor. 40 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN chromosome. Though the numbers of fragments present in different cells of an individual have been found to vary somewhat, the fragment is commonly trans- mitted through the gametes, being found in plants three and four generations removed from 47031-1. Unfortunately, the original plant and the fragment- bearing triploids were lost before pairing affinities of the fragment could be determined, and there are sufficient irregularities in the descendant hexaploids to make its identification in PMC's doubtful. Plants of D. tatsienense Franch., a species closely related to D. grandiflorum І, were found to possess a genome similarly differentiated except that in the indi- viduals surveyed the satellite is as often developed on the H chromosome as on the C chromosome (see 51-311-9, Table III). D. cheilanthum.—By contrast with D. grandiflorum, D. cheilanthum bears its satellites on the A and C chromosomes, that on the A being not only larger but also the more "active" of the two (Table III, 51-307-6 and 51-307-1). $e far as can be ascertained, the chromosomes of the two species are otherwise similar, except that the satellite-bearing arm of A is slightly longer here than in D. grandiflorum. SATELLITE EXPRESSION IN DELPHINIUM SPECIES AND HYBRIDS Fraction of chromosomes bearin : | sstellite* B Number Theoretic 1 Actual average Plant of satellite | . lite number | C | H | А | се number Diploids grandiflorum 51-312-6 0.94 0.48 = 100 2:2:0 1.8:1.0:0 51-315-6 0.91 0.62 d. 100 2:2:0 1.8:1.2:0 tatsienense 51-311-9 0.85 0.80 = 100 2:2:0 1.7:1.6:0 cheilanthum 51-307-6 0.47 = 0.95 100 2:0:2 1.0:0:1.9 51-307-1 0.49 a 0.98 100 2:0:2 1.0:0:1.9 Tetraploids Е. D. elatum 51-318-2 0.56 0.12 0.85 21 4:2:2 2.2:0.25:1.6 51-318-4 0.50 0.07 0.92 71 4:2:2 2.0:0.15:1.8 Triploids 48-27-3 0.61 0.33 0.05 100 3:2:1 1.8:0.67:0.05 50-15-3 0.52 0.34 0.63 100 3:2:1 1.5:0.68:0.63 Hexaploids mede ete Smith's Belladonna 0.63 0.10 0.84 25 6:4:2 Ms 51-232-4 0.45 0.25 0.13 60 6:4:2 2.7:1.0:0-3 wur p лш т 7 *Fraction is computed on the basis of the actual observed maximum number of satellites for that chromosome. 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. 139 Other diploids.—A third situation with respect to satellite number and position was found in D. cardinale (pl. 13) and D. nudicaule from California, and a fourth in D. Zalil Aitch. and Hemsl. (native to Persia and Turkestan) and D. sulpbureum Boiss. and Hausskn. (from north Syria). In the California species the satellite number is 5, and the satellites occur on the H and on two pairs of intermediate chromosomes, one of which may or may not correspond to C, the lengths of the chromosomes in these species being not exactly comparable to those of D. cheil- antbum and D. grandiflorum. ‘The third satellite is on the second shortest chromosome. Despite the high potential satellite number, the low number of nucleoli usually present at telophase and interphase (generally no more than 4) indicates that all are rarely active in a particular mitotic cycle. In D. Zalil and D. sulphureum satellites occur on A and on one of the intermediate chromosomes. However, the genome differs from that of D. cheilanthum in the arm lengths of the intermediate chromosomes, so that direct comparisons cannot be made. In studying satellite development in the above diploid species a situation was noted which has apparently not been described before, namely, that even within old diploid species differential "activity" of satellited chromosomes may be com- mon. With this in mind further variations in satellite development which are found in polyploid species may not be so unexpected. In order to avoid ambiguity here, I shall use the term “differential activity” to apply to the differences in satellite development such as characterize diploid species, and "amphiplasty" to refer to any pronounced alteration in differential activity which follows hybridiza- tion. Actually, however, it is not to be assumed on such scanty visual evidence that any physiological distinction is being made. Tetraploids: D. elatum.—1n D. elatum, the only tetraploid species observed, distinctions among the chromosomes are again difficult to make. There are 2 Pairs of long, medianly constricted A chromosomes, 2 pairs of long, sub-medianly constricted B chromosomes, 10 pairs of intermediate chromosomes with subtermi- nal constrictions, and two pairs of short chromosomes with subterminal constric- tions. Amphiplasty obscures the satellite situation, the more active A an satellites usually being the only ones present (Table Ш), but a study of а large number of cells, both in the species (accessions from Géteborg Botanic Garden, and the Royal Botanic Gardens at Kew) and of horticultural variants, indicates the presence of satellites on 1 А, 2 C's and 1 Н of the haploid complement. Although the number of satellites in the tetraploids parental to the triploids could not be determined statistically, camera-lucida drawings of the complements of 47-98-3 and Summer Skies indicate that these plants were typical in this respect (pl. 14). Asin D. cheilanthum, the satellited A chromosome stands out from its counterpart in the other genome by virtue of its longer satellited arm. Lewitsky (1931) noted this difference in length, but did not observe the satellite. The nature of the chromosome differentiation in D. elatum is thus strongly suggestive of an allopolyploid origin by hybridization and subsequent doubling of the chromosome number of diploid forms whose chromosome complements were of [Vor. 40 140 ANNALS OF THE MISSOURI BOTANICAL GARDEN the types represented in present-day species D. grandiflorum and D. cheilanthum. That the chromosomes designated at C are truly comparable in the two genomes and are not different members of the medium-sized group is corroborated by the fact that both possess a secondary constriction of the longer arm. Aneuploids are occasionally found, even among the plants from the botanic gardens. However, they do not possess the abnormal characteristics which ordi- narily distinguish aneuploids on the diploid level, and fertility, as judged by amount of seed produced, is apparently not greatly impaired. Triploids.—All plants of the four groups of triploid hybrids were examined and found to be essentially similar in so far as the morphology of the chromosomes is concerned (pl. 14). The maximum satellite number in each is 6: 1 A, 3 С% and 2 H's, as would be expected on the basis of the generally regular segregation of the parents. The precise situation as to satellite development has been thoroughly worked out only for two individuals, 48-27-3 and 50-15-3, most of the other plants having died before this part of the study was undertaken. An interesting case is 48-27-3, in which the satellited A chromosome, so active in D. elatum, is rarely developed. (Study of camera-lucida drawings of other plants of this line and of the very similar 48-6 line indicates that this is generally true of both groups.) However, in 50-15-3, the other triploid analyzed, the A satellite was observed in more than 80 per cent of the cells examined. Thus, in crosses of similar parentage, amphiplasty has been demonstrated in one case, but did not occur in another. With regard to the centric fragment of their pollen parent 47031-1, triploids 50-15-1 and 50-15-4 were found to possess the fragment in most cells; 50-15-3, 50-40-1, and 50-40-2 lack the fragment. Hexaploids.—The tendency already noted for partial suppression of the poten- tial satellite number was especially evident in the hexaploids, of which Smith’s Belladonna,’ Cliveden Beauty, Bellamosum, and Lamartine were observed in some detail, as well as in the derived hexaploids which are to be considered in chap. ү. In most cases, cumulative records on these plants indicated a possible satellite number of 2 A's, 6 C's, and 4 H's of the diploid complement. However, the normal satellite number in individual cells is often only half this number (pl. 15). When interplant variation in degree and type of amphiplasty was studied Smith's Belladonna was found to possess very active А SATs, but among its progeny obtained by selfing, all degrees of activity were found; this was also true of the Fı obtained by crossing this plant with Bellamosum. At the other е treme are certain individuals of Bellamosum, in which the satellited A can rarely be distinguished (Table III). Meiosis and Fertility.— Meiotic studies have been made of the triploid hybrids and their parents, where these have been available, as well as of Belladonna varieties, in an effort to compat? "The plant termed "Smith's Belladonna” th : А : the horti- cultural variety "Cliveden Beauty." throughout this study is a selection from TABLE ТУ. MEIOSIS OF THE TETRAPLOIDS eu ا‎ HRS ا‎ eS ah д ана | an Eno E р TE TET а 4 Ly м та € “ K][t1naonzas == Е ТЕ I Е $22 sequin Sede 3 jewsouge EZET nnd 2s Á[[t1n32n235 +“ epo 29qumN. TEL TEE Т e الا‎ * е NN ч ч a urirouqt д, єз + НЕЕ ILI p qt 25 iz а у. 2 IIL ‘ds I [jn =m" own E mer Aa == LASS a РАНА I6 TI 9 Hr IN ¥ n m~ = jewsou gá E < Á]Je1n35n21$ aa АНИ — 3 2 $122 39qumN Z & тартка о ENS jewsouge z ipe Á][tan3on2as 3 = $122 1241 М < = + I zz Ija zi T $22 jo 19qumN о + < 7 colon ee А РОН Cee x а ед < I O1 ‘II It on Е Павии n = ЕСЕСІ ЕКСЕ g IS T dg ә = Z pcr n Е. I91 чу 9 e- e| „Ше ec A I9'"Uu «| >> B Ф £ її ПП <| ^ 9 ^ кары o » тия | 9^ - IZI JI 9| = Ke ie _—_—_ a 5 IZ ‘IE SI ык : z тот“ Lj te || CS II 91 "m п IS I$ SEM - о © Е ү Е ут Бони a 75 = ыыы то лм м а 5 Е чөдө $777 “Е oo oo оо o0 оо || 00 1 ooo es "Bot cb ob ocBdd TT Olli i. a "LHOH VNNOGVTIJH X WüINIHdTIdG—H5Vv9 [#561 [Vor. 40 142 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN chromosome homologies in the triploids and Belladonna. Early prophases are usually difficult to stain by squash methods, and while fair preparations can be made by paraffin sections, using gentian violet stain, the high number of chromo- somes present makes prophases generally of less use than later stages, which can readily be stained by conventional squash methods. For the most part, configura- tions at diakinesis and metaphase were analyzed, and anaphase I and II were scored for structural abnormalities. However, counts of chromosome segregation were not made, since the frequent occurrence of aneuploids in all polyploid populations is sufficient evidence that hyperploid and hypoploid gametes are produced. The approximate percentage of "viable" pollen produced by certain key plants is given in Table 1. However, since samples from the same plant taken on dif- ferent days may show a wide range of viability, the values given can scarcely be considered absolute indications of fertility. Moreover, apparently good grains may not survive until the pollen tube has reached the ovule, not only on account of some deficiency of the grain, but also, perhaps, because of incompatibility with the stylar tissue. Certainly, the particular environmental conditions prevailing at the time of pollen formation and shedding are not the least important factor in the production of functional pollen and its further development. It has repeatedly happened that a plant which produced a large amount of well-filled, staining pollen grains and set a high percentage of viable seeds at one time, produced little pollen and set few seeds at another time, or again, produced apparently good pollen but set no seed. The immediate cause of impaired fertility of this sort is often 2 prolonged hot spell, but other factors may have a similar effect. Diploids.—Blue Butterfly is usually quite regular in meiosis. In almost every one of numerous cells examined at diakinesis and MI, pairing was in 8 bivalents (pl 15). Chiasma frequencies ranged from 2 to 3 for chromosomes А and B, to 1 or occasionally 2 for the others. Anaphase separation is quite regular, bridges occurring in less than 1.0 per cent of the cells examined. A regular second division is followed by the simultaneous cleavage of the pollen mother cell into tetrads. Unfortunately, metaphase pairing of 47031-1 was not observed during the first year and the plant was subsequently lost. In anaphase I, however, laggards (probably the fragment) were noted in 9 per cent of the cells, and bridges in 4 per cent of the cells. The amount of “viable” pollen was about 65 per cent. The presence of the fragment in two of the five triploid offspring of this plant is satis- factory indication that pollen fertility was not prevented by the possession of the fragment. However, whether the failure of the plant to set any seed when self- or cross-pollinated may indicate some deleterious effect of the fragment in the ovule cannot now be shown. This case thus appears to be quite different from that of the centric fragment studied by Rhoades in Zea (1940), where trans mission of the fragment was largely through the egg, fragment-bearing pollen grains usually not competing successfully with normal pollen grains. Tetraploids.—Meiosis of Summer Skies is not entirely regular, the occurrence of 2 to 4 univalents at metaphase I being rather frequent. Further evidence of 1953] GAGE—DELPHINIUM X BELLADONNA HORT. 143 structural hybridity is given by anaphase I and second-division configurations, bridges, fragments, and lagging chromosomes being present in 11.3 per cent of the А I and 23.3 per cent of the A II cells examined (Table IV). Pollen viability appeared to be rather low (Table I), but that the plant is highly fertile has been indicated by the amount of seed produced. 47-98-3 exhibited fairly regular pairing; only two of the twenty diakinesis and MI plates showed each 2 univalents. However, a higher proportion of the large chromosomes had reduced chiasma frequency, with sometimes only 1 chiasma per bivalent. Further evidence of chromosomal hybridity is seen in the rather high number of cells at anaphase I which showed visible abnormalities (44.0 per cent). Abnormalities of the second division were less frequent, being present in only four of twenty cells examined. Pollen viability was about 65 per cent. 47-2-3, the tetraploid parent of the 48-6 group, was not available for study. Triploids.—Representatives of the several crosses have been analyzed. Since the number of PMC's examined in each case have not been large enough to permit separate discussion, the data have been somewhat grouped. 48-27-: Numbers 2, 3, 5, 6, and 12 have been studied (Tables V and VI, pl. 15). Metaphase pairing is largely in bivalents and univalents, only 34 trivalents having been found in the 258 cells analyzed. Usually, the long chromosomes (А and B) were involved in the trivalent configurations, but trivalents formed by chromosomes of intermediate size were found at least twice. The low number of trivalents involving the intermediate chromosomes may be due in part to their observed low-chiasma frequency, but also to the presence of differentiated seg- ments. The amount of even the A and B chromosomes present in trivalents is only 7.2 per cent. In two individuals (48-27-3 and 48-27-5) bivalent formation is near the maximum number expected if normal chiasma frequencies prevail—that is, if those chromosomes which pair form the normal number of chiasmata. However, whether such pairing is ordinarily between the 8 chromosomes contributed by D. grandi- florum and 8 chromosomes of D. elatum, or whether pairing is primarily within the elatum complement only can not be determined precisely. The above-men- tioned occurrence of occasional trivalents does indicate that some mixture of the two sorts prevails. That intragenomic pairing may occur in D. elatum is already known from the work of Lawrence (1936) and Mehlquist (unpublished) ; but the morphological similarity of one D. elatum genome to that of D. grandiflorum I5 equally strong indication that pairing between these chromosomes cannot be discounted, : Despite the rather high proportion of bivalents which may be formed in cer- tain individuals, the frequency of univalents, which may sometimes number З Тапу as 22, is indicative of structural differentiation of the chromosomes. This is further substantiated by the high percentage of structural abnormalities observed at anaphases I and П. At least one bridge-fragment was observed, on the average, їп about 50 per cent of the cells examined, both at A I and at A П. Such config- [Vor. 40 144 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE VI MEIOTIC ANAPHASES OF THE TRIPLOIDS Anaphase I Anaphase II 2 "a 8 ч E: Я Я 3 я g я Е 5 E Е 5 Е ob a 5 50 t=] E Plant be. = 2 + = С] B an E э B a = - | Е B 8$ 21 * B E 50 & 9 “Ж о on о е $13.24 ов орна Be ET 39 @ - ед a = بم‎ a - рд a Е = 48-27-2 i i i rog 64 | 67.2 7 17 30 | 433 48-27-3 4 61 4192] 20451 5% i4 4 151 Hi 7 | 627 48-27-5 29 33 12 33 107 69.2 12 7 5 35 59 | 407 48-27-6 10 | 21 23 $5. 60: |. — | — do bo = е, 48-27-12 44 25 16 39 124 68 — == pa: 48-6-1 i ОТС Zii p |. 3 1 3 14 | 500 48-6-3 13 | 10 6 173 34 ОЛУ Wit ГИ 5 | 10 39 | 74.3 50-15-1 ZU] HEI E R4 ov Зее Бос] се. ЕЕ 1:19 Wr 50-15-3 14 1 4 | 28 47 | 40.5 || 11 B 4 1.23 41 | 46. 50-15-4 19 2 45 62 0.3 | — | — | — = ran Bees urations have been shown by others to be caused by crossing-over within inversions, by crossing-over involving certain types of duplication (McClintock, 1941) о as suggested by Emsweller and Jones (1938), by pairing between chromosomes whose insertions are not opposite. The number and behavior of the lagging chromosomes are of interest. The average number of laggards was less than one per cell in A I, and even lower in A IL As a result, except for the numerical imbalance of most of the gametes formed by these plants in natural consequence of their triploid constitution, the loss of univalent chromosomes, which is often characteristic of such plants, W?$ not pronounced. This point will be considered further in the treatment of the hexaploids. The number of microspores per tetrad is variable, 2 and 3 being the more com- mon variants, but a fifth micrograin is occasionally formed. Micronuclei - extremely common, but are usually included with one of the other nuclei within a common wall. The amount of apparently good pollen has been found to be quite small, and that which is formed is within the range of pollen size of the hexaploids, 518565008 that such pollen is actually unreduced (Table I). -6-: Numbers 1 and 3 of this line were briefly analyzed. There was арр?” ently a greater tendency to bivalent formation in 48-6-1, 7 II having been not in several cases, while 2-6 II were found in 48-6-3. One trivalent was noted i? 48-6-1. In anaphase I and anaphase II somewhat higher frequencies of abnor- malities were noted in 48-6-1 than in 48-6-3, 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. 145 Viable (unreduced) pollen grains were formed in 2.8 per cent of cases in 48-6-3 and 5.7 per cent in 48-6-1 (pl. 15). 50-15-: Numbers 1 and 4 have been examined and seem to differ somewhat in pairing relationships. Single trivalents involving the large chromosomes occurred in 5 of 18 figures in 50-15-1, whereas no trivalents were noted in the 15 cells of 50-15-4 analyzed. Structural abnormalities at A I were present in about 40 per cent of the cells of each, and there were many irregularities at А П. Pollen viability was determined at 1.5 per cent and 2.5 per cent for numbers 1 and 4 respectively. 50-40-: A survey of meiosis in 50-40-2 indicated that pairing was usually in 5 to 7 bivalents, with an occasional trivalent. Reduced chiasma frequency was evident in the А and B chromosomes, and abnormalities of later stages were frequent. Moerbeimi was lost early in the study, but preliminary observations indicated that meiosis was comparable to the three hybrid groups. Pollen viability was ex- tremely low, less than 0.5 per cent unreduced pollen grains having been found. Hexaploids.—Belladonna: Lawrence (1936) has reported that meiosis in this form is largely regular. In Cliveden Beauty, a light-blue Belladonna variety, 2 to 4 univalents (of small or intermediate chromosomes) were noted in 10 out of 16 MI cells, but the chiasma frequency of the А and B chromosomes was only slightly reduced. At anaphase I structural abnormalities were visible in 12.3 per cent of the cells. Anaphase II was quite regular, only 7 of 121 РМС being visibly abnormal. Individuals of this variety were found to have from 63.2 to 80.5 per cent of good pollen. Preparations of Smith’s Belladonna showed somewhat less regular pairing at metaphase, with up to 8 univalents and the A and B chromosomes often with but one chiasma per pair. The actual amount of pairing was somewhat higher, how- ever, for in diakinesis univalents were uncommon and the large chromosomes had mostly two or three chiasmata in each bivalent (Table VII, pl. 15). Table VII gives some indication of the irregularities which are found in D. Belladonna varieties, although the plants cited by no means represent a random sample. A certain number of univalents are almost always present, but they assort at random and do not commonly lag at the plate. Multivalents are rare, but were noted in 52-325-11 (Smith’s Belladonna Х self), where both the A chromosomes and intermediate chromosomes were observed in such configurations. However, although one A chromosome was lacking from this plant, the monosome Progeny of various crosses. Thus, in 51-224 (Smith's Belladonna X Bellamosum, a dark blue Belladonna segregate) 8 of the 15 plants in the sample possessed 48 chromosomes, and in 52-311, produced by selfing а 48-chromosome individual of ` [Vor. 40 146 ANNALS OF THE MISSOURI BOTANICAL GARDEN MEIOSIS IN BELLADONNA TYPES S96 abnormal cells Metaphase I Anaphase I Anaphase II E 2 THERE 2 E Plant 3 3 d 3 E х 3 E i E ER 13 31434 з Е, 8| | =| 4% Е 2 ГЕЗ : 1] © ы в 5 aj 3 «| Х| > 1 Е E E Z| 8s! 3| 8| 3 a: я 8 |] аа ОЕ 1 е Ж 1 =i zi 61 dai a | z | 81434 Smith's Belladonna* | 48 | 55 | 24—20 0 0 23 || 12 2 2 33.3 || Mostly regular Cliveden Beauty-5 48 | 15 | 24-22 0 0 23 || 81 3 3 7.4 || 121 4 3 52-325-11 47 (SA). 32 | 23-19 | 4 2 22 || 48 3 9 25.0 || — | — " 51-224-16 48 | 21 ] 24-22 0 0 23$ 129 |. A4 7 35.9 14 1 0 $1-232-47 49 | 45 | 24-19 0 0 23 ||100 9 | 34 43.0 A9 rtine 48 | 46 | 24-21 0 0 23 || 67 6 4 14.9 || 104 7 6 * Of 21 late prophases observed, 17 showed 24 II, and 3 had each 23 II and 2 I. One cell had 6 univalents. this population, chromosome numbers of 25 plants ranged from 41 to 49, only 7 individuals having the "normal" number. Again, among 25 individuals of cross 51-232 (Bellamosum X self) chromosome numbers ranged from 45 to 49, 11 plants having the 48 chromosomes expected. In a sample of 10 plants obtained by selfing Smith's Belladonna (not figured), numbers ranged from 46 to 48. In most of the above cases the additional or missing chromosomes were of the smaller classes, but one plant with 48 chromosomes (51-232-1) lacked an A; and 51-224- 13, with 45 chromosomes, lacked a B. None of the aneuploids were visibly ab- correlation between chromosome number and pollen viability. Evidently, like many other polyploids, D. Belladonna can withstand the loss of a certain amount of genetic material without suffering the consequent weaknesses and abnormal development common when such losses occur at the diploid level. Discussion and Summary.— The cytological information given above bears on the problem of the identity 1953] GAGE—DELPHINIUM X BELLADONNA HORT. 147 E ЕЕ ER Text-fig. 7. Frequency distributions of chromosome numbers in populations of D. Belladonna re of the derived hexaploids. Ordinates represent per cent of population; abscissa, number of chromosomes, A—51-224 D-—51-216 G—51-219 J—51-222 B—52-311 E—51-217 H—51-220 K—51-223 C—51-232 F—51-218 I—51-221 not equivalent to morphological loss of the satellite was made evident by analyses of certain plants in which the rare occurrence of the satellite proved that its usual absence does not represent permanent loss. This condition is thus different from the actual morphological differences in satellite disposition between the chromo- somes of the two diploids D. grandiflorum and D. cheilanthum. Recognition of this difference is important for the interpretation of the satellite Pattern in D. elatum, the various triploids, and D. Belladonna. Thus, D. elatum might be a derivative of two diploid species whose genomes were differentiated along the lines of present-day D. grandiflorum and D. cheilanthum. And, given D. elatum as the tetraploid parent of D. Belladonna, D. grandiflorum might be the diploid parent; but D. cheilanthum could not have been, unless drastic mor- Phological alterations of the chromosomes have ensued since the time of origin of Belladonna. If the alternative supposition is made, that D. cheilanthum was the diploid parent of Belladonna, again, as with the morphological characters, there is the problem of finding a tetraploid other than D. elatum in the gardens of Europe— а tetraploid, moreover, both of whose genomes of the haploid complement are . similarly differentiated. [Vor. 40 148 ANNALS OF THE MISSOURI BOTANICAL GARDEN Analysis of pairing homologies of the chromosomes has proved illuminating in some respects, though not so useful in others. This was not unexpected, since in polyploids above the tetraploid level all combinations of autopolyploidy, segmental allopolyploidy, and amphiploidy may exist within a single individual (Stebbins, 1947, 1950). The triploid hybrids were found to exhibit neither the complete absence of pairing characteristic of the progeny of crosses between very distantly related species, nor the relatively complete pairing of some segmental allopolyploids, or of autopolyploids. True, as has been pointed out, the low metaphase chiasma fre- quency of the chromosomes would ordinarily prevent trivalent formation by all except the large A and B chromosomes, but the actual number of univalents was ordinarily higher than the eight expected if heterogenetic pairing either between grandiflorum and elatum chromosomes or within the elatum complement were of regular occurrence. Actually, the formation of occasional trivalents by the large chromosomes indicates that such pairing as does occur is a mixture of the two sorts. It has repeatedly been noted that pairing of chromosomes in the F, of such a species cross cannot be taken to indicate that a polyploid derivative of the cross (here a hexaploid) would necessarily be characterized by corresponding multi- valent formation (Darlington, 1937; Goodspeed and Bradley, 1942; Clausen, Keck and Hiesey, 1945; Stebbins, 1947, 1950). Rather, after doubling has oc- curred, differential affinity of the chromosomes may result in regular bivalent formation, not only in allopolyploids (Clausen and Goodspeed, 1925; Buxton and Newton, 1928; Kagawa and Nakajima, 1933) but even in known autopolyploids (Skirm, 1942). Since each case must be considered individually and the direct hexaploid deriva- tive has in this instance not been produced, the triploids can best be described as being the sort of plants which might well give rise to segmental allopolyploids, the low chiasma frequency of whose chromosomes would, on the whole, lead to more-or-less regular bivalent formation in the hybrid. One should expect, then, relatively little segregation in future generations, and the plants would continue to possess the more-or-less intermediate appearance of the triploids. The cytology of Delphinium Belladonna itself suggests that this species behaves rather like a segmental allopolyploid, but with perhaps fewer aberrations than аге generally found in polyploids of this type. Thus, while at first metaphase, pairing is primarily by bivalents, still the formation of univalents is not uncommon. Multivalents are rarely foun А consequence of univalent formation in gametogenesis is the production of numerous aneuploids. 'The mere occurrence at the polyploid level of viable aneuploids of good fertility is not unknown (Sears, 1944; Clausen and Cameron, 1944), although their high frequency here was not anticipated. In Triticum aestivum and Nicotiana tabacum, most of the nomosomics and nullisomics are individually recognizable, and this has been found to be true in certain other poly- ploid hybrids where aneuploids have been found (Primula kewensis, Newton 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. 149 Pellew, 1929; Nicotiana tabacum, Clausen and Goodspeed, 1924; Nicotiana rustica, Lammerts, 1932; Nicotiana glauca X М. Langsdorffii, Kostoff, 1939). However, while a part of the variability within Belladonna may be owing to aneuploidy, still it has not been possible to isolate any characteristic monosomic phenotypes, even for the large chromosomes, suggesting that there must be a considerable amount: of duplicated chromosomal and genetic material in D. Belladonna. This conclusion is corroborated by a survey of the extent of hypoploidy which Belladonna tolerates. Plants having from 43 to 44 chromosomes, while apparently infertile, are not themselves weaker than their euploid sibs. In most allopolyploids with fully differentiated genomes, such aneuploids as occur are usually deficient for no more than one or two chromosomes (Newton and Pellew, 1929; Müntzing, 1937). It is in plants in which the genomes contain many homologous chromo- somes or chromosomal segments that extreme aneuploids have generally been found (Müntzing, 1937; Myers and Hill, 1940; Myers, 1947; Love and Sunneson, 1945). Since the univalent chromosomes are usually included in the daughter nuclei, the prevalence of hypoploids among the plants of all the Belladonna populations studied can be explained on the assumption that in these relatively balanced plants, a gamete deficient for one or several chromosomes is at an advantage over gametes containing chromosomes in excess. Therefore, those gametes which function would then include 24 or fewer chromosomes. It is not, however, to be concluded that Belladonna is insensitive to chromosome losses of indefinite extent: on the Whole, aneuploids with 46 or 47 chromosomes are the most frequent, so that functional gametes must not, as a rule, lack more than one or two chromosomes. Further, though intervarietal crosses are easily made, the germinability of seeds Produced is higher in selfs than in intervarietal crosses (Table XI). It seems, therefore, that such chromosomal differentiation as exists between varieties may result in sufficient deficiencies either in the embryo or in the endosperm that in consequence the hybrid seed is less viable. Finally, since fertility tends to be gen- erally higher in the more nearly euploid individuals, and the progeny of the aneuploids do not vary more in chromosome number than do the progeny of normal individuals, D. Belladonna Hort. maintains itself approximately at hexaploid level. Of the many allopolyploids which have been analyzed (Stebbins, 1947, 19 50) the one perhaps most similar to D. Belladonna is the segmental allopolyploid Anemone Jancewskii (= A. sylvestris X А. multifida var. magellanica), first Synthesized by Jancewski before 1892, resynthesized by Gajewski (1946), and turther, occasionally spontaneous in botanic gardens. Morphologically, the hybrid 1$ intermediate between the parents and is quite constant, even aneuploids not being of abnormal appearance. A cytological study of the parents and the hybrids by Gajewski revealed that in the triploid (24 chromosomes) at MI from 3 to 8 bivalents are formed, the remaining chromosomes being unpaired. In the F2 8neration produced by the functioning of unreduced gametes (Jancewski re- Ported two instances of somatic doubling but Gajewski did not obtain any hexa- [Vor. 40 150 . ANNALS OF THE MISSOURI BOTANICAL GARDEN ploids by this means), in addition to cells in which 24 bivalents occur, in other cells there were some univalents and from 1 to 3 multivalents. Seed and pollen fertility in different individuals of this generation ranged from rather high to rather low, and in the Ез there was a general increase in fertility. The chromo- some numbers were uniformly about 48 in Gajewski's plants, but in the accessions from botanic gardens they ranged from 42 to 48. The chief respect in which A. Jancewskii seems to differ from D. Belladonna and from the hybrids synthesized here is in the complete absence of trivalents in the triploid and in the higher frequency of multivalents in the hexaploid. The first may be due to greater dif- ferentiation of the parental genomes; the second may well be the result of the much higher chiasma frequency of the chromosomes of this hybrid. To summarize, then, although D. Belladonna has previously been considered an amphidiploid on the basis of its largely regular meiosis and its relatively constant appearance, intermediate between certain diploid and tetraploid species of cultiva- tion, the garden hexaploid has been shown to have somewhat less regular meiosis than is characteristic of true amphidiploids. "This is indicated both by the common occurrence of univalents and other irregularities in meiosis, and in the higher production of viable, fertile aneuploids than is characteristic of such plants. Rather, D. Belladonna may better be classed as a segmental allopolyploid, in which the low chiasma frequencies of the chromosomes tend to enforce bivalent formation, and morphological constancy is further enhanced by the quantitative mode of inheri- tance of many of the differentiating characters. DERIVED HEXAPLOIDS For descriptive purposes, all of those plants having in their ancestry one of the triploids and Belladonna have been termed "derived hexaploids.” The pedigrees of the major lines are given in text-fig. 2. First- generation Hybrids: Triploids< Belladonna Varieties. — e dangers of making decisions regarding the nature of polyploidy in 4 hybrid, whether it be a segmental allopolyploid or an amphiploid, have been point out by Stebbins (1947, 1950). Neither external morphology of the chromosomes nor the nature of chromosome association can be used as a single criterion 1 of the triploids has not been achieved, a compromise has been made in the produc- tion and study of hybrids between the triploids and Smith's Belladonna. By using the triploids as female parents and pollinating them with Belladonna 23 F, seeds were obtained, of which 7 yielded plants which reached maturity. Unfortunately, these hexaploids, which are of the greatest interest in this study» are not known in as great detail as is desirable, since all but one were lost in the 1953] 51 GAGE—DELPHINIUM X BELLADONNA HORT. TABLE VIII MEIOSIS IN THE DERIVED HEXAPLOIDS Metaphase I Anaphase I Anaphase II Е * 5 i- Ф £ " & к: i3 i 3 4 Chromosome] ,, 3 ы ue A я 3 ч Мыс мш ра | 8 аера ва аа ЕЗ Б дш €) 2| Зач ||| за 3 Ф во Е ча s „д ud E 5 „д8 Е 50 3 E м © я - ad = g Da ы ы g g ная e hi D] з ч © ы "- Б ч © aa a Ф.н O d wa E в E Я 8 в Е я Е i з | 238 $ у ате) n аа * Z| 28 | Ro eee а ow Of Os] €^ | x1 01 Ба Ж 51-213-3 48, 2F 45 24-19 3 1 23 1139 6 | 13 13.7 60 9 | 2 18.3 51-216-6 49, 2Е 6 | 24—20 2 0 — | — Largely regular سے ا — اس‎ 51-216-10 50 (7А) |29 | 24-22 8 2 | 24| 45 1 4 1.1 || About 15% irreg. 51-216-16 50 (7А) 31 24—20 4 0 24 65 9 9 27.7 — — — — 51-219-4 46 55 22-17 0 0 21 50 8 2 20.0 37 2 2 11.2 51-220-4 48 10 23—20 1 0 — 25 3 2 20.0 — — — — 51-221-11 48 20 24—21 0 0 23 82 8 8 19.5 70 5 13 25.7 51-222-21 47 33 23—20 1 0 21 83 17 4 25.0 45 7 5 26.6 51-222-38 48 23 4—21 0 0 23 32 7 5 37.3 22 7 3 45.4 51-223-5 47 33 23-21.. 2 1 23 — — — — — — — — 51-223-31 44 7 22—1 0 0 61 8 2 16.7 36 7 5 25.0 [Vor. 40 152 ANNALS OF THE MISSOURI BOTANICAL GARDEN unusually severe winter of 1951-52 following their first flowering, and most of the meiotic material available for study was a few buds embedded in paraffin. Of the plants which were surveyed, 50-20-1 (48-6-2 X Smith's Belladonna) formed mostly bivalents at MI, but 6-12 univalents were commonly present, and reduced chiasma frequency in the long chromosomes was observed. At first ana- phase, abnormalities were observed in 28.0 per cent of the cells studied, but in only 10.0 per cent of cells at anaphase П. Pollen viability was 41.4 рег cent, and а fair amount of seed was set both by selfing and by backcrossing to Belladonna. 51-209-1 (48-27-5 X Smith's Belladonna) exhibited fairly regular pairing at MI. Usually no more than 4 univalents were found, and frequently 24 bivalents were present. Anaphase I was visibly normal in about 45 per cent of all cases, and AII was somewhat more regular. Pollen viability was in the neighborhood of 85 per cent. This plant produced a heavy set of viable seed, whether selfed or further backcrossed to Belladonna. 51-213-1 (50-15-4 Ж Smith's Belladonna) was studied in sectioned material, in which only approximate counts could be obtained. There was apparently rela- tively good pairing at MI—many PMC's appeared to be quite regular, and usually no more than 4 univalents were evident. Several times, a single trivalent involving one of the larger chromosomes was observed. АГз were regular in 60 per cent of the cases, as were most of the AII's. Pollen viability was as high as 91 per cent. Again, both selfs and backcrosses to Belladonna proved successful. А sister seed- ling, 51-213-2, showed metaphase pairing with generally 2—4 univalents, the re- maining chromosomes being in bivalent configurations. AI segregation in this plant appeared to be about 55 per cent normal. 51-213-3, studied in some detail (Table VIII), had at least 25 bivalents in two-thirds of the cells at MI, and both anaphase divisions were highly regular. Little trivalent formation was recorded, however. Two other hexaploids, 51-211-1 and 51-211-2 (50-15-1 X Cliveden Beauty); could not be analyzed; of these, the former yielded a good set of seed upon selling: The other plant was not pollinated. The advanced generations among the derived hexaploids had their origin, 3 may be seen in text-fig. 2, from five Е; individuals. Of these, two of the parental triploids had different white elatum parents but the same blue grandiflorum parent, while the triploid parent of the third line was the offspring of an azure elatum variety and the fragment-bearing cobalt diploid, 47031-1. In general, as will be seen, the lines derived from the last cross are distinct in cytological behavior from the other two. Somatic Chromosome Morphology.— Because of the number of plants represented by these populations, studies of somatic chromosomes have been restricted to root-tip counts, The total number of each of the easily identified A and B chromosomes was counted, gross morpho- logical alterations of the chromosomes were noted, and in instances where staining was good enough, average satellite number and disposition were determined. — Nm Bom ел М. Су ОО D m “~N ~ --oo0o0-o0oo0 q1uxu$ej]p| * © 12101. = тосола EDERT EE D: -m 22430 LRReecclecce = edd < = кк = a 92 о © = © ©+зс о о mo я я = yZ № -"-cooonuocococo a 5 [5 `6 E E as o ooooocooococ & | E >= a t 5 vs © ooocococcooooo a н 2d a Á[uo А 3024321 "^ oocooocoooo < N Sz Áquo pU DEG щ 5 рю]4пэчу < e a8 = [201L n No \0 \о © о К. со са н ^ =“ : n S 3940 [М ооо ооо Z < m к г [| = gz © oocooo-oooo p 5 z `6 t Б УЕ о ^"ooooooo-oc — 5 > = о [2 vs © ооооноооо н е о Áquo Шат че Se OOS © с т Áquo pio[dnouy o он м о о омо № ЮМ. 900 Oy © -— сч c6 Ym сч we сє сч N C4 СЧ СЧ CN. mM t r ee ttt — ہس سم ہے س ہے‎ жч =ч е сч; A WA м м м м м м м м 'LHOH VNNOQGVTTI8 X JWüINIHdTAIQ—45Vv5 [2561 [Vor. 40 154 ANNALS OF THE MISSOURI BOTANICAL GARDEN As to the number of satellites present, the chief difference noted among the various populations was in the expression of the A-SAT, which is potentially present on two of the 6 A chromosomes in all lines, since the triploid and the Belladonna parent each should have contributed one. However, while plants in the three populations descended from the 51-213 individuals usually revealed one or two satellited A's, this satellite was not strongly manifested by the other popu- lations. Apparently, this difference had its origin in the triploid parents, since, as was mentioned above, the 50-15 individuals were characterized by possessing a more "active" A than the other lines. Aneuploidy is common in all populations, as it is within Belladonna; however, the various lines have characteristic differences (text-fig. 7, Tables IX and X). Thus, while Belladonna populations contain many hypoploids with as few as 41 chromosomes and only occasional hyperploids, this pattern occurred only in 51-219 and 51-220 (51-202-3 X self, and Х Smith) among the derived hexaploids. А somewhat intermediate situation was found in 51-222 and 51-223 (51-209-1 X self, and X Smith) and in 51-221 (51-202-7 X 50-20-1) in which plants with 48 chromosomes were frequent and both hypoploids and hyperploids occurred. A third situation was found in lines 51-216, 51-217, and 51-218, derived from ‘the fragment-bearing individuals 51-213-1 and 51-213-2. In these populations, hyperploid individuals were most numerous, though an occasional plant with as few as 45 or 46 chromosomes was found. In addition, the centric fragment was quite common, being found in 17 out of 24 individuals of 51-216, and 23 of 26 individuals of 51-218. In 51-217, which was a backcross to Smith's Belladonna, the fragment occurred in 4 of 8 individuals examined. The exact number of fragments present in each individual is not given, since the fragment does not always segregate regularly and may occasionally vary from root-tip to root-tip IRREGULAR DISTRIBUTION OF CHROMOSOMES IN THE HEXAPLOIDS Number chromosomes Number ch es deficient in escent : Number Line of plants : А В Other A B Other 51-216 24 0 с З {з : i» 51-217 9 0 ó А e A 51-218 26 0 i n : : He 51—219 12 0 1 18 0 d 0 51—220 7 0 $ 12 4 : i 51-221 24 0 { ; о : 51-222 45 0 1 18 4 i 6 51—223 20 0 М i : : : 51-224 15 0 x y А 0 : 52-311 25 0 1 49 б 1 0 51-232 25 1 F hes : : : Total 232 1 6 173 19 5 51 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. 155 of a given plant. It is not known to what extent it may be possible to accumulate fragments in future generations, but in samples of two Ез populations, each of whose progenitors bore two fragments, no more than three fragments were ob- served in a total of fifteen individuals; on the other hand, all but one of the remainder possessed one or two fragments. Since transmission of the fragment through both the pollen and the ovule has been demonstrated (in the pollen, on the diploid and hexaploid levels, and in the ovule, on the triploid and hexaploid levels), it may well be that the failure to find many individuals with an augmented fragment number reflects some lethal effect of numerous fragments. It may be noted that the formation of isochromosomes by misdivision of the fragment, such as was noted by Darlington in Fritillaria (1940) and by Rhoades in Zea (1940) has never been observed here, although such a chromosome should be easily recog- nized by its distinct morphology. Fertility of tbe Derived Hexaploids.— А systematic study of all the plants has not been made, since they number now well over 1000. Аз has been noted above, the initial triploid hexaploid hybrids were of good fertility. In succeeding generations (to the F4 and various back- crosses to Belladonna) plants exhibiting all degrees of fertility have been found, as evidenced particularly by seed set. Detailed meiotic studies have been made in only a few of these plants ( Table VIII), but the degree of pairing has generally been found to agree well with that characteristic of D. Belladonna. ‘Thus, 51-221-11 forms no more than 6 uni- valents and usually only 2 at MI; no multivalents were noted; and both anaphases are fairly regular. A similar situation, with slightly more numerous irregularities in the anaphases, was found in 51-222-38. In aneuploid individuals, irregularities were, to be sure, more frequent. Thus, 51-216-10 and 51-216-16, both of which Possessed an A chromosome in excess as well as one other excess chromosome, often formed 24 bivalents, but were also characterized by low multivalent formation, chiefly involving the extra A chromosome. А hypoploid (51-219-4; 46 chromo- somes) averaged 21 bivalents, but the anaphases were not especially irregular. As in Belladonna, the number of laggards at anaphase was always much less than that of univalents at metaphase. While infertile plants have been found both among the aneuploids and among apparently normal plants with 48 chromosomes, a study of seed germinability among those plants which have set some seed shows that, on the whole, the highest number of viable seeds were produced by plants with 48 chromosomes. However, even in this group, germinability ranged from 0 to about 90 per cent, and, with опе exception, the production of seed with at least 40 per cent germinability was confined to plants having 47—49 chromosomes (Table XI). Segregation in the Derived Н exaploids.— On the whole, each line within this group exhibits considerable uniformity by the Ез (or equivalent) generation. However, there are few individuals in each 156 [Vor. 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN TA SEED GERMINABILITY IN DELPHINIUM BELLADONNA AND DERIVED HEXAPLOIDS Chromosome umber Number % Germin- Рим umber seeds sown seedlings ability Smith X self 27 21 78 Smith X Bellamosum 48 M ? 86 25 29 Lamartine X self 24 14 57 Lamartine Х Smith 48 X 48 52 8 17 216-3 X self 8 4 0 0 51-216-4 X self 49, 2F 2 0 0 51-216-12 X self 48, 2F 75 15 20 51-216-13 X self 52 15 0 0 51-218-2 X self 49, F 8 3 37 51-218-9 X self 48 14 2 14 51-218-16 X self 48, 3F 42 23 55 51-218-18 X self 49, 2F 2 0 0 51-218-19 X self 46, 2F 29 18 62 51-218-29 X self 50, 2F 68 23 34 51-218-39 X self 48, F 8 2 25 51-221-2 X self 49 50 30 60 51-222-11 X self 48 3 0 0 51-222-13 X self 47 36 4 11 51-222-19 X self 48 11 0 0 51-222-28 X self 48 4 0 0 51-222-7 X self 48 200 165 82 51-222-8 X self 48 47 21 44 51-222-14 X self 48 23 17 73 51-222-36 X $ 48 12 0 0 1-222-37 X self 48 32 14 44 51-222-43 X self 47 22 3 13 1-222-46 X self 48 8 1 13 51-222-50 46 33 16 48 51-222-54 X self 49 20 1 5 51-222-61 X self 48 22 13 59 51-222-67 X self 48 5 0 0 51-222-77 48 9 4 45 51-222-95 elf 49 15 7 47 51-222-100 X self 48 ра 1 14 51-223-1 X self 49 3 0 0 51-223-21 X self 48 10 6 60 51-223-22 X self 48 15 5 33 51-223-38 X self 48 14 5 34 51-223-44 X self 47 7 0 0 group which deviate noticeably from the rest, especially fee SEE in leaf shape. In only one instance (an Fs family derived from 5 1-209) have more than a few individuals deviated in the direction of either D. grandiflorum or D. elatum, and the variation here is in the direction of rather more finely dissected leaves than generally occur in D. Belladonna. Taken as a whole, however, the general aspect of the assemblage is that of a Belladonna population in which a few individuals are more or 169 branched, shorter or taller, or with leaves more or less finely dissected than in D. Belladonna Hort. (pl. 10). 1953] GAGE—DELPHINIUM X BELLADONNA HORT. 157 Color inheritance in these forms is such as to raise some interesting questions, but detailed treatment of this subject will be reserved for future consideration. Discussion and Summary.— Although the analysis of meiosis in the first-generation hybrids was not com- plete, it is still clear from the above that while somewhat more irregular than in Belladonna, the meiotic process does not appear to be greatly disturbed, and in only one individual (51-213-1) was the regular production of multivalents noted. Chromosome numbers among the progeny of these plants suggest for the various crosses what homologies and disharmonies may have been operative in their parents. Crosses 51-219-, 51-220-, and 51-223- are most similar to Belladonna in that chiefly euploids and hypoploids were produced. In the first two of these lines, which represent the self and backcross to Smith's Belladonna, respectively, of the Р» of the cross 48-6-2 X Smith's Belladonna, there has been no intentional building up of the Belladonna complement by backcrossing. Therefore, unless the introduced chromosomes of the triploid have been appreciably eliminated, no strong barriers of chromosomal differentiation appear to exist between the parental triploid and Belladonna. The aneuploidy thus seems to be of the same nature as in D. Belladonna Hort. On the other hand, 51-223 is a first backcross to Smith's Belladonna of the original hybrid between 48-27-5 and Smith's Belladonna. Since the same hybrid When selfed gave rise to line 51-222 (in which both hyperploids and hypoploids were frequent, and in which excessive numbers of supernumerary А chromosomes were found), it seems in this case that occasional formation of multivalents in- volving the large chromosomes must occur, and perhaps that more pronounced structural differences may exist between the parental genomes than in the previous instance. The first conclusion is based on the occurrence of 4 А chromosomes out of a total of 10 chromosomes extra in the hyperploid individuals of line 51-222, more than three times the expectancy if this chromosome were segregated irregu- larly only as often as the small chromosomes, and even more unlikely as а random event if the higher chiasma frequency of this chromosome is considered. In the absence of any indication of multivalent formation by the smaller chromosomes, it is necessary to assume that some other factor is operative in Producing individuals possessing these chromosomes in excess. This may be ac- counted for if gametes hyperploid for these chromosomes are not at the selective disadvantage which appears to occur in D. Belladonna. Such an hypothesis can t be explained if certain euploid combinations in these individuals are actually deficient for necessary genetic material which may be supplied by the extra chromosomes. The extreme frequency of hyperploidy in the 51-213 derivatives, where hyper- Ploids represent the bulk of the populations, seems to indicate both high multi- valent formation by the A chromosomes, occasional multivalent formation by the B chromosomes, and to a greater extent than in the preceding case, the operation of some factors which favor the functioning of gametes hyperploid for the smaller [Vor. 40 158 ANNALS OF THE MISSOURI BOTANICAL GARDEN chromosomes. This is particularly true of 51-216 and 51-217, the Ез and first backcross respectively of 51-213-1, the plant in which multivalents were noted above. That transmission of the excess large chromosomes may be through the pollen grain as well as through the egg is suggested by the fact that a comparable excess of large chromosomes was present in 51-217, the backcross line, as compared with 51-216. As to the fertility of the derived hexaploids, it has already been noted that euploid chromosome numbers and fertility are not necessarily related, but that, on the whole, more plants which yield seeds with good germination have about 48 chromosomes than have chromosomes either much in excess or in deficiency of this number. The tendency to preserve the hexaploid number appears to be almost as strong as in the Belladonna lines themselves. So far as the homologies of the genomes derived from triploid and hexaploid sources are concerned, it seems that they must be very nearly similar, since there as been no strong morphological or cytological disharmony in the derived hexa- ploids as compared with D. Belladonna. In fact, the differences are very slight indeed if one contrasts the behavior of these plants with that of the intervarietal crosses between Belladonna and Bellamosum. Although the latter variety was derived from the former, yet reduced seed germinability and more frequent aneuploid production have been found to occur in their hybrids than characterize either variety when it is selfed, GENERAL Discussion It is concluded that D. Belladonna Hort. is an allopolyploid hybrid between the tetraploid species D. elatum L. and the diploid species complex of D. grandi- orum L. In general, D. Belladonna may best be classed as a complex segmental allopolyploid, but it should be emphasized that classification is especially difficult since strong chromosomal barriers to hybridization do not seem to exist between even distantly related species of Delphinium (e.g., D. elatum and D. nudicaule ot D. cardinale). Unlike many other segmental allopolyploids, however, D. Bella- donna is easily distinguished from both parents. It has already been suggested that regularity in pairing, enforced by differential affinity of the chromosomes, as well as by the not insignificant effect of low chiasma frequency, is probably re- sponsible in large part for this behavior. Moreover, the quantitative mode first occurrence of fertility was reported; second, it has been subject to the care and selection of gardeners and plant breeders over the years, so that the difficulties (for instance, multivalent formation by the large chromosomes) such a hybrid would encounter during the first few generations in nature were mostly obviated. 1953] GAGE—DELPHINIUM X BELLADONNA HORT. 159 Several explanations may be offered to account for the elatum tendencies noted in the experimentally produced triploids. First, while D. grandiflorum has a num- ber of varieties, those varieties grown in gardens are, on the whole, similar with respect to the features which they contribute to the phenotype of Belladonna— the large self-colored bee, the wide open flower, the few-flowered inflorescence, the finely dissected leaves, and much-branched habit. D. elatum, on the other hand, exists in many natural and horticultural variants; probably a D. elatum bred for a showy "bee," with a relatively open inflorescence and rather narrow leaf seg- ments, would yield offspring of the type characteristic of Belladonna when such a variety was crossed with grandiflorum. However, the elatum variety Summer Skies, seed parent of triploids 50-15, has a small bee, broad leaf segments, and very dense strong main spike with numerous flowers; the triploids plotted in text-fig. 5 are indeed intermediate with respect to their known parents. All one can say of the presumptive elatum parent of Belladonna is that it was almost certainly a smaller, less extreme variety than those grown today, judging from historical plates of Delphiniums cultivated in the middle of the last century. The question of the nomenclature of the triploid and hexaploid garden Del- phiniums may be viewed in three very different aspects: first, as it concerns the taxonomist, second, the geneticist, and third, the gardener or nurseryman. While the first two views may be reconciled to a certain extent, the unfortunate truth is that the people most concerned with these Delphiniums (which are, after all, cultivated plants) will probably perpetuate the past errors and contribute new cases of mistaken identity in the future. For the geneticist, it is desirable that the exact parentage, insofar as it is pos- sible, be given for every hybrid where it is known. Convenience in the herbarium, on the other hand, requires that some category be devised which will adequately contain the assemblage. Thus, according to the International Code of Botanical Nomenclature (Lanjouw, ed., 1952), "hybrids or putative hybrids between two species of the same genus are designated by a formula, and whenever it seems useful ог necessary, by a name." In the latter case, the specific name is to be preceded Y "X." Further, where different hybrids of the same parentage exist, they may be united in a collective taxon in which the forms are recognized as nothomorphs, "designated by an epithet preceded by the binary name of the hybrid group and the term "nothomorph." While the general practice in systematics is to give Priority to the earliest validly published name, for cultivated plants provision is made that the most generally employed name may be used. Of the early names applied to hybrids of D. grandiflorum and D. elatum, only D. formosum and D. Belladonna are in wide usage. Since the name Delphinium formosum was given in 1856 to a wild species (Boissier and Huet), it seems advisable to maintain D. X Belladonna Hort. ex Bergm. (see footnote, p. 119) as the epithet of the hexa- Ploid Delphiniums and their triploid allies. Varieties such as Lamartine, Capri, Moerbeimi, and the triploids and derived hexaploids of this study may then be Considered as nothomorphs of this species. [Vor. 40 160 ANNALS OF THE MISSOURI BOTANICAL GARDEN SUMMARY 1. Previous workers have postulated that D. Belladonna Hort. originated either as a variety of D. cbeilantbum Fisch. ex DC., or as а hybrid of D. cheil- antbum with some unknown other species, or by hybridization between such diploid and tetraploid species as D. grandiflorum L. and D. elatum L. It is pointed out that solution of the problem must be based upon a combination of methods, including historical research, morphological analysis, and cytogenetic data. 2. A study of the taxonomic and horticultural literature indicates that D. cheilanthum is a coherent diploid species, rare in gardens, native to Siberia, Mon- golia, and Kumaon (China), with relatives in adjacent areas; D. Belladonna, on the other hand, is a hexaploid, known only in gardens, where it has arisen upon different occasions. Hybrids between various lines of D. elatum and D. grandiflorum var. cbinensis have been produced and have been found to be morphologically com- parable to D. X Belladonna Hort. ex Bergm. and, moreover, within the limits im- posed by their triploid condition, interfertile with the "species." 4. Character analysis of the various species and hybrids indicates that both Belladonna and the experimentally produced triploids are intermediate with regards to D. elatum and D. grandiflorum, but that D. cheilanthum is morphologically and cytologically distinct from D. Belladonna and could not have been the diploid parent of the latter if D. elatum was the tetraploid parent. 5. Methods are given for the study of somatic and meiotic chromosomes and of pollen mounts. 6. It is pointed out that for the species and hybrids studied, size of pollen grain may be used as an indicator of polyploid level in both living plants herbarium material. 7. Study of the morphology of the somatic chromosomes has shown that the genomes of the two diploid types which have been considered possible parents of D. Belladonna may de distinguished on the basis of their satellite distribution. n D. grandiflorum, satellites occur on chromosomes C and Н, while in D. < antbum the A and the C chromosomes are satellited. Further, the length of the satellited arm of chromosome A in D. cheilanthum is longer than the correspon’ ing arm of the same chromosome in D. grandiflorum, suggesting that there bis been a translocation of the satellite-bearing segment in the evolution of the genom* 8. Differential activity of the satellites was found to characterize even diploid species: in D. grandiflorum the C satellite is about twice as “active” ^ the H satellite; in the closely related D. tatsienense, both are equally active. >: D. cheilanthum the A satellite is about twice as active as the C satellite. 9. The tetraploid species D. elatum was found to have a gametic complement corresponding to the summation of the grandiflorum and the cheilanthum genomes: Amphiplasty of the H satellite occurs. 10. In all of the experimentally produced triploids the potential satellite number was found to be 1 A, 3 C’s, and 2 H's, but in triploids of lines 48-27 and 1953] GAGE—DELPHINIUM X BELLADONNA HORT. 161 48-6, the A satellite was rarely evident on account of amphiplasty. The mor- phology of the chromosomes of Belladonna supports the hypothesis of similar parentage for these types, since the total Belladonna complement has double the potential satellite number of the triploids. In both the triploids and in Belladonna, amphiplasty of the A satellite was found in some individuals. 11. Meiosis in D. grandiflorum and D. elatum is largely regular, the chromo- somes forming bivalents (rarely some univalents in the latter species), and assorting on the whole regularly. 12. In the triploid hybrids, prophase and metaphase pairing is chiefly in univalents and bivalents, trivalents being formed only rarely and only by the A or B chromosomes. Chiasma frequency is somewhat reduced and both inter- genomal and intra-genomal pairing of the chromosomes occurs. Bridges and frag- ments are characteristic of both anaphase divisions, although the number of chromosomes lagging at anaphase is considerably less than that of univalents at metaphase. Micronuclei are frequent, and rare viable unreduced pollen grains are produced. 13. Meiosis in Belladonna is more or less regular, but the production of uni- valents commonly occurs, and aneuploid gametes are produced and function. Chromosome numbers as low as 41 have been found, but ordinarily only those plants with about the hexaploid number are fully fertile. It is suggested that D. Belladonna is a segmental allopolyploid which preserves a uniform appearance because low chiasma frequencies of the chromosomes enforce bivalent formation and because of the quantitative mode of inheritance of many characters. 14. Lines of derived hexaploids have been produced by pollinating various triploids with pollen of Smith's Belladonna, and further selfing or backcrossing of the plants produced. On the whole, these lines are only slightly more variable than is D. Belladonna, and fertility is only slightly impaired as compared with that "species." A small amount of multivalent formation characterizes some of es. _ 15. It is concluded that D. X Belladonna Hort. ex Bergm. embraces the en- ure assemblage of present-day varieties derived from crossing D. elatum with D. grandiflorum, as well as а number of hybrids of the last century which are no longer in cultivation. 16. A telocentric fragment has been transmitted from a D. grandiflorum Pollen parent through four generations. No isochromosomes have been observed. BrBLIOGRAPHY ее, (1907). Roy. Hort. Soc. Jour. 32:lxxviii. m duds » (1911). Delphinium Moorheimii. Gard. Chron. Ш, 50:352-353. ее Oe Miscellaneous annuals etc. at Wisley. Delphinium. Roy. Hort. Soc. 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Bruscia ; (1943). peter induced tetraploidy in el phinium В Jou E Hed. 34:187—192. Meurman, O., and E. Ta (1939). Stia on the chromosome morphology and structural ridity > А, genus Clematis. Cytologia 10:1-14 Meyer, к es 945). Prefixing with paradichloedbenzene to facilitate chromosome study. Stain Pini. Moore, T (аз). Delphinium cheilanthum var. Hendersonii А. Henfr. Gard. Mag. Bot. 2 üntzing, А. (1932). Cytogenetic investigations on synthetic Galeopsis Prosar Hereditas té: diis: 154 — ——5 (1937). The effects of chromosomal variation in Dactylis. “ат, d HT Myers, У. M. ed ). Crilogr and genetics of forage grasses. Bot. —421. 7 ——, and H. D. Hill (1940). Studies of chromosomal association ==” fedet н“ трлр of пасагу іп autotetraploid grass species, nbus grass, tall oat grass, and crested whea Brass. he 102:236—255. Navashin, м" (19 34). Chromosome alterations caused by hybridization and their bearing upon certain ке genetic problems. Cytologia 5:169—203. AN 2 С. F., and С. Pellew (1929). Primula kewensis and its derivatives. Jour. Genet. 20: Paxton V. ODE Delphinium Barlowii. Paxt. Mag. B , . Mag. Bot. рь С. A. (1949). Delphiniums, Their History and Culti tie Eyre & Spottiswoode, London. 939 i h сораса, ). Was ist eigentlich unsere Rittersporne? Gartenschónh. 20:283—284. ш, Кр 40). СР Chromosomenzahlen von Delphinien und ihre Auswertung für die ung der Gartenformen. Die Gartenbauwissenschaft 14:642—650. Regd, p nmi utiles ung дег. Baikalien, Dahurien, und am Amur ..... Gesammelten Rh . Abt. Soc. Imp. Nat. Mace Bull. 34: 1-211. : Ў "d › M. M. (1940). — a a арша. seem in maize with reference to t t Mak, Bc ri В. (1933)? "The first red Delphinium," in Delpbinium, L. Leonian, ed. sum 35737. ischkin, В. К. (1937). “Del hinium," in V. L. Komarov, Flora "URSS 7:101 pie E. s дч, ploi — of wheat. II. Additional chromo- berrations in Tri vulgare. Genetics 29:232—24 Schrank, P. von (1819). Vorrei, cheilanthes.” Plantae asain Horti Academici Monacencis. Sim, б ©. Y. (1942). Bivalent pairing in an induced tetraploid of Tradescantia. Genetics 27:635- [Vor. 40 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN Stebbins, G. L. (19 m - be жи their classification and significance, in М. Demerec, Advances in Genetics 1:4 ‚ (1950). Variation pene. dum on in Plants. Columbia Univ. Pres, New York. —— ———, (1952). The — - ыы plants = weeds. Evolution 6:445—448. Tjebbes, K. (1927). The chrom es of three Delphinium species. a E e MP. Tournefort, J. P. (1700). Delph Вы um. Р ФА Aloüetie, in in Inst. Rei Herb. Trautvetter, E. R. (1847). Florula taimyrensis phanerogama, in A. T. аа, Sibirische Reise 1:17—64 » (1877a). Plantas Sibiriae borealis ab A. Czekanowski et F. Mueller. Hortus Petropol. Acta 5:1-1 —— ——, (18 77b). Flora riparia Kolymensis. lbid. 5:497-574. Turczaninow, N. (1842). Flora Baicalensi-dahurica . . . Part L Soc. Imp. Nat. Moscou, Bull. 15 Li Vanderbilt, N. Е. (1933). American Pioneers. I. The Vanderbilts. In Delphinium. L. Leonian ed." Van r ht 1857). Delphinium formosum, Hortul. Fl. des — en - ^ і 1783; Wilde, E. к ). Studies of се genus -— een Exp Wod APPENDIX CHROMOSOME NUMBERS OCCURRING IN THE SPECIES OF DELPHINIUM О Ru NaN ых Species 2n number Author and date Ajacis L. 16 reges 1927; Tjebbes, 1927; Gregory, 1941 Andersoni Gray 16 Lew: al, azureum Michx.* 32? pao 1931; Gregory, 1 Belladonna Hort. 48 Langlet, 1927; Lawrence НН Propach, 1939, 1940 lochmanae Greene 16 LH а " ay brachycentrum Ledeb. 16 Langlet, Brunonianum Royle 16 IE UY Propach, 1940 bulleyanum Forrest 32 Gregory, 1941 californicum T. & G. 16 Lewis е? al, 19 cardinale Hook. 16 Bonk 1940; Mehlquist, 1943; Lewis e£ = Ре cardiopetalum D.C. 16 Tjebbes, Pe Lewitsky, 1931; Gregory, 1 arolinianum Walt.* 32? Gregory, casbimirianum Royle Propach, 1940 саисаѕі Меу. 16 Gregory, 1941 cheilanthum Fisch. ex DC 16 Propach, 193 Consolida L. 16 Hocquette, ЧИ т) ebbes, 1927; MP 1927; 1; Gregory, 194 cuyamacae Abrams 16 Lewis e£ al, ч decorum Fisch. and Mey, ssp. Tracyi Ewan 16 Lewis e£ al, 1951 Delaveyi Franch. 16 Lewitsky, 1931; Propach, 1940 Dubmbergi Huth 32 Lewitsky, 1931 16 Propach, or 32 Lewitsky, 1931; Propach, n O 1941 e ак. & Ки. 32 сон, DAS Propach, 1 flexuosum Raf. 32 Gregory, 1941 formosum Boiss. & Huet 32 Эзел 1941 gayanum Willmott 16 Gregory, 1941 glaucu $. 16 Lewis e£ al, 1951 gracilentum Greene 16 Lewis ef al, 1951 grandiflorum L. 16 Propach, 1940 dp Qu Mr >. — ener iier igo: has determined these American species as diploid. His studies of plants tive habitat, whereas the other determinations were probably made from material ae out ү т European botanical gardens, which often con fuse D. elatum with D. carolinianum and D. azureum. 1953] GAGE—DELPHINIUM Х BELLADONNA HORT. 165 Species 2n number Author and date үе Ewan 16,32 | Lewis e£ al, 1951 p pem ene 16,32 | Lewis e£ al, 1951 besperium 16 Lewis e£ al, 1951 besperium н Зог Ewan 16 Lewis e£ al, 1951 iliense Huth 16 Propach, 1940 inopinum (= D. parishii var. inopinum Jeps.) 16 Lewis e£ al, 1951 Lamartine Hort. 48 Lawrence, 1936 Moerbeimii Hort. 24 Lawrence, 1936 nudicaule Torr. & Gray 16 Tjebbes, 1927; viri 1940; Gregory, 1941; Lewis et al, 1 Nuttallianum Pritz. 16 Lewis e£ al, 195 Т ientale (?) orientale Losc.; ‚ orientale S. Gay) 16 Beckma = 1928; n 1941 Parisbii Gray 16 Lewis, 951 Parryi Gray 16 Lewis, т al. 1951 Parryi ssp. seditiosum Jeps.) Ewan 16 Lewis, т 1951 batens Benth. 16 Lewis, 1951 Penardi Huth 16 Ск, ч ЪЪ 16 Gregory, 1941 bictum Willd. 16 Gregory, 1941 bolycladon Eastw. 16 Lewis e£ al, 1951 Purpus dg 16 Lewis e£ al, 195 recurvatum Greene 16 Lewis e£ al, 1951 Ruysii Ho 32 Lawrence, 1936 scopulorum Gray 16 G 1941 Specios ?) Ф speciosum Janka — m L.; ; speciosum "Bieb. 16 Langlet, 1927 Stap bis sagria L. ! 16 унво A и Langlet, 1927; Lewitsky, 1931; ; Gregory, ips sulph cn Boiss. & Hausskn 16 Gage, this p fatsienen е Franch. 16 Lewitsky, 1931 Propach, 1940 tric corne: 16 Gregory, 1 trolliifolium Gay 16 Lewis e£ x ibi truncatum Lang. 16 Langlet, 1927 Ашын, Curran 16 Lewis e£ al, 1951 umbraculorum 16 is e£ al, 1951 айн ots Torr. & Gray 16,32 | Lewis ef al, 1951 Yunnanense Franch 16 regory, 194 Zalil Aitch. & Hemsl. 16 Gage, this paper ee ee ee ence CHROMOSOME NUMBERS OCCURRING IN THE HYBRIDS DETERMINED BY H. PROPACH, 1940 с Variety Breeder or Source Chromosome Number п. Eisberg E. Benary, Erfurt 16 Blauglut K. Foerster, Bornim 24 Blautanne » 24 Al e x = Berghimmel Ж » 1а 4 » Blickfang x 166 [Vor. 40, 1953] ANNALS OF THE MISSOURI BOTANICAL GARDEN in blaues Wunder De Enzianherold Enzianturm Ernst von Borsig Fón ;lasfenster Gletscherwasser Gnom Grossenwahn Gute Nacht ichflam Quem Clothilde empelgong aei ad У 1 jov musoz g ~ پا‎ p Fig. 2. Delphinium Hendersoni, from Revue Horticole. 1854. Fig. 1. Delpbinium cheilanthum Rariores Horti Academici i Monacensis,’ pl. 52. K. Foerster, Bornim ээ EXPLANATION OF PLATE PLATE 7 Fisch., PRS from von Schrank, ‘Plantae 1819 PLATE 7 ANN. Mo. Bor. Garp., Vor. 40, 1953 رر ررر As‏ a PS PS EE a Ki GAGE- DELPHINIUM BELLADONNA HOR T. PLATE 8 ANN. Mo. Вот. Garp., Vor. 40, 1953 PHINIUM roawosum He 170 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE | Qu PLATE? — Figs. 1 and 2. Habit and détail of inflorescence of Delbbiniu cheilan ex DC., grown at the Missouri Botanical Garden rom seed received from U Figs. 3 and 4. Habit and detail of E srendif lorum Loy г. chi Fisch. "Blue Butterfly. €— mes emen GAGE—DELPHINIUM X BELLADONNA HORT. 6 ALVIg ESEL ‘OF “IOA *auvo ‘104 ‘OW ‘NNY 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 10 ОИЕ. Delphinium elatum L. Plant pet SUE tri ploids. PLATE 10 "^ wn = = + E Ке > а “ < O ы о 25 0 — 2; Z z < DELPHINIUM Р І HLOR ANNALS OF THE MISSOURI BOTANICAL GARDEN | И T ANN. Мо. Вот. Garp., Vor. 40, 1953 PLATE 11 GAGE—DELPHINIUM X BELLADONNA HORT. E д a Farad? an Habit ма Summer Skies," seed paren | Figs. 3 and 4. Inflorescence and detail of triploid 50- ANNALS OF THE MISSOURI BOTANICAL GARDEN ст ALVIg €£€61 ‘OF "IOA “AUVO ‘104 ‘OWN ‘NNY 3 BELLADONNA HORT. GAGE—DELPHINIUM x 2 [VoL. 40, 1953] 178 ANNALS OF THE MISSOURI BOTANICAL GARDEN d EXPLANATION OF PLATE PLATE 13 ey X 1600; camera-lucida drawings, X 45 Fig Ideogram of the haploid chromosome complement of D. ООО v PLA ла form "Blue Butterfly. Fig. 2. Ideogram of the haploid chromosome complement of D. cheilanthum. Fig. 3. Photograph of somatic metaphase of a white variety of D. grandiflora var chinense, showing satellites on one C and two H's. Fig. 4. Photograph of somatic аи, of the same plant, showing heterocteon tll regions as well as three of the satellite Fig. 5. Somatic metaphase of D. cheilanthum, showing the four satellited c somes. Fig. 6. Camera-lucida drawing of the cell in fig. 5. 18. 7. Photograph of metaphase chromosomes of "Blue Butterfly" from a root-tip preparation. ig. 8. Camera-lucida drawing of somatic chromosomes of "Blue Butterfly" showing satellites on both C's and both H's. it 9. Camera-lucida drawing of somatic metaphase of 47031-1, showing fra. 10. Somatic rompe of D. cardinale showing satellites оп six chrom of d diploid complemen Fig. 11. Drawing of the cell in fig. 10. ANN. Мо. Вот. GARD., Vor. 40, 1953 PLaTE 13 ABCDEFGH| ABCDEFGH ST eren ППШ | GRANDIFLORUM | CHEILANTHUM GAGE—DELPHINIUM X BELLADONNA HORT. [Vor. 40, 1953] 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 14 Magnification of photographs as indicated; camera-lucida drawings, X 1350. Fig. 1. Photograph of somatic chromosomes of 48-27-2. X 1500 u 2. Drawing of the cell in fig. 1. Camera-lucida drawing of the pt in a root-tip preparation of 48- 6-2, p Nee satellites on one A, three C's, and one H Fig. 4. Drawing of un somatic chromosomes of "Summer Skies," showing satellites on two A’s, two C's, and two H's. 5. Camera-lucida drawing of ke m pex му him of tetraploid 47-98-3. NE visible on two A's, three C's, and t Fig. 6. Photograph of the cell in fig. 7. X 1500. Fig. 7. Camera-lucida drawing of MI of triploid 48-27-3, showing 8 II and 8 I. Fig. 8. MI of triploid 48-27-5, showing maximum amount of pairing observe ed: EN 5 П, and 7 I. Five bivalents of the intermediate chromosomes have two chiasmata. Fig. 9. MI of triploid 48-27-5, showing greatly reduced pairing: 5 П and 14 I Fig. 10. Pollen of "Cliveden Beauty." Pollen mostly viable. Х 350. ie 11. Pollen of triploid 48-6-1, showing one unreduced grain. X 350. 12. Pollen of 50-20-1, an п between Smith's Belladonna and triploid 48-6- 2, NADA aborted pollen grains. X 3 ANN. Mo. Вот. Gard., Vor. 40, 1953 PLATE 14 GAGE—DELPHINIUM x BELLADONNA HORT. [Vor. 182 ANNAIS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PEATE 15 Photographs, X 1600; camera-lucida drawings, X 1450. Fig. 1. Diakinesis and early metaphase I of "Blue Butterfly," showing eight. Fig. 2. Photograph of the somatic chromosomes of D. Belladonna. Fig. 3. Anaphase I of 47031-1 with lagging fragment. Fig. 4. Diakinesis of "Summer Skies," showing essentially regular pairing Fig Photograph of MI of Smith's Belladonna, showing eight univalen two of da la arge bivalents with two chiasmata. This was among the most extreme of lack of pairing observed in this plant. Fig. 6. Drawing of the cell in fig. 2, showing Deli a one А, six c one Fig. 7. Camera-lucida drawing of diakinesis of Smith's Belladonna, showing six valents. : e Fig. 8. Camera-lucida drawing of the cell in fig. 5. ANN. Мо. Вот. GARD., Vor. 40, 1953 PLATE 15 GAGE—DELPHINIUM X BELLADONNA HORT. OF THE MISSOURI BOTANICAL GARDEN = | Emeritus Director — d SA ; ; GEORGE T. Moore AME ees ` Acting Director т наз шик ы чс, Тонм S. LEHMANN | EINE < Assistant Director —— Pow I EDGAR ANDERSON © "ucc = Executive Assistant pir LEICESTER B. Faust 4 ps ROBERT Е. Woonson, da | _ — Curator of the Herbarium —— Henry N. ANDREWS, ROLLA М. Taxon, | | = Elmer's “Leaflets of Philippine Botany" АН the available sets of Elmer’s “Leaflets of Philippine Botany” have been deposited at the Missouri Botanical Garden. Several complete sets are available. Each set consists of ten volumes and an index, unbound, a total of 3,936 pages of descrip- tions of Philippine plants, many of them new species. Each set $20.00 plus postage. Volumes from the few incomplete sets, $2.00 plus postage. Missourr BOTANICAL GARDEN 2315 Tower Grove Ave., St. Louis 10, Mo. Number 3 Annals of the Missouri Botani Annals of the Missouri Botanical Garden E Quarterly Journal containing Scientific Contributions from the ouri Botanical Garden and the Henry Shaw School of Botany of a University in affiliation with the Missouri Botanical Garden. Information The $ OF THE Missoumr: BOTANICAL GARDEN appears four times = the cider year: TON May, September, and November. Four umbers constitute a volum Subscription Price __ пс per volume Single Numbers 2.50 each Cont of previous issues of the ANNALS OF THE Missouri BOTANICAL Gaam a are е Бат in the Agricultural Index, published. 5 the H. W. Wilson Company. Met west Ten i TET E a oerte a DULCE a рода: И КЫШ сы а ы Ө, ы РОТЕ EUR P р дани EET Va с a ur v LC В | Th! МАТЕЈЕ ui Annals of the Missouri Botanical Garden Vol. 40 SEPTEMBER, 1953 No. 3 SOME AMERICAN MEDULLOSAS? HENRY М. ANDREWS лмо SERGIUS H. MAMAY? The continued discovery of new species of the polystelic stems referable to the genus Medullosa emphasizes the conspicuous role that these plants played in the Pennsylvanian floras of the United States. Almost every large collection of coal balls that is studied reveals a specimen or two that differs in some way from those previously described. Although it has been impossible to define species limits with definite clarity when the specimens are fragmentary, poorly preserved, or represent only a short segment of the stem, it seems more than likely that most recorded American species are valid. Those of us who have worked with these plants assumed some time ago that аз the sources of material were more extensively explored our efforts would tend _ toward working out fine details rather than reporting ever more bizarre forms. Actually the reverse has proved to be the case, and the occurrence in recent col- lections of specimens of greater stem length has permitted more satisfactory study of variable characters such as stelar number. One of the most notable contributions to our knowledge of the Medullosas is the recent description of M. heterostelica by Stewart and Delevoryas (1952). Although it is а bi-stelar stem the two steles divide in the nodal region to form а complex of twenty or more, some of which contribute to the petiole traces, whereas others reunite, the stem becoming bi-stelar again above the node. Ha this specimen been found in a more fragmentary condition, descriptions of several “species” might well have resulted. As it is, the evidence suggests that stelar number is a reliable taxonomic character provided that a representative specimen of the stem is available. The present account deals with several Medullosa specimens that have come to light in the paleobotanical laboratory of Washington University since Baxter’s contribution in 1949. Certain of these add to our knowledge of the distribution | Of previously described species; others are distinctly new and present what we —— z Publication authorized by the Director, U. S. Geological Survey. U. $. Geological Survey, Washington 25, D. C. (183) Issued September 30, 1953. [Vor. 40 184 ANNALS OF THE MISSOURI BOTANICAL GARDEN believe to be highly significant information bearing on the problem of the tax- onomic significance of stelar number. MEDULLOSA THOMPSONI Andrews Two specimens referable to this species are present in our collections; both come from southeastern Kansas. One (No. 721) presents three steles (fig. 2), measuring 5 X 2.5 mm., 6 X 2.5 mm., and 2 mm. in diameter respectively and compares closely with the type specimen described from Iowa (Andrews, 1945). The second specimen is of interest because of its length and the type of stelar changes that occur. This stem, found in coal ball No. 713, is approximately 23 cm. long. Мо recognizable plant remains were evident on the exterior of the coal ball to guide the initial cut, and by chance the stem was dissected in a nearly median longitudinal plane. This seemed unfortunate at first, although it has materially aided us in following the changing stelar morphology through the length of the specimen. The plane of this initial cut accounts for the conspicuous break in certain of the cross-section illustrations (figs. 5-7). Text-figures 1 and 2 show, respectively, a profile of the coal ball and a sketch of the stelar system indicating the divisions and fusions that take place. Їп the A1-2 and A2-2 blocks three steles (fig. 6) are present. These are enclosed by a clearly defined internal periderm that does not exceed 0.5 mm. in width and con- sists of rather thin-walled nearly cubical cells that present essentially the same appearance in transverse and radial sections. The steles at this point measure 15 X 9.5 mm., 11.5 Х 6 mm., and 20 Х 7 mm. In passing through the A1-1 and A2-1 blocks two of the steles fuse; the resultant two steles (fig. 7) measure 14 X 7.3 and 21.5 Х 8 mm. Further divisions and fusions are indicated in text-fig. 2 as follows: In passing through the B blocks a stele divides, revealing three steles on the upper surface; in passing through the C blocks a fusion takes place, revealing two steles оп the lower surface of D1 and D2; finally a small stele separates from one of the two steles, revealing three steles on the lower sur- face of the E blocks. Although the phloem is partly preserved, all stelar dimen- sions refer to the wood only. The secretory canals, which average about 0.3 mm. in diameter, are confined chiefly to the fiber zone of the cortex. The primary body (fig. 3) contained a considerable amount of parenchyma, although this has decayed for the most part, leaving only the primary tracheiós. The departing primary petiole strands are large and accompanied by little or no secondary wood. Through the length of this specimen there are at least els petioles that depart from the stem. It is especially important to note this, — obviously the type of nodal anatomy here is quite in contrast with that described for Medullosa heterostelica. This point will be considered in further detail under the description of the specimen that we are assigning to M. heterostelica. The affinities of this stem seem to lie closest to Medullosa distelica, M. anglict and M. thompsonii, but particularly the last. It differs from M. distelica in that 1953] ANDREWS AND MAMAY— AMERICAN MEDULLOSAS 185 у Тех Lt. x Profile of Ee ball No. fe contain- ing specimen of Medullosa thompson. Numbers han kon of blacks i in pesca wa jme 1). [Vor. 40 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN it presents a less constant association of secretory canals and fiber strands; an internal periderm which attains only about one-third to one-fifth the thickness of that of distelica; and in the stelar system the endocentric development of the secondary wood is not nearly so strongly pronounced nor as constant in our specimen. In comparing it with M. anglica it also appears to possess less abundant secretory canals associated with the fiber strands (Scott, 1899, pl. 12, fig. 14), somewhat smaller steles, and the secondary wood around the departing leaf traces is not retained in the Kansas fossil. Almost every Medullosan stem discovered in American coal balls display: characteristics which tempt one to set up a new specific entity, or which at least render it difficult to assign the specimen with confidence to an established species. e four species assigned by Schopf (1939) to his subgenus anglorota as well as M. thompsonii (Andrews, 1945) and the species described more recently by Bax- ter (1949) are certainly all closely related. With some species the relationship is so close as to render very problematical the dividing line between species and varieties; we refer here to such specimens as M. anglica and its varietal forms (vat. thiesseni, Schopf, 1939; var. ioensis, Andrews and Kernen, 1946), M. thompsonii, М. elongata, and М. distelica. Distinctive characters are somewhat more рго- nounced in certain others such as M. endocentrica and М. primaeva. The taxonomic treatment of the specimen described above is admittedly some- what arbitrary. Its affinity lies with the anglica-distelica-thompsonii complex, and the most expedient way of handling it appears to be to assign it to the last species mentioned. It differs from previously described specimens of this species only in its somewhat larger size. It is of real interest, however, for the informa- tion it conveys regarding the stelar anatomy. A seemingly significant length of stem is present to give some indication of what characters may be taxonomically dependable. Some variation in the centricity of the secondary xylem is evident, but it could at no point be closely compared with the extreme excentricity foun in M. distelica and M. endocentrica. Within certain limits, which admittedly cannot be defined quantitatively, the degree and form of excentricity in Medullosa stems appear to be significant. Locality: Strip mine of the Pittsburgh & Midway Coal Co., approximately four miles south of West Mineral, Kans. Horizon: Fleming Coal, Cherokee shale, Des Moines group, middle Pennsyl- vanian. MEDULLOSA DISTELICA Schopf Another Medullosa specimen (No. 688) from the West Mineral, Kans., locality appears to be referable to this two-stelar species described by Schopf (1939) from the No. 6 coal of Illinois. It is basically a two-stelar specimen but presents certain variations which seem significant in furthering our knowledge of what may be considered to be specific boundaries in the genus. 1953] ANDREWS AND MAMAY—AMERICAN MEDULLOSAS 187 | = Ј | | m i pec & A tei © о ELLE s] E | nd F | D ee 8 | үн а ы) : : i га B 5.5 3. Profile of coal ball No. Text-fig. 4. Profile of coal ball No. 81 men of Medullosa containing specimen of Medullosa betero- umbers indicate thickness of blocks ers. distelica. Numbers indicate thickness of stelica. N the blocks in centimeters. i i It has been possible to follow the stem through a distance of approximately 10 cm. (text-fig. 3). The stelar system and the enclosing periderm are quite well Preserved, but the cortical tissues have been distorted and a considerable amount lost. In block A the stem is composed of two strongly endocentric steles (66.4). In the stem's course through the lower part of block B a small stele separates from one of the original two, producing a three-stelar system. After another ^ cm. the two-stelar phase is resumed, but since the preservation is poor beyond this point it has not been possible to determine whether any other changes take place, : The two principal steles measure about 18 X 6 mm. and 14 X 5 mm.; taking into account the obvious crushing, they measured approximately 18 X 8 mm. and [Vor. 40 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN 14 Х 7 mm. in life. The small branch stele is nearly round in cross-section, measuring 5 mm. in diameter. The primary body of each stele consists of scattered groups of tracheids and abundant parenchyma (fig. 9). А similar ratio occurs in Medullosa endocentrica (Baxter, 1949), presenting a rather striking contrast to certain other species such as M. primaeva and M. thompsonii. In the type specimen of M. distelica (Schopf, 1939) the preservation of the primary wood was poor. The position of the protoxylem elements was not deter- mined; and even in the well-preserved primary wood of the present specimen it is not possible to locate with certainty any regular position of protoxylem ele- ments, so few and scattered are the primary tracheids. The organization of secretory canals and fiber strands in the outer cortex or rind seems quite distinctive. An especially well-preserved portion of a petiole is shown in fig. 17, although essentially the same relationship may be observed in the stem. Starting at the outside it may be noted (fig. 10) that the secretory canals reach to within less than 1 mm. of the periphery. Here they are minute, the smallest being about 0.1 mm. in diameter, and are closely associated with the fiber strands. The canals increase in diameter toward the inside, reaching a maximum of approximately 2.5 mm. Within the rind proper the canals are conspicuously enclosed on three sides by a fibrous sheath. Only the outermost canals have an inconspicuous sheath or none at all. Locality: Strip mine of the Pittsburgh & Midway Coal Co., approximately four miles south of West Mineral, Kans. Horizon: Fleming Coal, Cherokee shale, Des Moines group, middle Pennsyl- vanian. MEDULLOSA HETEROSTELICA Stewart & Delevoryas Shortly after Stewart and Delevoryas (1952) described M. heterostelica from West Mineral, Kans., the present writers collected from the same locality a coal ball (No. 819) containing a stem which displays strikingly similar gross mor- phology. The stem has two steles which divide into more than fourteen, presum- ably in the nodal region. It differs from the type specimen, however, in two ways: 1. The two original steles in No. 819 measure about 16 mm. in diametet (xylem only) and are radially symmetrical with reference to both primary and secondary wood. The steles of the type specimen measure 5 X 8 mm., being less than one-half the diameter of those in our specimen. 2. The over-all size and form comparison is also reflected in the primary wood. The ratio of tracheids and parenchyma in the primary wood of the two seems to be comparable; but in No. 819 the size of the tracheids is conspicuously greater, and the primary body is nearly circular, whereas that of the type specimen is elongate. A wholly satisfactory comparison of the two is impossible since the cortical tissues and leaf bases of our specimen are not preserved. It is, therefore, only on 1953] ? ANDREWS AND MAMAY—AMERICAN MEDULLOSAS 189 the characters cited above that a decision as to specific identity can be made. The chief point of interest that the specimen presents is the remarkable multi- plication of steles in what we assume is the nodal region. This is in notable con- trast to the stelar anatomy of any other species of Medullosa, and it is of course important to determine whether it is a specific character in itself. The evidence to date indicates that some species did not follow this pattern, whereas with others (based on short or fragmentary specimens) we cannot be certain. A profile of the coal ball is shown in text-fig. 4. The stem appears about mid- way through block A and runs through the remainder of the specimen, totaling about 15 cm. As an introduction to a more detailed description it may be noted first that the stelar system consists of two steles, each of which gives off numerous midway through block A two steles make their appearance (fig. 11). They measure approximately 16 mm. in diameter, are essentially circular in cross- section, and have uniformly developed secondary wood. The primary wood con- sists of tracheids with a few parenchyma cells and measures about 2.5 mm. in diameter. Other than fragmentary pieces of the cortex, no tissues are preserved outside the xylem. These two steles are arbitrarily designated А and B for the purpose of identification in this description. The two steles remain unchanged until about midway through block D, at Which point (text fig. 5-2) a stele departs from A. Another small separate stele may also be noted here; this and several others were observed in a peripheral Position and were not traced to an attachment with A or B; lack of attachment is due apparently to faulty preservation. At a slightly higher level, midway through block E, several branch steles are now evident (text fig. 5-3). These rep- resent additional branches from А and B and the divisions of branches themselves. It will be noted from the text figures as well as the photographs that many of the steles are only partly preserved. Most notable is the gradual loss of stele A. The preservation is poor at the level shown in text-fig. 5-5; at the level of text- fig. 5-5 only half of it is present, and by the time block J is reached (text-fig. 5-7) it has disappeared. Stele B retains essentially the same size and form through the specimen and is at all times dominant in relation to its branches; it seems likely, judging from the portion of it that is preserved, that stele A followed the Same pattern. The preservation in block H is very poor, and little of significance remains beyond block J. For the most part, the branch steles shown on the left side of the text-figures come from stele A and those on the right side from stele B. A union of the two "Systems," however, is evident in text-fig. 5-6 in the form of the elongate stele in the upper part of the figure. : The maximum number of steles present at any one point can be determined only approximately, because of faulty preservation and in part because of the Prolific anastomosing nature of the system. For example, in text-fig. 5-6 a more [Vor. 40 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN WIP к Text-fig. 5. Medullosa beterostelica. Transverse diagrams of specimen No. 819 showing changes in the stelar form through blocks C-] (see text-fig. 4). Several of the poorly preserved steles are omitted. 1, C-t2; 2, D-t17; 5, E-t14; 4, F-t16; 5, F-t9; 6, G-t1; 7, J-t1. or less 7-shaped stele may be noted encompassing the right side of the figure whereas 16 mm. above (text-fig. 5-7) this has divided into five or six steles. At the latter point, which may be taken as representative of the more highly divided part of the stem, there are about fourteen whole or fragmentary steles. So poorly is the stem preserved beyond block J that we are unable to determine whether or not it passes back into the two-stelar phase as reported for the р specimen. The lack of cortical tissues and petiole bases also makes it impossible to determine which steles contribute directly or indirectly to the petiole trac system. 1953] ANDREWS AND MAMAY—AMERICAN MEDULLOSAS 191 The only previously described Medullosa specimen with which this may be com- pared closely is M. beferostelica. As stated above, it differs most notably in size, the two central steles being twice as great in diameter as those of the type speci- men. Assuming that the multiple stelar condition in our specimen is also a nodal region, the size contrast is comparable here. This region is described as being approximately 5.5 cm. long in the type specimen (Stewart and Delevoryas, 1952, p. 508, figs. 5 and 12). In our specimen the multiple-stelar phase starts in block D and runs through J, a distance of 9.3 cm., and undoubtedly ran for several centimeters more if indeed it did again assume the two-stelar phase. Aside from the size there is nothing on which to base a specific difference, and for that reason our specimen is assigned to M. heterostelica. Locality: Strip mine of the Pittsburgh & Midway Coal Co., approximately four miles south of West Mineral, Kans. Horizon: Fleming Coal, Cherokee shale, Des Moines group, middle Pennsyl- уашап. MEDULLosa grandis sp. nov. The most unique medullosan specimen that we have acquired in recent years is No. 718 from the West Mineral, Kans. locality. It is especially interesting by virtue of its large size and by its numerous steles which anastomose freely and present much anatomical variation. With the exception of Medullosa moei it is much larger than any of the previously reported Carboniferous species. The specimen composes the major portion of a coal ball about 30 cm. long and 28 X 12 cm. in its largest transverse dimensions. The coal ball tapers to a very blunt point at the А end (text-fig. 6) and the stem decreases correspondingly in size; that is, the decrease is normal and not due simply to loss of part of the stelar system. This small end is also heavily pyritized and less well preserved than the rest of the specimen. The other end is represented by a broken surface. If, as seems likely, this specimen constitutes the apical portion of a stem, then it clearly represents a tree of some magnitude; and if the stem continued to increase in diameter for an appreciable distance at the rate shown in our fragment it must ave soon attained the dimensions of a large forest tree. In order to present a clear picture of the complex organization of the stelar system a restoration of the major portion of it is shown in text-fig. 7. This is accompanied, in text-fig. 8, with diagrams of the system at the four points indi- cated. All the numerous smaller steles that show in text-fig. 8 are not included in the restoration (text-fig. 7). Text-fig. 6 is a profile of the coal ball, which may be referred to for the position of the text and plate figures. A casual glance at the text-figures reveals considerable variation in the size and shape of the steles as well as the relative amount of primary wood that is Present. A striking feature is the great variation in the structure of the secondary Wood itself, in which the different ratios of parenchyma and tracheids present distinctly atypical steles. It is of course likely that many of the smaller strands [Vor. 40 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN A-4 4 A-3 3 a LER Text-fig. 6. Profile of coal ball No. 718 containing the specimen of M. grandis. Numbers in the right column indicate thickness in centimeters. contribute to the vascular supply of the petioles. Of necessity we are therefore using the term “stele” to include all the vascular strands with secondary W (secondary phloem is present but poorly preserved). In tracing the over-all pattern of the stelar anatomy it is significant to note that it is composed of two fairly distinct systems, thus following in a general o the type of organization described for Medullosa beferostelica. Stelar branching and fusion take place so rapidly that the system presents an intricate anastomosing network. The block is so large that it did not seem practicable to run a complet? 1953] 195 ANDREWS AND MAMAY— AMERICAN MEDULLOSAS um or iui \ 1 ! ^ A restoration of the stelar system и of Medullosa grandis. Text-fig [Vor. 40 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN series of sections through the specimen. Text-fig. 7 presents, therefore, a slightly less complex picture than that actually represented by the number of steles involved and the branching pattern. The numbers assigned to the individual steles in text-fig. 8 A—D are arbitrary and are intended to facilitate following the steles through the portion of the stem that is represented. Те seems somewhat simpler to start with the smaller end, presumably the apex of the stem. We may first follow the course of steles 1, 2, and 3 from A to B, where steles 1 and 2 have fused and stele 5 is evidently pre- paring to divide. In C the three steles are shown fused into a single irregular- shaped stele. Finally in D this has now divided into four distinct steles. These fusions and divisions take place through a distance of approximately 10 cm. Returning to text-fig. 8-A we may now trace through the second "system" of steles, represented here by numbers 5, 6, 7; these also increase markedly in size in passing through B to C. Because of faulty preservation it was not possible to interpret precisely the changes that ensue below this point. At least one branch stele departs from number 5, and in D steles 5 and 6 are fused to form a very irregular-shaped wood mass. There can be little doubt that the numerous smaller steles shown to the right are a part of this stelar "system," but they have not been traced in actual connection. . In addition to the more conspicuous steles considered above there are numer- ous smaller ones all of which display a remarkable range of variation in their organization. Certain of these features may now be considered. Figure 13, which represents a single stele from the cross-section shown in fig. 1, may be taken as typical of the organization of the larger "normal" vascular elements of the stem. The stele is slightly crushed, although it is evident that there was no great abundance of primary wood. In fig. 14 is illustrated a portion of another stele of generally comparable dimensions. It is evident here that the primary wood is composed chiefly of tracheids with little admixed parenchyma. There is nothing exceptional about the organization of the secondary wood of the stele shown in fig. 13. The pitting in the radial walls of the tracheids is typically pteridospermous, and the rays are rather narrow and of great height. Several of the larger steles compare with the one described above, but others vary much in their course through the stem with respect to the relative abundance of parenchyma and tracheids, and this difference may be noted in both primary and secondary wood. Figure 16 shows a portion of the large stele at the upper right of fig. 1. Te central region of this stele is occupied by parenchyma exclusively, whereas the im- mediately surrounding secondary tissue consists chiefly of parenchyma with a few radially aligned rows of tracheids, a feature that is common in the secondary xylem of some of the large Permian Medullosas. Toward the periphery of : stele the organization compares closely with that observed in the stele shown 3? fig. 13. It should be kept in mind-here that the stele shown in fig. 16 is directly connected with the “normal” stele shown in fig. 13. 1953] ANDREWS AND MAMAY—AMERICAN MEDULLOSAS 195 сь & aR S n 718 through blocks A-C (See text-fig. 6). Text-fig. 8. Medullosa grandis. Transverse diagrams of specime showing changes in the stelar form А, A-t2; B, B2-t1; C, В1-с1; D, C-tl [Vor. 40 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN Another predominantly parenchymatous stele is illustrated in fig. 15. Here there is a large pith composed chiefly of parenchyma but with some groups of tracheids scattered through it. The tracheid-parenchyma cell ratio in the second- ary wood is variable, as is indicated in the figure. A small branch stele preparing to depart is shown at B. : Most of the extra-stelar tissues of the specimen have been lost, but the frag- ments of the characteristic medullosan internal periderm suggest the enclosure of the stelar system here as in other species. There is a rather poorly preserved groundwork of parenchyma, and scattered through this are numerous secretory canals, each enclosed by a "periderm" ring. Several small traces are present, also enclosed in the same tissue (fig. 8). There is no evidence suggesting that any portion of this specimen represents a nodal region. Diagnosis: Stem large, greatest known traverse dimensions of the stelar sys- tem 12 X 28 cm. Steles numerous (18—20), extremely variable in size (up to 6 cm. in diameter), shape, and xylem-parenchyma ratio. Steles anastomosing frequently, but generally arranged in two systems, one somewhat more complex than the other. Extra-stelar organization unknown. Locality: Strip mine of the Pittsburgh & Midway Coal Co., approximately four miles south of West Mineral, Kans. Horizon: Fleming Coal, Cherokee shale, Des Moines group, middle Pennsyl- vanian. Type specimen is deposited in the Paleozoic plant collections of the U. $. National Museum. Medullosa noei Steidtmann Our recent coal-ball collections from southern Illinois include two specimens of the large tri-stelar М. noei (Steidtmann, 1937, 1944). They are approximately 43 cm. and 30 cm. long; although this is intended only as a preliminary account, since the two have not been studied critically, they appear to follow the general stelar pattern described above for M. thompsonii. They show no evidence of the multiplicity of stelar branching and fusion found in M. heterostelica and M. grandis. One of the specimens showing the complete stelar system is illustrated in fig. 18; the original specimen was a fragmentary one, although the concept of its stelar organization (Steidtmann, 1937, fig. A) is confirmed by our specimens. Discussion: Several contributors to the literature on the Medullosas have been tempted zo speculate concerning the relationships of the now numerous species included in the genus. A total of no less than 44 species and varieties have been recorded to date and since the stelar anatomy is nearly as diverse as the number of entities recorded it is understandable that paleobotanists find it intriguing to propose evolutionary lines in the group. Although the primary purpose of this paper is to present 1953] ANDREWS AND MAMAY— AMERICAN MEDULLOSAS 197 what appear to be significant factual contributions, a few speculatory comments seem to be in order. One point of anatomical significance is well illustrated by the assemblage of specimens discussed in this paper, namely, the contrast in the relative constancy of stelar number among certain of the better-known species. Medullosa endocentrica Baxter may be taken as a striking example of a species in which the stelar number remains constant. The single available specimen was some 12 cm. long, a significant length considering the small diameter of the plant. In this specimen the tri-stelar form remains constant throughout its length, there being no evidence of stelar fusion, division, nodal complexities, or notable changes in shape or size of the individual steles. Based on observations of a specimen some 23 ст. long, M. thompsonii also displays a relatively simple stelar pattern. There is some stelar division and fusion; this, however, is not apparently related to the position of the nodes, and the number of steles varies only from two to three (text-fig. 2). The stelar complexity of M. grandis and M. heterostelica presents, at least on casual observation, a striking contrast to the organization found in M. endo- centrica and M. tbompsonii. 'The contrast may cause one to question the inclusion of all four species within one genus. The complex stelar form of M. grandis and M. heterostelica does, however, mask an apparently basic pattern of organization. We have demonstrated that, so far as preservation allows, our specimen of M. heterostelica contains a two-stelar system. That is, the two “original” steles each divide profusely, but the two resultant groups of steles remain more or less dis- tinct. In М. grandis, also, the stelar system appears to be organized into two &roups, although we are unable to determine whether or not the specimen at our disposal represents a nodal region comparable with that of М. heterostelica. In view of the rather significant size of the specimen of M. grandis, it seems likely that the stem did not possess the two-stelar internodal simplicity of M. hetero- Stelicg, One point can be stated with certainty concerning the interrelationships of the Medullosas is that we can guess, and guess only vaguely, concerning possible lines of development. The extent to which competent workers have disagreed in their Concepts of racial development in the genus is indicated by comparing the Phyletic chart of Baxter (1949, p. 309) with that presented by Stewart and Delevoryas (1952, р. 514). In the latter, Sucliffia is assigned а basal position, Whereas Baxter suggests that Sutcliffia represents a specialized side branch from a Medullosa of the M. anglica type. It is also significant that the discovery of M. heterostelica caused Stewart and Delevoryas (1952) to modify rather radically the Concept presented in Stewart’s earlier paper of 1951. It is thus evident that the discovery of each new species of Medullosa may result in a new phyletic scheme for the genus, particularly in view of the unique ‘pecimens that have been discovered in recent years. Our feeling is that inter- [Vor. 40 198 ANNALS OF THE MISSOURI BOTANICAL GARDEN relationships within the genus are actually much more complex than has been implied by previous workers. Evidence in support of this view is contributed by the recent report of M. olseniae from the Permian of Texas (Roberts and Barg- hoorn, 1952). It is a multi-stelar stem, the steles being arranged in a peripheral group resembling M. leuckarti or M. solmsi, but subsidiary steles are reported to be lacking. It is thus difficult to “fit” it into the top of either of the two branches of the genus indicated in the chart presented by Stewart and Delevoryas. Although we shall refrain from commenting further on the evolution of the genus as a whole, we should like to conclude with a comment on the type of anatomy presented by the grandis-heterostelica complex. It has been assumed by many previous workers that the Medullosas evolved from a single-steled plant of the Heterangium type or one in which a single stele appears dominant, as in Sut- clifia. Referring for the moment to the type of anatomy exhibited by M. grandis, there appear to be at least two possible explanations for its origin: First, in view of the two distinct stelar "systems" represented, Medullosa grandis may be considered as having evolved from a two-stelar ancestor with stelar proliferations resulting in this type of multi-stelar stem. Further modifica- tions of the M. grandis pattern may then have resulted in the diverse types found in the Permian. Then, unless we assume а phyletic return to the two-stelar form, M. grandis may be considered as an advanced species having evolved from the anglica-tbompsonii-distelica-etc. group. Stratigraphic relations are not very help- ful here. The exact position of M. grandis in relation to the species cited above is not known, but it seems likely that M. anglica is of earlier origin, while M. thompsonii and M. distelica are of later origin (see Andrews, 1951, p. 433). | Second is the possibility that M. grandis is a distinctly primitive Medullosa in which we may see the beginnings of dominance being assumed by a few steles. In this case M. heterostelica may represent a species in which the multi-stel phase is confined to the nodal region and in turn the two- to three-stelar species have evolved (as suggested by Stewart and Delevoryas) by the complete loss of the multi-stelar phase. This suggestion next requires some tentative explanation concerning the origin of M. grandis, and it is at this point that we would diverge most radically from previous views. 4 In recent years several poly-stelic plants have been reported from Devonian and lowermost Carboniferous horizons such as Xenocladia (Arnold, 1952), Pietz- schia (Gothan, 1927; and Read and Campbell, 1939), S£eloxylon (Solms-Laubach, 1896, and Read and Campbell, 1939), and Cladoxylon (Bertrand, 1935). Ий not implied that M. grandis may be traced directly to any of these plants, but 1t is suggested that such a stem type may represent а medullosan ancestor. Acknowledgement: This study has been aided in large part by a grant from the Permanent Science Fund of The American Academy of Arts and Sciences. А portion of the work was also carried out during my tenure, at Harvard University, as a Fellow of the John Simon Guggenheim Memorial Foundation. I am sincerely grateful to several institutions.—H.N.A. 1953] ANDREWS AND MAMAY—AMERICAN MEDULLOSAS 199 References: mm H. N. (1945). Sg aah ve to our ae = American те Floras. УП. ера stems from Іоу и Gard z 1 . American coal- ball û ot. T ev. 17:430—4 Kernen, Jules A. e a x our И. of American Carbon- iferous lors. УШ. Aut ммен rom Iowa. Ann. Mo. Bot. Gard. 33:141—146. Arnold, C. A. (1952). Observations on fossil pe from the Devonian of eastern North America. enocladia medullosina Arnold. Univ. Michigan, Contr. Mus. Paleont. 9:297—309. yes Preridosperm. stems and fructifications with particular reference e Medullosae. Ann. M zm 36:287—352 hada Paul (1935). vi win a l'étude des Cladoxylées de Saalfeld. Palaeontographica 1 eee Pflanzen aus dem Oberdevon von Wildenfels. Abh. Süchs. Landesanstalt 3: Read, С. and Campbell S. (1939). Preliminary account of the New Albany shale flora. Amer, Midl. Nat. 21:43 Roberts, David C., and Elso 5: ا‎ (1952). меотов манер a Permian Medullosa from north central Texas. Harvard ‚ Bot. Mus. Leafl. —200. ume. T M. (1939). тта distelice, a new а of d anglica group of Medullosa. r. Jour. Bot. 26 Aid Ee D. H. (1 92), On t не and affinities of fossil plants from the Palaeozoic rocks. Ш. Medullosa endi a new representative of the Cycadofilices. Roy. Soc. London, Philos. Trans. 191:81—12 Solms-Laubach, H. (1896). Ueber die seinerzeit von Unger beschreibenen strukturbietenden Planzenreste > a von Saalfeld in Thiringen. Preuss. Geol. Landesanstalt Abh., Folge 23 Seddemann, Waldo E a 937 7). A preliminary report on the anatomy and affinities of Medullosa у. from the Pennsylvanian of Illinois. Amer. Jour. Bot. 24:124—125. ; > (1944). The PY. and affinities of Medullosa moei Steidtmann, and associated foliage, roots, and seeds. Univ. Mic higan, Contrib. Mus. Paleont. 6:131-166. jf Stewart, Wilson N. (1951). Medulio pandurata sp. nov. from the McLeansboro group of Illinois. Jour. Bot. 38:709-7 ‚ and Fag ыйлаш Theodor (1952). Bases for determing relationships among the Medullo- Saceae, Ibid. 39 200 Fig. Natural t size. [Vor. 40, 1953] ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 16 Medullosa grandis Andrews & Mamay. Transverse section, peel 718-C-tl. PLATE 16 1953 , Ann. Mo. Bor. GARD., Vor. 40 ANDREWS лхо MAMAY — AMERICAN MEDULLOSAS 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN _ Ann. Мо. Bo Garp., Vor. 40, 1953 ^ 3% у 3 УКА SM “ is * ness А2 ә, vee Vesey “© s Дб бе, LI 2% ‘tia ^ aa, к/з, 14; Thay ALME. 2797020655 АННЕ r1 нае iv - ТЕ. DH £ 2; s> рте? "жэ чл, IEEE Fosse” ыр не. ze а = > m Te I Н LIT rs qe ETE EE 7 e Te er ANDREWS Asp MAMAY — AMERICAN MEDULLO = Ls mas PLATE 17 Ann. Mo. Вот. Garp., Vor. 40, 1953 Prate 18 v uda e" РТА =... ANDREWS anp MAMAY — AMERICAN MEDULLOSAS [Уот. 40, 1953] ANDREWS AND MAMAY—AMERICAN MEDULLOSAS 203 EXPLANATION OF PLATE PLATE 18 Fig. 4. Medullosa ранг vies Transverse section of stelar system of specimen No. 688. Peel 688-A-t1 Fig. 5. Medullosa ко Andre Transverse section of specimen No. 713 showing two-stelar phase. Peels 713-D2- bi г 713-01-63. X 4. i Ann. Мо. Bor. Garp., Vor. 40, 1953 PLATE 19 ANDREWS anp MAMAY — AMERICAN MEDULLOSAS Ann. Mo. Bor. Garb., Vor. 40, 1953 PLATE 20 AN, Mi OS Й VEINS + pe, 0/0, (LER IN 2 2 cnn: - „> Pup, «Фал»; ася PE -% геу MN at ANDREWS лмо MAMAY — AMERICAN MEDULLOSAS [Vor. 40, 1953] ANDREWS AND MAMAY— AMERICAN MEDULLOSAS 205 EXPLANATION OF PLATE PLATE 20 . 8. Medullosa grandis. Transverse section of a small vascular strand, presum- چ‎ а Dui trace, surrounded by "periderm." Peel 718-В1-11. 55. Fig. 9. Medullosa distelica. Transverse section of primary wood. Peel 688-C-b2. 60. 10. о distelica. 'Transverse section of the outer cortex of the petiole. ы ne -B-t13. . Medullosa beterostelica Stewart and ЕЕ Transverse section show- ing the two-stelar phase. Peel 819-B-t3. Х 2.4 [Vor. 40, 1953] 206 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION ОЕ PLATE PLATE 21 . 12. Medullosa н Transverse section of the stem showing the multi- stelar phase. Peel 819-G-t3. x 3 PLATE 21 Ann. Mo. Bor. GARD., Vor. 40, 1953 DULLOSAS AMERICAN ME ANDREWS лмо MAMAY „оз ees, "rn 25 1A is EDULLOSA еа REM XA. SS 2 ow Sow RET DEDI edd rA E y + AMERICAN M 5 = "rd \ НАГА vit i: Hie > WWWA C. SECRETUM OQ. cm ui din t arig A PY Liei A ^x AY Я ` SN АЫ A "ridus TION WS AND МАМ ELLO gni Sea T» Фә 40, 1953 S 3 ANDRE #22: EEE: 2 и күм, ү Же ador ні g^ tida М » Vor. е. TARD., MONS G [Vor. 40, 1953] ANDREWS AND MAMAY—AMERICAN MEDULLOSAS 207 EXPLANATION ОЕ PLATE PLATE 22 Medullosa grandis Fig. 13. А single stele. Peel 718-C-t2. X 3. Fig. 14. Portion of a single stele showing primary wood. Peel 718-B2-tl. X 6.5. [Vor. 40, 1953] 208 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 23 Medullosa grandis Fig. 15. A -n stele showing parenchymatous primary and secondary wood. Peel 718-C-t2. X7 Fig. 16. Portion of " ор wood of a large itele showing high ratio of paren- chyma. Peel 718-C-t2. PLATE 23 Ann. Mo. Bor. Garp., Vor. 40, 1953 $ D di zi ms : я А“ m LS ` ANDREWS anp MAMAY — AMERICAN МЕ DULLOSAS + co DM < n © № = z В [22| = 2 T £ x 3 = < | > < 2. < = В . GARD., VoL. 40, 1953 WS А ANN. Mo. Bo [Vor. 40, 1953] ANDREWS AND MAMAY—AMERICAN MEDULLOSAS 209 EXPLANATION OF PLATE PLATE 24 Fig. 17. Medullosa distelica. ‘Transverse section of portion of a petiole. Peel 688- A-t4. X 5.5. B Fig. 18. Medullosa noei Steidtmann. Transverse section of stelar system. Peel 687- "tl. X 0.9. THE DISTRIBUTION OF DIOSPYROS VIRGINIANA L.* HARRY R. SKALLERUP** While the factors responsible for the present distribution of D. virginiana are in themselves not unique, the study of the literature pertaining to the native per- simmon is particularly rewarding. Because the persimmon is both a fruit and forest tree the information obtained is increased two-fold. The combination of botanical, horticultural, and dendrological data answers some of the questions in regard to the determination and interpretation of the many aspects of its life phenomena. Ап attempt will be made here to present the factors affecting the macro- and micro-distribution of D. virginiana and to review and supplement certain aspects of the literature relating to them. The genus Diospyros has a fossil record extending to the Mid-Cretaceous Era, and is represented by leaves, wood, calices, and even fossilized fruit, each having been reported from various locations throughout the world. Species evidently grew in Greenland, Siberia, Canada, northern Japan, Alaska, the British Isles, and in North Africa and Arabia (Berry, 1923). Although no major geographic region of the United States is without one or more fossil representatives of this genus, only one, based upon wood from a Pleistocene deposit in Louisiana, has been desig- nated D. virginiana (La Motte, 1952). Yet it is reasonable to assume, in the light of leaf variation evident within the extant species, that perhaps a few others of the North American forms based on leaf remains and present as far back as the Cretaceous (Berry, 1923) may also be D. virginiana. Indeed, the present distribu- tion pattern of D. virginiana indicates a very long history. In common with other plants of chiefly tropical affinity, it appears to have been dispersed throughout eastern United States from ancient centers in the Appalachian-Ozark highlands (Fernald, 1931). Diospyros virginiana is now known only from the United States, and occurs Within the area depicted in fig. l. Each dot represents a county from which a collection has been reported. The apparent low density of collections in the South- fast can best be accounted for by a lack of records from this area. Thus, although only a few collections are represented from Mississippi, Alabama, and Louisiana, floras of these states list D. virginiana as occurring throughout the area. Moreover, à survey of the commercial stands of persimmon timber in the United States showed these states to be especially well represented (Fletcher, 1915). The critical collec- tions at the periphery of its range are likely to define its extent fairly well, since specimens representing the rare species and new range extension are collected with a frequency disproportionate to their occurrence in the natural vegetation. —— р aster's thesis lt udes so 5 4 А А » Gard Versity of Minnesota. Formerly Graduate Research Assistant, Missouri Botanical Garden. (211) [Vor. 40 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN . j 25 CRAY 2 isotheri Л МП] arent OP OLDER GLACIATION EXTENT OF WISCONSIN М GLACIATION A e 248 © beh. а; 30 inc of hes rainfall Fig. 1. Range of Diospyros virginiana L. Glacial and climatic boundaries drawn after Flint (1945) and Kincer (1922, 1928), respectively. The study of the distribution of a tree such as D. virginiana has its disad- vantages as well as its advantages. Specimens may be reported or collected from trees introduced outside the natural range, and although the trees may be hardy in these sites they could not have survived in nature as seedlings. Also, reports of non-fruiting specimens, particularly in the older horticultural accounts whic ascribe non-fruiting to cold temperatures, may be erroneous due to the fact that D. virginiana is dioecious and some of the trees in question may be staminate. Diospyros virginiana is found growing widely in areas of older glaciation (the Nebraskan, Kansan, and Illinoisan) and in the unglaciated area of the South. the north and west of the older glaciated area D. virginiana appears to be limit by temperature and rainfall, respectively. Few specimens were found north of the 25° F. February isotherm as determined by Kincer (1928). Inasmuch as individual trees transplanted north of the 25° isotherm are able to survive and bear fruit, it seems that the photoperiod is not a factor in flower formation, and there- fore does not limit its distribution (Naudin, 1880; Harrington, 1900; Trabut, 1926; Baldwin and Culp, 1941). It is possible that seedlings in the wild are unable to withstand the cold and to become established in such areas. This is supported by the report of Aikman and Boyd (1941) that persimmon trees planted for soil con- 1Of the series of maps prepared by Kincer (1928), February represents the coldest winter month for a period of 20 or more years. 1953] SKALLERUP—DIOSPYROS VIRGINIANA Г. 213 servation purposes in southeastern Iowa (at the northwest limit of the range) needed a protected site if they were to survive. In determining which factors might account for the western limits of the species, two possibilities appear: both the line of 30 inches of annual precipitation and an increase of 1000 feet in elevation running northward from the Edwards Plateau coincide fairly well with the distribution of D. virginiana. Since the per- simmon is found commonly in the southern Appalachians (Eddy, 1927), it is more probable that the rainfall is limiting. (In the more northern mountains of Penn- sylvania and West Virginia, persimmon is reported to be more rare than in the lowlands—letters from J. A. Small, E. L. Core—but here the lower winter temper- atures and high elevations are probably responsible.) While it is not known at which phase of its growth the moisture requirement may be critical for D. virginiana, in other fruit trees it has been found that an annual precipitation of at least 30 inches is required by mature individuals (Magness and Traub, 1941). While temperature and rainfall thus appear to limit the greater part of the range of D. virginiana, there is a conspicuous absence of specimens from the area of the last or Wisconsin glaciation within the compass of the 25° isotherm. That this area does not differ significantly in climate fails to explain the absence of D. vir- giniana in the later glaciated region. Berry (1923) states that during Pleistocene times the native persimmon retreated far to the south of the ice front, only to spread north again in the “wake of the ice sheet.” With regard for the dangers in assuming that post-glacial expansion is still going on (Deevey, 1949), it would appear that D. virginiana had advanced into the glaciated area, only alternately to withdraw and advance in response to the Wisconsin glaciation, so that now it occupies the northern boundary of the older glaciation (where it is not limited by low temperatures) and has made but slight advances into Indiana, New Jersey, and Connecticut, the areas of the more recent glaciation. The isolated specimens in Indiana, Ohio, and New York (fig. 1) probably represent collections from culti- vated trees, as does a specimen from Massachusetts (not shown on map—letter, R. Rollins, 1933). Baldwin and Culp in 1941 made a cytological study of persimmon from selected stations throughout the range and found that two chromosome races are Present: 2n — 60 and 2n — 90. Considering other members of the genus ( Table T) it appears that the basic number of chromosomes is 15; accordingly the races of D. virginiana are regarded by Baldwin and Culp as tetraploid and hexaploid. Their relative distribution is interesting: the 60-chromosome race has a continuous dis- tribution in the central and southeastern parts of the range, while the 90-chromo- some race occupies a peripheral position throughout the rest of the range. Several additional counts were made in this study. Collections of seeds gathered in St. Louis, Franklin, and Bollinger counties in Missouri, and from Washington unty, Arkansas, were grown in the experimental greenhouse at the Missouri Botanical Garden. Chromosome counts made from aceto-lacmoid root tip squashes [Vor. 40 214 ANNALS OF THE MISSOURI BOTANICAL GARDEN SUMMARY OF KNOWN CHROMOSOME NUMBERS IN DIOSPYROS* Species 2n Source D. discolor 30 Namikawa, Sisa, and Ani (1932) D. texana 30 Baldwin and Culp (1941) D. lotus 30 Namikawa and Higas shi (1928) D. kaki 90 Namikawa and Higashi (1928) D. virginiana 60, 90 Baldwin and Culp (1941) * After Baldwin and Culp, 1941. showed these to have 90 chromosomes—the number reported by Baldwin and Culp in Crawford County, Kans., and Jasper County, Mo. While it would be premature to draw conclusions as to the meaning of the chromosome races in this species, it is of interest to note that the 60-chromosome individuals are found in and about the ancient eastern center of the species. The occurrence of higher polyploids on the periphery of the range is subject to two interpretations: (1) that they are autopolyploid derivatives which have been able to colonize areas not so accessible to the lower polyploid (Cain, 1944; Stebbins, 1950); (2) that the 90- chromosome race is actually of hybrid origin (with an extinct species or race) and is spreading eastward along the northern boundary of tetraploid D. virginiana. In either case, studies of both the morphology and cytology of plants of known chromosome number will be required before the question can be answered even tentatively. In the course of horticultural practice D. virginiana has been introduced into climates outside of its natural range with varying degrees of success. It had introduced into England before 1629. Parkinson’s ‘Paradisi in Sole Paradisus Terrestris’ gives the first account of cultivated persimmon trees (Lotus virginiana). One specimen given to George III by the Duke of Argyle (ca. 1790) and grown in the old Kew Arboretum was of considerable dimensions in 1895. At that time it was “apparently as contented [there] as in its native habitat” (For. and Gard., 1895). Diospyros virginiana has also been grown successfully in British Guiana (Hiern, 1873); it fruits regularly in the warmer European climates, although it is irregular in more severe climes where it has been reported to withstand tempera- tures as low as —25° C. (Naudin, 1880; Trabut, 1926): FACTORS AFFECTING LOCAL DISTRIBUTION Although the persimmon is frequently cited in the literature as a pioneer species —a tree that is among the first to grow in almost any situation—a survey of the flora growing spontaneously within the present city limits of St. Louis failed to show any persimmon trees growing in those abandoned places where a “pioneer” would be expected. That persimmons did grow in the area now incorporated 25 St. Louis proper is borne out by a few, but reliable, references. One of the most 1953] : SKALLERUP—DIOSPYROS VIRGINIANA L. 215 z OLD FLORISSANT ROAD e. CLAYTON —® ROAD © / ST 1001$ [| ГД $ 1.5; 56 P4 O 0.4.22 | 3 —_—_— 0 N 61-67 e/ | TELEGRAPH} | {ROAD || | Fig. 2. Map showing occurrence of D. virginiana in the St. Louis area. Black dots represent sites where it was found closest to the city; circles represent apparently favorable sites where it did not occur. interesting is the narrative by Elizabeth Kennerly Russell, “Persimmon Hill’. Although written in the style of a saga, its title refers to the actual Kennerly mansion, Có£e Plaquemine, built in 1832 on a hilly tract of land "five miles north- West of town... where Taylor and Kennerly Avenues cross today," and named for the persimmon trees commanding the site (Russell, 1948). Prior to this time the land outlying St. Louis that was not upland forest had been kept in prairie by the Indians who annually burned the vegetation in order to flush out game. Ву 1836, the prairies had practically disappeared, due to the cessation of the autumnal fires and were succeeded by young forests (Beilmann and Brenner, 1951). Nicholas Riehl, collecting in 1838, gathered persimmon specimens from the "fórets" of St. Louis; while a visiting farmer, Patrick Shirreff, tasted his first per- simmon upon travelling Old St. Charles Road, only to mention its disagreeable taste in a letter written home, dated 1835 (Stevens, 1906). Coincidental in time and place with references to the flora are necessarily those to the birds and animals of the region about which William Kennerly reminisces “. . . we learned the ways [Vor. 40 216 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE II Road Sites where D. virginiana was found | Sites where D. virginiana was not 70 (represented by black dots, fig. 2) found pest 5 circles, fig. 2) E US 6I (1) Clone, on top of , lope at edge | Mixed oak association II. Telegraph Road (US. 66 и $. 50 V. Clayton Road VI. Alternate U.S. 40 . Old Florissant . Missouri 99 . Missouri 109 (not on m *1. Acer Neg 2. Celtis oc 4. Quer ap) eee alis 3. Pla valle кь Cus sp. of old field wi (2) Individuals in grove with 2, 4, 8 Scattered, on top of ravine, s.w. side in oods mixed oak w (1 Jr бе 1). (2) Scat- tered, in grove with Pinus echinala, 1, 2, and 6 yim on slope = old field at edge of grove of Ulm Scattered, in stand of 1, 2, 3, 4, 8, and Cornus sp. E bc slope of old field, with 5, ropogon virginicus, Aster s e» КЎ in mixed oak asso- cia attered, along ditch with 4, 3 us glabra, Populus de ltoldes. PRA pecimen of D. = var. pubes- cens found her "wes side of road, with 2, 4, ; Rbus Heb: Prunus sp Scattered, glade with Bumelia lanuginosa, TUE canadensis Flood plain of Mississippi River Mixed oak association Along River des Peres Brier thicket in recently cleared field (1) de pu of Missouri River at St. (2) dee p association Mixed oak association Dense grove of Ulmus at edge of cultivated field uds of animal life in the woods and meadows . . . hoofprints of the deer as well as the bobcat’s heavier tread . . .” (Russell, 1948). Dos al albid Symphoricarbor < а Ulm we could read the small dainty Wild turkey, deer and abounding game were also written of by Flagg in 1836 (Thwaites, 1896—1901; see also Beilmann and Brenner, 1951, for a more complete account). However, with the expansion of the city, persimmon became scarce within its limits. In 1911 the Engelmann Botanical Club listed D. virginiana occurring in St. Louis proper only in such floral sanctuaries as Missouri Botanical Garden and in Tower Grove and Forest parks (Engelmann Bot. Club, 1911). j Unfortunately the origin of these ney like plants in many parks and botanical gardens, is speculative. Amon ut trees in the nursery of Tower Grove Park in 1871, fifty were ии being persimmons (Shaw, 1871). Henry arida ‚ in addition to purchasing the Riehl collection herbarium specimens, p bought the first trees planted in his Garden from the Riehl Nursery (Spaulding, 1909). 1953] SKALLERUP—DIOSPYROS VIRGINIANA L. 217 Ironically, a year later persimmon was recommended as a native tree suitable for growing in St. Louis (Jensen, 1912). To determine the present occurrence of persimmon in the St. Louis area, a reconnaissance was made along the main highways radiating from the city. Although every side road could not be followed, and doubtless, in the forced pace of highway traffic, some trees were overlooked, a fairly complete idea of its local behavior was obtained. Figure 2 and Table II summarize the data obtained. It can be seen that persimmon does not occur within the city limits, and is primarily an upland tree which prefers an open drained habitat. Diospyros virginiana was usually found growing in clones or scattered in open situations where there is little competition; or else it was found scattered with other second growth trees or in young mixed-oak forests (pls. 25 and 26). In the former instance, the trees were young and were initiating growth, while in the latter instances they were compara- tively old or dying and represented the last of their number. The present local ab- sence of persimmon within the metropolitan area can not be explained on the basis of unfavorable habitat brought about by smoke or industrial gases, for trees are suc- cessfully grown as ornamentals in the city parks and at the Missouri Botanical Garden. Rather, the lack of dispersal agents for its seeds appears to be the under- lying cause. Persimmon is poorly equipped for dispersal by means other than animals, or, indirectly, by man. The fruit, а true berry, ranges from 94 to 172 inches in diameter, and bears proportionately large seeds, four to eight in number. Once separated from the pulp, the seeds are not light enough to be scattered by wind or to float upon water. In the city they can still be seen under cultivated trees, un- touched by squirrels, rabbits, or rats, as late as June. Tables Ш and IV list the animals that eat persimmon fruits in the wild. Although these data do not show diet preferences, the fact that the animals listed do eat persimmon fruit indicates that they act in a large degree as dispersal agents for its seeds, since it is unlikely that the seeds are dispersed by any other means. In addition to the animals listed, the pine mouse will use the seeds as food (Martin, 1951), while the meadow mouse is known to include persimmon and Bumelia seeds among its caches, and thus figure in its dispersal (L. С. Brenner, personal communication, 1953). While certain birds eat the flesh or pulp of the fruit, it is doubtful whether any listed in Table III B (except the turkey now rare in Missouri) is able to swallow or void the large seeds. This is in contrast to sassafras (Sassafras albidum (Nutt.) Nees); а "pioneer" species which does grow spontaneously within the city, whose fruit and seeds are eaten by birds and thus is scattered more widely than persimmon. [Vor. 40 218 ANNALS OF THE MISSOURI BOTANICAL GARDEN ABLE III LIST OF ANIMALS AND BIRDS EATING PERSIMMON FRUIT* | Per cent of Diet 1 Species | Southeast | Northeast А. ANIMALS Red Fox 10—25 0.5—2 Стау Еох 2—5 0.5—2 White-tailed Deer 0.5-25 Not recorded Opossum 2-5 Not recorded 5—10 5—10 Ring-tailed Cat (Texas only) 5—10 n, Stripe d Skunk 0.5—2 Not recorded В. Binps Wild Turkey Not recorded 0.5-2 Catbird Not recorded ed — Not recorded Yellow-bellied Sap-sucker Not recorded Food item of undetermined extent Myrtle Warbler 0.5-2 edar Waxwing 2-5 ite d Mockingbird . Not recorded Food item P iia СЛ extent * Based upon stomach and crop analyses, droppings, and field observations (after Martin, 1951). TA OCCURRENCE OF PERSIMMON IN MISSOURI WILDLIFE* Species Samples examined Occurrence of persimmon Deer 348 stomachs 40 Coyote 680 stomachs 51 Red Fox 731 stomachs 31 Gray Fox 251 stomachs 56 Quail 5,472 crops 33 Quail 1,358 droppings (roost) 1 * Prepared from letter, L. J. Korschgen, Missouri Conservation Commission, 1953. HABITAT TYPES As mentioned above, D. virginiana was found either initiating growth и abandoned places or as а minor element in mixed-oak associations. To interpret its behavior in these habitat types, data will be presented from sites where more com- plete studies were made. Old Fields.—Many authors list D. virginiana as occurring in clones along roadsides, in old fields, and other waste places (Sargent, 1894; Eddy, 1927; уха Dersal, 1938; Steyermark, 1940). Personal observations were made along roadsi and in old fields at the Missouri Botanical Garden Arboretum, Gray Summit, where 1953] : SKALLERUP—DIOSPYROS VIRGINIANA L. 219 information concerning the history of the land is fairly complete. Here, three patterns of "field invasion" were found to be present. These may be characterized as: (1) Clonal: trees of varying ages, all of the same sex. (2) Clonal: trees of almost the same age, all of the same sex. (3) Scattered: trees of varying ages, both sexes present. (1). A group of the first type was found in a sloping field, abandoned since 1924, which, except for the eradication of elm, had been allowed to grow wild (A. P. Beilmann, personal communication, 1953). An apparently injured, double trunked tree, near a shallow gully, was the oldest element present (c. 20 years). Growing in association with it were Andropogon virginicus, Allium sp., Panicum, sp. Aster sp., Solidago sp., young Juniperus virginiana, Cornus racemosa, and Parthenocissus sp. In somewhat elongated concentric arcs about the old tree were other persimmon trees of different ages, the younger ones being on the periphery, but more numerous down the slope (pl. 25). Trees old enough to bear fruit showed the single peduncles characteristic of pistillate trees. Upon digging near the bases of several older trees, long horizontal spreading roots? connecting the older trees were found at a depth of a foot (pl. 25). Several roots were followed for their entire lengths and were found to branch repeatedly. These gave rise to several small trees, one of which is shown in pl. 25. The diameter of the root remained almost constant (34 inch) except near the tips, where normal growth occurred. "There were no evidences of decay or separation of root branches; nor were the branches themselves chance root grafts. From observations of other sites similar to this one (Table II) it was noted that the advance of the younger trees is usually more rapid down the slope, probably due to the greater development of the root system in this direction, a consequence of gravitational influence or greater soil moisture. Although the aspect of this site was largely clonal, a few of the smaller trees may have been seedlings, since seeds collected here were found to be viable. (2). The second group was observed along a roadside near an abandoned field of similar history to the one above, except for possible disturbances due to road repairs. Other species growing at this site included: Rhus glabra, Sympboricar pus orbiculatus, Rubus sp., Andropogon virginicus, and Rudbeckia sp., as well as a few Very young Juniperus virginiana trees. The largest persimmon trees examined here were all essentially the same age (about 15 years). They exhibited the uniform growth characteristics and the same sex of trees vegetatively propagated from a common stock, yet no "parent" tree as in the above situation was present ( pl. 26). These trees were also found to be connected by stolons, while some individuals had their own groups of younger trees about them. Seeds found at this location were also viable. It is conceivable that this situation was brought about as a result of the — There are references to persimmons as "stoloniferous" (Sargent, 1894; c.f. also, Holm, 1909). However, a complete description of this condition was not found in the literature. The roots described here did not appear to differ in gross structure from the normal roots. [Vor. 40 220 ANNALS OF THE MISSOURI BOTANICAL GARDEN roots of an older persimmon tree having been accidentally cut into pieces and dis- tributed, so that groups similar to those of the first type were concentrated in this small area. That this is possible is demonstrated by a common horticultural prac- tice in which cuttings of persimmon are made for propagative purposes and vigorous sprouts are obtained from them. Further, decapitated trees have been reported to send up root suckers within two months after being injured (Fletcher, 1915). (3). On the top of the slope of the same field, but away from the road, a third pattern was found. Неге trees of varying ages and forms were scattered in groups. Most of the trees were from 15 to 25 years old, although some seedlings were also found. The trees were both staminate and pistillate, and occurred in sporadic groups over the field. Most groups were composed of two or more trees, although some lone trees were noted. Ecologically, this site is part of the “highway” system used by deer and other animals at the Arboretum (Beilmann, personal communi- cation, 1953). The scattered, but grouped trees closely reflect the way seeds are dropped by animals. Yet it is difficult to account for their non-clonal habit. The most likely explanation is that none of the trees were ever injured to the extent that they began to sucker. No plowing had been done in this field for over 25 years and the trees were not planted out. Injury by cattle grazing (besides trampling) is speculative; there are reports that cattle will not browse persimmon leaves or twigs (presumably because of calcium oxalate crystals and tannins present—Deam, 1932; Van Dersal, 1938); however, they are known to eat the leaves when suitable forage is not available (Brenner, personal communication, 1953), and many wild animals are reported to eat the buds and leaves (Missouri Conservation Bulletin, 1940). Forest associations.—From observations at the Missouri Botanical Garden Arboretum and in the St. Louis area, persimmon was never found upon a river flood plain (Table II), nor were mature trees ever found in a pure stand. Persimmon is frequently mentioned in the literature as existing in pure stands and as reaching its greatest proportions in the basin of the Mississippi River and along the Wabash River Valley. Ridgway (1882, 1894) and Sargent (1894) are probably the un- cited sources for these references. However, a more recent survey of the hard- woods growing in Louisiana listed persimmon as only a minor species in all the habitats in which it was found (Lentz, 1929; Hepting, 1936), while Miller and Tehon (1929) report that no persimmon trees as described by Ridgway are now known in the Wabash Valley. Evidently, persimmon was more of a dominant tree (as were others) in the primeval forest than it is in the second growth timber of today. That D. virginiana is only a minor element in present-day associations is veri- fied by other authors. In the Ozark and adjacent plains region, Steyermark (1940) found D. virginiana occurring as a pioneer element on dolomitic glades, 25 2 temporary dominant on bald knobs and bare limestone areas, and, frequently, 1953] SKALLERUP—DIOSPYROS VIRGINIANA L. 221 initiating oak-hickory associations on the acid substrata of the prairie. However, he did not list it as a member in the building of a flood plain climax, in associations where burning-over had recently taken place, or in drainage regions. Palmer (1921) did not mention D. virginiana as a conspicuous element of the flora of the Ozarks. Although Lentz lists the persimmon as growing in both the well- and poorly-drained bottomlands of northeastern and southern Louisiana, it was not mentioned as occurring in the Louisiana swamps or in lands subject to overflow. In the Carolinas, Eddy (1927) found it growing at the edge of alder swamps and on the coastal plain, but again, only as a minor or subdominant species. his evidence indicates the wide range of ecological situations in which D. virginiana is able to initiate growth; yet, because of its slow rate of growth, it is shaded out by faster-growing trees. Brenner (1942, 1952) made comprehensive observations of the mixed oak associations in his environmental and quadrat studies at the Gray Summit Arboretum. An example of the short duration of the per- simmon in an open woodland can be found from data reported by Brenner from а quadrat staked out in an oak coppice in 1937. At that time, numerous persimmon and other heliophilous trees were keeping pace in growth with the white oaks, the dominant tree. Since then, the white oaks made considerable growth, with the result that, 15 years later, one-fourth of the persimmons were dead and the remaining ones were in poor condition. SUMMARY 1. Although formerly members of the genus Diospyros ranged far to the north, the native persimmon now appears to be limited in its spread to the north by an annual temperature of 25° F., and to the west by rainfall of 30 inches annually. Within these limits, however, it appears uncommon in the area of the Wisconsin glaciation. 2. Supplementary chromosome counts from Missouri (three counties) and Arkansas (one county) are in agreement with previously recorded determinations. 3. Local absence of D. virginiana is attributed to eradication of the trees that were present in what is now the city. Dispersal of seeds is largely through the activities of animals, and for this reason persimmon is not known as a pioneer species within urban limits. . Three types of “field invasion” by persimmon in abandoned places are described. The ability of the roots to sucker upon injury accounts for the dense clones often found in these situations, although non-clonal situations also exist. 5. Although persimmon once was known to occur in pure, dense stands, more recent reports indicate that this is now not the case. Throughout its range Persimmon is reported as a minor species in older associations. ACKNOWLEDGMENTS I wish to express my gratitude to Edgar Anderson, Missouri Botanical Garden, Whose suggestions and encouragement made this study possible. Thanks are given to those already cited by name in the text for data and information received and [Vor. 40 222 ANNALS OF THE MISSOURI BOTANICAL GARDEN used in this paper, and also to the following: L. E. Arnold, University of Florida; J. T. Baldwin, College of William and Mary; G. N. Jones, University of Illinois; B. Long, Academy of Natural Sciences of Philadelphia; G. B. Ownbey, University of Minnesota; B. C. Tharp, University of Texas; R. F. Thorne, State University of Iowa. I am indebted also to C. J. Felix, Washington University, for a valuable contribution. BIBLIOGRAPHY Anon. ees The persimmon, Gard. and For. ——,‚ (1909). Missouri has the une sedation ач in the world. St. Louis Globe-Democtat. Jan. Е ———, (1915). An unusual persimmon tree. Jour. Hered. 6:525, fig. I7. Aikman, ү M., and I. І. Boyd ge ic эзше батады studies of plants for soil conservation purposes in southern Iowa. Iowa Aca . 48:117-121. Bailey, L. H. Quem Sketch "p L^ ga e of our native fruits. pp. 433-441. 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Agr., Bur. — . Bull. 141: Gürke, M. f: Ebenacea In Engler was Prantl, Die Natiirliche НЕТ 41:153-165. ague, S. M. (1911). А morphological s study zi prion virginiana 34-44. Harrington, F. O. (1 (20). The wa. lowa State Hort Pup ре Hedrick, U. P. (1919). ан поїез оп S editis кабы N. У. Agr. Exp. бе, эы 2:2 Непгу, А. нее ). Knospenbilder. Nov. Act. Acad. Caes. Leop. get Nat; Cur. 12: peo Hepting, G ex — following fire in young Mississippi delta hardwoods. U.S. Dept. Agr. h. Bull. 494:1— or Y. P. (1873). x “monograph of the Ebenaceae. Trans. Cambridge Phil. Soc, 12:27— Hodgson, R. W. и, Te situation, "s © "ig tion, and parthenocarpy in the ад per si s Proc. t SER 372 Holm, Theodore ا‎ a eed plants ts North America, No. 31: Diospyros virginiana L. ate Mus. Bull. Hume, Н. es (1914). Piae persimmons. Jour. Hered. Jensen, Г. Р. (1912). Native мы pn ralized plants ot ved ME for ornamental planting. о. State Bd. пот. рес. 519 وا ,410:52-57 Kincer, J. B. (1922). Precipitation and и Atlas of American Agriculture, Part II, С чаар наа Uus t. Agr. Adv. Sheets eon oom (1928). Atlas of (pet Agriculture. Clim e, temperature, sunshine and wind. LaMotte, R. B ( feed Catalogue of the Cenozoic i Rn 'of North America. Geol. Soc. Heg "n Mem. 51:146-14 Lentz, G. H. (1929). Summary of first year's hardwood investigations in Louisiana. Jour. For. 6 27:486—494, Lowe, Е. М. (1 Aug Plants of ret Miss. State Geol. Surv. Bull. 17:222. т re a and H. P. 6 ub (1941). Climatic ry de ions of fruit d pe crops. In Climate onde U. 8 god Agr. pp. 409—4 arr BE Hoe ш ш to Ámerica. Ha Oud 377. Marshall, H H. (1785). Arbustrum pg ericanum. p. 40. J. C а b os delphia. Martin Hill, et al Ei 51). American wildlife iid таң, рр. 219—227, 268—270, 358—369. McGraw- ‘New — Е т tus we Agricultural exploration in the fruit and nut orchards of China. U.S. Dept. Bur. Plant us e 204:10-1 М Miller, "Ro B., and L n (1929). "The native naturalized trees of Illinois. Ш. Nat. Hist. Surv. Bull. is des 267. Missouri Conservation Commission (1940). Conservation Bull. No. : а пена С. (188 0). Quelques remarques au su и de н ' Diospyros) ooh à Pair e dans des fene de l'Europe. Nouv. Arch. Mus. Hist. Nat. Pa ris П, 3:2 Озо, ^ d al (1947). The National EAT E sath 4. р. 1435. J. B. enge: (^ Phila- Palmer, | E. 3. (1921). The forest flora of the Ozark region. Jour. Arn. Arb. 2:216— xe. and J. A. ge srg (1935). An annotated catalogue of the ера es of Mis- Ann. Mo. Bot. Gard. 22:365—759 Parkinson, J. (1629). Paradisi i in Sole Paradisus бле A рр. 569—570. London гоми, Е. (1892). Histologie ни des ées. а vr d Lyon 6: 1-150. ellett, F. С. (1920). American honey plants. Hamilton Pharmacopoeia of the United States of America КТ. 5th fmi rev. ed., p. 58. Lippincott Co., Philadelphia. —— 5 (1882). 6th decennial rev. ed. Wood Co Papas F. P. (1883). Resources of the southern fields je Lian pp. 386—388. Charleston, S Г du P Нес ы Page (1763). The history of Louisiana or of the western parts of Virginia an P. 15. London. on Im Р. as 16). Flor. ra Am e septentrionalis 1:2 Ph ipis nesque, C. E. elphia. Ridgway, R. e а Media ка Fora 133-13 К ге г Wabash Valley and White River valleys in Illinois and In ive . Proc. U. S. Nat. feas enemy. (1894). Additional pnus on the native trees of d s lovi Wabash Valley. Ibid. 17:409— [Vor. 40, 1353] 224 ANNALS OF THE MISSOURI BOTANICAL GARDEN Russell, E. K. (1948). Persimmon Hill. 9—70. Univ. Okla. Sargent, C. S. (1894). The Silva of Marche merica 6:1-10. Cambridge, Mais: ————, eun Notes on North American trees. УШ. Jour. Arn. Arb. 2:1 — Е. (1913). Les cultures sur le littoral v la мифа Ped йн, Algerie. zs B. Balt et fils, (HS (1932) RB doy Me of Ohio vascular plants. Ohio Biol. Surv. Bull. 23:173. Shaw, H. Sod 1). Second Annual Report of the Commissioners of Tower Grove Park. p. 10. St. Loui ае Harry R. (1953). Our "pec persim . Gard. Bull. J. K үз Мо. mall, 01). сдаю" and trees of the S sik Dura: re “Bull, T в р P 356. 21). Old trails and n а. Jour. №. Y. Bot —— (19 t. Gar Spaulding, F. e т А е айка чәй sketch of botany at St. Louis, Mo. Т; рор Sci. Month. Stebbins, с. L m d Variation and evolution in plants. pp. 350—381. Columbia Univ. Press че F., ж (194 g> — 2 tot Hed — of Seed and Plant Materials in the Trade, pp. “91, » Mass Stev: в : 1906). f аны et St. 3 с ТЫШ . Louis. Steyerms р » A. (1940). Studies of the лода = einen I. Field Mus. Nat. Hist., Bot. ‚ Si 1940). "Plants new or apiid unrepcrted from Missouri. рано 42:101. 41 he persimmon tree. Мо. Bot. Gard. Bull. 37:192-19 Thwaites, к. xm (1896-1901). jena Relations ‘end “Alli ed Documents i: 137; 65:73; 66:229. Burrows Bros., Cleveland — — ——, (1904-1907). Early herd Shap ды 321, sits A. H. Clark Co., E Trabut, L (1926). Des Diospyros co v. Bot. Appl. Agr. Col. 6:541—547. Tr and O. M. Hadley (1896). rev pes Presse Indiana Ai Exp. Sta. Bull. тарж Van Dersal, В. (1938). dote woody plants of the United States, their erosion control and wild een values. U.S. D ч TM ied er 117-118 Watts, R. L. (189 im gr. . Sta. Bull. 1 White, O. E. (1926 sc ; Geosraphieal diriciburica © e cold resisting characte of certain herbaceous perennial and woody plant groups. Brooklyn Bot. Gard. —10. 194 Temperature. reaction, mutation, and на оцон in plant groups. Eighth A m. Sci. Cong. 3:287-2 Proc Yasui, K. (1915). Studies of Diospyros hoki Г. Bot. Gaz. 60:362-375. EXPLANATION OF PLATE PLATE 25 Diospyros virginiana L. Fig. 1. Fig. 2. Portion of a “stoloniferous” root from the above situation showing its branch- ing nature and one young tree attached to it. (Length of root approximately 6 feet-) Clonal type 1: Trees are of varying ages, and same sex. ANN. Mo. Bor. Ganp., Vor. 40, 1953 PLATE 25 SKALLERUP—DIOSPY ROS VIRGINIANA L. PLATE 26 ANN. Mo. Bor. Garp., Vor. 40, 1953 mE sag SKALLERUP—DIOSPYROS VIRGINIANA L. [Vor. 40, 1953] SKALLERUP—DIOSPYROS VIRGINIANA L. 225 EXPLANATION OF PLATE PLATE 26 Diospyros virginiana L. Fig. 1. Clonal type 2: Trees are essentially the same age, and are of the same sex. Fig. 2. Tree group characteristic of the third type. Fig. 3. Persimmon in mixed oak association. WILDWOOD A STUDY IN HISTORICAL ECOLOGY ALFRED G. ETTER* PROLOGUE This is the story of a gathering of towering trees, old spreading oaks and sky- penetrating hickories that shelter the past in their shade. It is one of the few old woods that remain on the rich uplands that lie northwest of St. Louis between the Missouri and Mississippi rivers. Most of the scattered timber that early settlers found on the headwaters of the creeks in this area was cleared to make fields, or cut for cabins, firewood, fence-posts, lumber, cooperage, or one of the many other purposes for which wood was used during that period of American expan- sion which so depended on its omnipresence. A few wood-lots, through the smile of fortune, were preserved on tracts of land belonging to large estates. Wildwood was one of these estates, and it took its name from one of these old woods. HisTory In the earliest days of the frontier when the land took men and hid them in the shadows of its trees, or lost them on its seas of grass, Wildwood’s acres stood ignorant of everything but Indians. Then French forts began springing up along the near-by Mississippi. The French town of St. Louis was founded in 1764. Shortly thereafter Spain assumed control of this Upper Louisiana region. Then near the end of the century the Spanish regime began to yield to the pressure of the American movement west. Grants of land were made by the Spanish to many early frontiersmen and explorers, among them Daniel Boone. Settlements were also recipients. The villa ge of Florissant, a few miles north of Wildwood, received a large gift of land for its common fields and wood-lots, about 1785. It was around the time of Christmas in 1799 that the Spanish Lt. Governor, Don Carlos Dehault Delassus, granted to John Wendel Engel a tract of 800 arpens (680 acres) lying on the river Maline. Wildwood (to be) was in the northwest corner of this grant, and so first became a parcel in the domain of Man. By 1804 John Engel was apparently stirred by the ever-ready lust for money, for in Janu- ary he sold 600 arpens of his land to James Rankin for 500 dollars. The other 200 arpens, including Wildwood, he reserved for himself for cultivation. The Spanish regime ended officially in March 1804, and the great territory of Louisi- ana became American. Whether or not this influenced Engel in some manner 1s hot certain, but in August of that year he completely gave up his ideas of farm- ing by the river Maline and sold to Simon Wood the remaining land, including that destined to become Wildwood. During his brief period of residence Engel had built a house and cleared ten arpens of land and put it under fence. The 200 ые B. Clopton Experimental Farm, Clarksville, Мо. Washington University School of edicine, (227) [Vor. 40 228 ANNALS OF THE MISSOURI BOTANICAL GARDEN arpens, house, and improvements, together with a spring, went for the price of 200 dollars. After the transfer of the capital of the Territory of Louisiana to the City of St. Louis, the old fur post became an active and growing city and the headquarters for expeditions and vagrant dignitaries of every description. Simon Wood held his property for sixteen years, but during this time entered into some contract with a gentleman named Josiah McClenahan. Wood could not fulfill the terms, and was obliged in the spring of 1820 to sell his farm to McClenahan for 500 dollars. John Mullanphy had entered into a covenant previously for the purchase of the same land from McClenahan for 750 dollars, as soon as the latter should obtain a clear title to it. This shrewd deal was accomplished just before Missouri received its statehood in 1821. In 1830 Mullanphy provided that the 200 arpens on Maline Creek should be the property of his young grandson Owen Chambers. It was agreed that should Owen die before the age of twenty-one the land was to revert to Owen's brothers and sisters (of whom there were to be twelve)! Owen did die before he was 21, and in 1850 it was necessary to divide the Maline property. Fortunately Mullan- phy had meanwhile procured the 600 arpens of land which were a part of the original grant, so that when the division became necessary there were 800 arpens to be divided among the eight children interested. To make a complicated story simple, of the resulting eight lots, lots Nos. 7 and 8 contained the 200 arpens of Simon Wood's farm. Lot 7 went to Ellen Chambers LaMotte, wife of J. H. LaMotte, ultimate owner of Wildwood. Lot 8 went to John Chambers, who in 1851 sold the property to LaMotte, and so the original plot of 200 arpens was once more consolidated. The price paid for 110 acres was $3,292. At the time of the purchase LaMotte was a major in the United States Army and only five years before had been actively engaged in the battles of the Mexican war. He continued his service until 1856, striving assiduously (as he expressed it in letters written at the time and preserved in the library of the Missouri His- torical Society) to make the necessary preparations for a happy life on his Maline land. These Preparations included the careful sale of many of his investments 10 Property in St. Louis and elsewhere. During this time his farm was run by various tenants, as it had been in the years before it came under his ownership. When he resigned from the army he Proceeded immediately with his plans for the trans- formation of these 220 acres into a home and estate. Work was begun on the construction of his seventeen-room home. Stables, quarters for slaves, and mis- cellaneous buildings slowly began to rise from mere vision to the reality of Wild- wood, country estate of J. H. LaMotte. Then came the interruption of the Civil War and its attendant complications. Actually it was not until 1867 that the LaMotte family was established in the ael home. At that time LaMotte, visiting in Georgetown, received a letter from his son, Frank, to the effect that “We are well and comfortably fixed at Wildw 1953] ETTER—WILDWOOD, AN ECOLOGICAL STUDY 229 plenty to eat, drink and smoke. Carpets down, potatoes all in, fences up. Harry and mother ride out frequently and enjoy the country very much . . . Uncle is furnishing our apartments in princely style—marble mantels, paper etc." It was in keeping with the wishes of Major LaMotte that the woods around the house were never cut, though horses, cows and sheep grazed at various times through the undergrowth. Meadows of timothy and fields of corn occupied the flat valleys north of the river Maline. On fertile slopes tobacco was planted and nearer the house were peach orchards, rows of potatoes, and sun-filled patches of watermelons for which the Major became famous. The raising of horses became his particular pastime, and their grazing, combined with that of the sheep and cows, soon made a spacious park from the woodland about the home. The Victorian refinement of the secluded Wildwood home held sway in lux- urious comfort, peace and plenty while Major LaMotte lived. After his death in 1892, his wife and few remaining members of his family continued to live in the home, but his many activities were no longer attended. A tenant farmer ran a few cows in the extensive pastures, but apparently not enough to keep many young trees from sprouting. In 1911 Mrs. LaMotte died and the property went to her daughter, who passed away within a month. Her husband lived alone in the man- sion for a few years, leading an almost hermit-like existence, neglecting the home and the fields. At his death, the home stood alone in the woods, forgotten except by vagrants who found the old place a haven on winter nights. There was no caretaker and no interested heir. In the meantime a new tenant farmer had moved on the property and began building up his dairy herd. According to his recollection, about 1920 he found it necessary to enlarge the available pasturage. Portions of the woods that had grown scrubby were cleared, and the really dense areas in the valley that had never been pastured at all were also cleaned out. Fallen trees and logs were burned, and the cows turned in to finish the work. Mr. S. still resides on the acreage and pastures his cows in Wildwood. Until 1929 the LaMotte mansion at Wildwood remained abandoned. At that time it was sold to the present owner who removed seven rooms and restored the test. His purchase included only that land in the immediate vicinity of the house. The remainder (about 200 acres) was sold at public auction in 1954 for $60,000. The purchaser intended to build a private home on the grounds, but his plans did not mature, and during 1946 а “Рог Sale" sign went up, offering 180 acres for $180,000! The price of this land had thus risen from approximately $1 an acre in 1804, to $3.00 and then $4.50 in 1820, $30.00 in 1851, $300.00 in 1934, and $1000.00 in 1946. Once it was free! : The pressure of suburban development has finally risen to the point where Wildwood can scarcely hope to exist longer as a unit, much less as a farm, within City limits. In this respect, this study is essentially an obituary. [Vor. 40, 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN 4 Cage ee У [OK 793 КОЛ? e z 9 Я Cansar. бм і Рів. Prairie and timber areas northwest of St. Louis as recorded by surveyors іп 1817-1818. Timbered areas dotted; prairie areas unmarked; no survey data were at hand for the extreme northeast corner of map or for St. Louis proper. Original Spanish grant outlined in heavy black; id blac marked i i EARLY DESCRIPTION Figure 1 shows the original Spanish grant in heavy outline, and Wildwood (the wood-lot and residence area) in solid black. Maline Creek drains eastward through river-bottom woods into the Mississippi River, while to the west Cold- water Creek flows north into the Missouri. Wildwood is seen to lie between branches of Maline Creek, and consists of sloping land above cultivated bottoms to the east and south. In 1817, shortly before the founding of the State of Missouri, extensive suf- veys were made of township and range lines in this area and details of topography and vegetation were recorded by the surveyors. From this material, preserved in the St. Louis County Engineer's office, it was possible to prepare fig. 1, whic shows the distribution of forested and prairie land at that time. Hickories and Blackjack Oaks seemed to be the predominant types of trees. There is a strong tendency for the timber to be confined largely to the dissected headwaters of Maline Creek, and Gingras Creek and the River Des Peres to the south and yer е surveyor, as he progressed along the range line of Section 12 a few chains east of Wildwood made the following remark: “Тапа rolling, soil rich and good for farming, thickly covered with oak and hickory". (Section 12 is indicated on fig. 1.) Significantly, however, he stated that halfway along this section line he built a mound (for a marker) in a “prairie,” presumably a grass area surround by woods. Along the north boundary of the section the notation, "Prairie, soil, scattering blackjacks and hickories,” was made. On the west border 1953] ETTER—WILDWOOD, AN ECOLOGICAL STUDY 231 country was “Gently rolling, тер: timber, blackjacks and oaks”. It is thus vividly demonstrated that Wildwood was at the edge of the prairie. This fact plays a significant part in the abe of Wildwood's vegetation. On fig. 1 there also appears a soil boundary line. This was taken from the soil survey of the county made in 1923 (Krusekopf). It represents the eastern- most limits (in the county) of the Tama silt loam, a rather well-developed prairie soil. To the east is what was then called the Memphis, a forest soil. Apparently those grasslands described by the old surveyors as being east and south of this soil boundary line had not been established for a long enough period to have their inevitable effect on the soil. The grass was invading land which inherently belonged to timber. Fortunately, one of the early Government surveyors that worked in the region, a Mr. R. W. Wells, was interested in this prairie-forest problem. In his article (Wells, 1819) “Оп the origin of prairies," he made the following statement: ...the writer has seen, in the country between the Missouri and Mississippi, [pre азы the area with which si are concerned] wur unusually dry sea erg more than one ry undred acres of woodland together converted [by fire] into prairie. An again , where p Brass has been prevented from burning by accidental causes, or the prairie has been de- u rge he e, it will assume, in a f those places. Now the s вое ountry for several miles is covered with a growth of trees of four or five riches Жайды, near the towns where the burning first ceased, and Eh dea diminishing in size as you recede, until you at length gain the open prairies. Nearly twenty years later another writer made observations which provide an interesting sequel to Mr. Wells’ description. In his diary of a trip through the outskirts of St. Louis in 1836 Edmund Flagg has written: The face of the c выт is neither uniform nor broken, but undulates imperceptibly away, clothed in dense forests of blackjack oak, jd wi with thickets of the wild plum, the crabapple and the оно үт years ago » MM plain was a treeless, shrubless waste without a solitary farmhouse to break the notony. But the annual fires were stopped, a young forest sprang into а delightful villas and country seats are now gleaming from the dark i in all directions. It is difficult now for us to visualize our predecessors of 100 years ago living in a country of scrub oak, but such seems to have been the case. This change Was not a uniform one, however. As Wells pointed out, it often began in the Vicinity of settlements. It undoubtedly moved faster on some terraine than on Others. "That there was still considerable prairie left northwest of St. Louis at the time Flagg made his trip is shown by a letter which Evert A. Duyckinck wrote in 1837 in which he described a trip he made from St. Louis to Florissant: Bive you an early account of our yesterdays visit to the humorous ps ре: tow e аа situated in the midst of the flowers of the Prairie We ia 6 ыа six o’clock. . . . We stopped оп the rough edge of the Prairie which lies pagum the City where the scrub oak and marshy weeds ill set ой by the thick fog gave us im- [Vor. 40, 232 ANNALS OF THE MISSOURI BOTANICAL GARDEN prenio n of the beautiful Ama und ачк that lay berend us in nthe fields to give so pretty name to little “Floris ssant. Here we breakfasted. ant 9 way again in the field. The mist gradually ro ү сы 2 TÉ I had m y бы rst gt Prairie аве азау & broken into innumerable landscapes with its "г fields « e hel and trees. A smooth level road lays through the Prairie which is rarely enclosed by fences and as rked it was a ride through a garden or the private lane on a E Bridge ici. MEE 1944) WILDWOOD ‘TODAY Wildwood was once one of those unfenced fields of trees. Today it still persists, though it is fenced and surrounded by suburbs instead of by prairie. It is mature now, even over-mature as the large number of dead and fallen trees on the diagramatic representation of the woods in fig. 2 show. Nearly one fifth of the trees had died in 1946. At the present writing the number has greatly in- creased. It is predominantly an oak-hickory woods. A complete tree-by-tree survey of the entire 41.5 acres in 1946 showed that Pignut Hickory, Black and Shingle oaks made up 54 per cent of the total. Twenty-one other species made up the rest of the population (Table I1). A glance at fig. 2 suggests the open quality of the woods and shows the cow trails that have resulted from years of TABLE I ANALYSIS OF WILDWOOD BY TREE SPECIES (INCLUDING DEAD TREES) Frequency Species Dead Alive Total % of | (96 of zones Total | in which found) Black ice ( e nigra) 4 4 .5 13 Mockernut Hickory (Carya tomentosa) 2 2 3 6 Peni” Hickory Сы ovalis) 17 82 99 12.4 100 Shagbark Hickory ( Carya ovata) 1 1 1 White Oak (Quercus alba 1 15 16 24 25 Post Oak (Quercus stellata) 3 3 4 13 Ке4 (Quercus ти 4 5 25 Black Oak (Quercus v * 101 101 12.6 81 Black Jack (Quercus marilandica) 1 30 8 76 Shingle Oak (Quer. 35 197 232 29.1 100 Slippery Elm (Ulmus та 3 36 » is 50 ries ds (Celtis etapa d 11 11 1.4 13 y (М ; 13 asai А rot albidu ar я и 5 e Sycamore (Platanus occidentalis) 2 2 E n Black Chingy (Prunus serotina) 1 2 3 4 13 lack Locust (Robinia a Pseudo-Acacia) 2 2 3 13 Ailanthus (Ailanthus altissima) 2 2 3 : Sugar Maple (Acer s rum) 3 8 11 1.4 13 es Maple (Acer saccharinum) 2 2 3 13 -Elder (Acer Negundo) 3 3 4 B die Dogwood (Cormus racemosa) 2 2 3 " Persimmon (Diospyros virginiana) 2 72 74 9.3 $ Oak, dead, species ert ified 30 30 3.8 6; Unidentified dead t 26 4l e TOTAL 119 678 797 * Included under dead oaks, below. 1953] ETTER— WILDWOOD, AN ECOLOGICAL STUDY 233 grazing. To locate particular areas and to get an idea as to which species were most widely distributed, the wood-lot was roughly divided into 16 zones by superimposing a grid. Table I shows the calculated frequency of the different species. Shingle Oak and Pignut Hickory occurred in all zones and Black Oak in all but three. Persimmon and Sassafras were the next most abundant trees though only the Sassafras was widely distributed, most of the persimmons being in a single copse. MANSION ig 1 э 3 4 Fig. 2. Diagrammatic representation of Wildwood. Solid black—Live t Dotted L к paths Radiating brancher Standing dead trees чекати сла — Пат 4 Ргопе trunks— ен Horizontal and гас lines divide Solid lines—Drainag, Wildwood into zones [Vor. 40, 234 ANNALS OF THE MISSOURI BOTANICAL GARDEN AGES AND THE PAST In order to learn in more detail what Wildwood's role may have been in the transition from prairie to forest and to attempt to find out what influence suc- cessive phases of exploitation may have had, an intensive study was made of the ages of the trees in the woods. In some cases ages were obtained by simple stump counts, but ordinarily the procedure used was as follows: With an increment borer a core was procured from the desired tree, but since the borer available was only 10 inches long, it was only in smaller trees that ages could be read direct. Ordinarily it was necessary to procure cores and stump counts from a number of different sizes of trees of a given variety. Measurements of growth in years per inch were then made and averaged to reduce the error due to skewed boring, tree shape and diameter, and habit of growth. Ап index in years per inch was thus obtained for 16 different tree species (Table II). The circumferences of 222 trees selected at random in all parts of the woods were then taken. These circum- ferences were converted to diameters and ages, according to the calculated growth index for the species. These ages were then tabulated as averages of successive twenty-year periods taken every five years. These averages were adjusted so that they represented each species in proportion to the number of individuals of that species in the total woods population in 1946. The resulting graphs are shown in figs 3 and 4. TABLE II DIAMETERS, HEIGHTS, AND GROWTH RATES OF TREE SPECIES WITHIN WILDWOOD d^. +H n Rate } © $ E growth i Ei È © $ E E yrs. per in = 2 Б = 5 987 ex [s = + y ES E و‎ pecies SEG o бмо |" A bo ak sag |$| а | 5| e388- | S.| وة‎ ed 5% |< a | £4) fea) 83 | *5|.85| Eg оз oo j| + Dem d l С] о os а о ag >j oj < 2 Bel >ле Ла эрсә ISSR | 3 | G8] <: | АЗ | 5 Е > © o > © © © о о 8 i сты. Tad шыла dli Тг |: н Z > = я $ $ R $ N © a 13,200 r 32 5 4 6 3 6 6 2 13,800 г 24 1 3 5 + 1 5 2 2 1 14,400 г 24 1 5 3 3 4 1 1 3 1 2 15,000 r 27 E 3 1 1 2 4 8 2 2 15,600 г 30 2 6 5 3 3 6 1 2 1 1 16,200 r 15 2 1 1 6 1 1 1 1 1 16,800 r 15 1 3 2 1 1 3 1 3 17,400 r 64 4 3 ^ 7 11 9 10 9 4 3 18,000 r 65 + 5 5 3 10 8 9 10 6 3 20,000 r 88 5 d 6 7 14 11 14 13 11 22,200 г 10 1 5 1 1 1 1 25,800 г 19 1 1 3 3 2 4 3 9 2,400 r 313 133 38 16 15 34 23 17 13 12 12 3,000 г 4 2 Anther 5,400 г 4 1 2 1 Anther 6,600 r 7 3 1 1 1 1 739 DISCUSSION The cytological data were recorded here for the purpose of giving background to the radiation effects mentioned. There is little need to discuss the effects of x-rays on meiotic chromosomes; this has been done at length by Lea (1947), and Rick (1943) who studied Petunia. In view of the extensive damage to chromatin in plants showing x-ray effects it is hardly plausible to assign such a character as light spots of the leaf to a particular radiation-induced chromosomal aberration. The widespread x-ray-induced leaf abnormalities may well attest to the large number of loci concerned in leaf development. It is suggested that in the leaf abnormalities considered here cells often appear undifferentiated rather than differentiated abnormally. In plant No. 40 (pl. 30, fig. 6), the mesophyll cells resemble the undifferentiated cells of tobacco (Avery, 1933) and apple (MacDaniels and Cowart, 1944) near the marginal meristem of the young leaf. In plant No. 78 (pl. 30, fig. 4), the mesophyll reflects somewhat the columnar condition of mesophyll in the young apple leaf during rapid cell division (MacDaniels and Cowart, 1944). In plant No. 26 (pl. 30, figs. 5 and 7), the mesophyll is irregular with some of the upper cells becoming columnar but mesophyll (plant 26) is clarified by Avery (1933) who points out that, in tobacco, epidermal cells acquire their characteristic appearance very early but that [Vor. 40 266 ANNALS OF THE MISSOURI BOTANICAL GARDEN Q 10_ 20 .30 40 50 60 70 80. 90. 100 £ 13,200; 13,800; 14,400r 15,000; 15,699; 16,200r 16,800; 17,400; 18,000r 20,000; 22,200; 25,800r $ 2,400; $ 3,0007 p ANTER 6,600 ^ . — Text-fig. 2. individuals; those : of Phenotypicaly normal plants. Figures at left represent numbers at top, per cent of non-staining pollen. palisade cells do not undergo similar development until the leaf is 4-5 mm. ae Here it seems there has been time enough for some epidermal development béto mesophyll development has been halted. 2d No phenotypic differences were observed between plants grown from $ : arising from treated sperms and untreated eggs and plants from seed from un treated sperm and treated eggs. : dut In general, it appeared that visible radiation effects were associated with a uf degree of pollen non-viability although there were some affected plants w Ж a smaller percentage of non-staining pollen than some with no visible effects. 5 pollen non-viability could occur independently of visible radiation effects apparent from text-fig. 2. 1953] MC QUADE—X-RAY EFFECTS IN PETUNIA 267 It is not suggested that the histograms present a full picture of x-ray effects, since they do not account for the seeds that failed to germinate or individuals that died in the seedling stage. The estimation of pollen non-viability is probably conservative in that pollen grains which stained would not necessarily germinate, or if they germinated might not function properly because of damage of a sort not immediately detectable. For example, plant 37, 20,000 r, with pollen grains of which 48.04 per cent were non-staining did not set seed when selfed; plant 36, 20,000 r, with pollen 64.06 per cent non-staining, set two fruits. SUMMARY 1. Nine hundred and seven of 945 Petunia plants were grown to maturity from seed arising from x-irradiated sperms and untreated eggs or untreated sperms and x-irradiated eggs. Of these, 206 plants showed x-ray effects; 38 died. No effects were apparent in 739 plants. 2. Cytological observations and observations on pollen viability indicated that phenotypic effects of x-ray treatment were usually accompanied by high percentages of pollen non-viability. It was also demonstrated that x-ray treat- ment often resulted in high percentages of pollen non-viability in plants whose phenotypes were normal. No phenotypic differences were observed in plants in which x-ray effects were brought about through treatment of pollen as opposed to ovaries. 4. The leaf morphology resulting from x-ray treatments described above were considered. BIBLIOGRAPHY Avery, = T Je (1933). Structure and development of the tobacco leaf. Am. Jour. E ri 565-592. Lea, D, 947). Actions Аш radiations on living cells. ТҺе MacMillan Со. New Y MacDaniels, L L. H., and Е. Е. Cowart (1944). The development and structure of he apple leaf, Cornell Univ. Agr. Ex ы "Sta . Mem. 258:1-29. McQuade, man A. (1952). The induction of parthenocarpy in Petunia, Ann. Мо. Bot. Gard. 39:97—11 Rick, Charles м. (1943). Cytogenetic consequences of X-ray treatment of pollen in Petunia. Bot. Gaz. 104:528—540. [Vor. 40, 1953] 268 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE 28 1. 20,000 r (Plant 23. See also pl. 30, fig. 1). Light and dark areas on leaf. X about М. 15,600 г (Plant 25). Leaf margins lobed. X about М. 20,000 г. Leaf margins involute. X about М. 18,000 г. Elliptical leaf. X about №. 20,000 г. Spatulate leaf. X about №. . Control X about М. “м > wn» ANN. Мо. Вот. Garp., Vor. 40, 1953 PLATE 28 McQUADE—X-RAY EFFECTS IN PETUNIA ANN. Мо. Вот. GARD., Vor. 40, 1953 PLATE 29 McQUADE—X-RAY EFFECTS IN PETUNIA [Vor. 40, 1953] MC QUADE—X-RAY EFFECTS IN PETUNIA 269 EXPLANATION OF PLATE 29 1. 20,000 г. Calyx members have given way to petaloid structures. In the flower at left, the smaller petaloid structure subtends an anther. th flowers and petaloid structures show characteristic spotting. About natural size. 2. 20,000 г. Globose X about 12. 3. 25,800 г. Abundance of weak leaders. X about 72. 4. 16,200 r. Partial development and distortion of flowers. X about 3⁄4. 5. 17,400 г. Fasciated stem. X about 1⁄2. 6. 9 3,000 г. White flecking of flowers. X about 14. [Vor. 40, 1953] 270 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE 30 ,000 r (Plant 23). Free-hand section through adjacent light and dark areas on ue es (see pl. 28, fig. 1). The thin area at left is a light spot on the leaf. X about . 18,000 r (Plant 29). Section pou : dark area of leaf where palisade and spongy mesophiytle are recognizable. X about 3. 18,000 r (Plant 29). Section through light spot of same leaf shown in fig. 2. X about 70. 4. 20,000 г (Plant 78). Palisade cells squat rather than columnar. X about 70. 5. 25,800 r (Plant 26). Curvature of leaf brought about through difference in eine of upper and lower epidermal layers. Mesophyll cells tend to be homogeneous. EI 6. 20,000 r (Plant 40). Homogeneous mesophyll. X about 70. 7. 25,800 r (Plant ree Detail of fig. 5. Note development of upper as opposed to lower epidermis. X about 35. ANN. Mo. Bor. GARD., Vor. 40, 1953 PLATE 30 McQUADE—X-RAY EFFECTS IN PETUNIA SOME LICHENS ОЕ TROPICAL AFRICA CARROLL WILLIAM DODGE The following account is based largely upon two collections from Nigeria and Cameroons by C. A. Thorold; from Sierra Leone by F. C. Deighton and others, and from Uganda on orchid roots by H. A. Omastin (sent by Sir Edward Salisbury, Royal Botanic Gardens, Kew); one from Nyasaland by Г. J. Brass (Vernay Expedition to Nyasaland) sent by the New York Botanical Garden; and a few from other sources. Using Zahlbruckner's Catalogus Lichenum Universalis and other sources, in all the genera represented, I made keys to the species described from tropical Africa, here defined as the area between 15° N. and 15° S. latitudes and between 20° W. and 55° E. longitudes, but including all of Mozambique and Nyasaland in the east and Angola in the west. I also included the neighboring islands: the Cormoro Archipelago and Socotra, off the east coast, and Ascension, St. Helena, Sáo Thomé, Annobon, Ilha Principe, and Fernando Po, off the west coast. Unfor- tunately, I have not had access to Stirton’s papers in the Transactions of the Glasgow Society of Field Naturalists, which may have included some species from this area. I am including these keys at the appropriate places in hope that they may be useful to other lichenologists who may study material from this area. The countries and islands mentioned in the keys are those from which the species were described and do not indicate the geographic distribution of the species in question. Perhaps the most interesting results are from Acharian species based on the collections of Afzelius in 1792-94 from Sierra Leone and Guinea. I have been unable to locate information regarding Afzelius’ journey, but assume his "Guinea" Was the Guinea Coast proper of geographers of that time, which included the modern Gold and Ivory Coasts although it may have included the whole coast from Senegal to Nigeria. The Nigerian and Cameroons collections are interesting as wholly collected on the bark of Theobroma (the foliicolous species having been sent to Santesson and published in his Foliicolous Lichens I (Symb. Bot. Upsal. 12:1:1-590. 1952). I know of по other such extensive collections from the bark of a single species of tree. In contrast to my experience with collecting on this substrate in Central America, Leptogium is represented by a few unidentifi- able scraps, and the Pannariaceae are absent, both the most conspicuous groups in Central America. Below is a list of the species found on Theobroma: PYRENULACEAE TRYPETHELIACEAE Pyrenula eucalypta Vainio Melanotheca cameroonensis Dodge Pyrenula fuscolurida Vainio Melanotheca nigeriensis Dodge ird beteroclita A La is Dodge yrenula mamillana (Ach.) Trev. Pyrenula trombetana Vainio ж, Dod Antbracotbecium nigeriensis Dodge Pyrenastrum parathelioides ge Issued December 18, 1953. (271) [Vor. 40, ive ANNALS OF THE MISSOURI BOTANICAL GARDEN Сүр ТЛАСЕАЕ 'THELOTREMACEAE Tylopboron ascidioides Vainio Ocellularia cavata (Ach.) Müll. Arg. odge Ocellularia scolecospora D Ocellularia trypanea (Ach.) Dodge CRYPTOTHECIEAE Thelotrema cameroonensis Dodge Crybtotbecia nigeriensis Dodge Tremotylium africanum Ris. Cryptothecia Thoroldi Dodge COLLEMACEAE Collema nigrescens var. minutum Hue OPEGRAPHACEAE LECIDEACEAE Opegrapha nigeriensis Dodge Lecidea granifera (Ach.) Vainio Opegrapha prosodea Ach. — nigeriensis Dodge ci idea rubina Ach. Lecidea tenuis Müll. Arg. а Bacidia golungensis (Vainio) Zahlbr. mE Bacidia nigeriensis Dodge Graphis nigeriensis Dodge Gra PHYLLOPSORACEAE Graphis T boroldi Dodge Pbyllopsora Buettneri (Müll. Arg.) Zahlbr. Phaeographis lynceodes (Nyl.) Zahlbr. wor r Phaeographis ochracea Dodge LECANORACEAE Graphina ulcerata (Vainio) Zahlbr. Lecanora aequinoctialis Stzbgr. Phaeographina deducta (Nyl.) Zahlbr. BLASTENIACEAE Bombyliospora nigeriensis Dodge CHIODECTONACEAE Bombyliospora Thoroldi Dodge Sarcographa labyrinthica (Ach.) Mill. Arg. PHYSCIACEAE Sarcographa Thoroldi Dodge Physcia Poncinsii Hue The usual techniques have been employed in this study, and colors have been recorded from Ridgway's Color Standards and Color Nomenclature. In some of the Graphidaceae with very thin thalli or where the thallus is endophloeodal, the color recorded may owe nearly as much to the color of the underlying bark as to the thallus, and other colors may be expected if the species is found on barks of other species. In the keys, the colors are those of the original descriptions or those recorded by other lichenologists based on the type specimen. The data are still too few to warrant subdivision into floras. From the data at hand, it would seem that the East and West African floras are distinct, but little collecting has been done in the central portion of the continent. There is no apparent relation between the flora of North Africa from Morocco to Egypt and that of Sierra Leone. There is a less marked division between Sierra Leone and Liberia, although a few species apparently extend from Sierra Leone all the way to Angola. In the east there seems to be a division between Abyssinia, Somaliland and Socotra island and the area from Kenya southward. PvRENULACEAE Thallus crustose, uniform, epi- or endophloeodal; cortex often poorly developed or absent; algae Trentepohlia; perithecia solitary in thalline warts, rarely aggtes- gated and somewhat concrescent, but not immersed in a stroma or pseudostroma (distinction from the Trypetheliaceae), erect with a central ostiole; spermatia usually exobasidial. I have referred here three species of Polyblastiopsis and one of Pseudopyrenula with a single perithecium in a true stroma. The absence of a pseudostroma has been used to separate this family from the Trypetheliaceae, but it seems ikely that when we know more of the development and interrelationships we will com- 1953] DODGE—LICHENS OF TROPICAL AFRICA 273 bine these families and split on other combinations of characters as suggested by G. T. Johnson (Ann. Mo. Bot. Gard. 27:1-43. 1940). The phylogenetic sig- nificance of the stroma has also been questioned in the related Pyrenomycetes (Munk, Dansk Bot. Arkiv 15:2:1—163. 1953). 2 EE Paraphyses branched and anastomosing, persistent or evanescent Paraphyses unbranched and free 6 Monobldstia Riddle 2. Ascospores muriform Polyblastiopsis Zahlbr. 3. Spore protoplasts cylindric with thin septa z 4 3% Spore protoplasts rounded to lenticular. Pseudop la Müll. Arg. à atia exobasidial or unknown 5 idi Sperm 4. Spermatia endobasidia 5. Ascospores ellipsoid to fusiform 5. Ascospores acicular, often helically twisted eptorhapis Koerb \ . Ascospores unicellular Coccotrema Mill. 6. Ascospores se е Ascospores murif 11 7. Asci with 1—8 ascospores - 8 7. Asci with many asc Thelopsis ay. ospores 8. Spore protoplasts cylindric, septa thin 8. Spore protoplasts rounded to lenticular, brown . Asci soon evanescent, ascospores acicular. Asci not soon evanescent 10. Ascospores hyaline 10. Ascospores brown 11. Ascospores hyaline, protoplasts appearing сиђіса!............----------------------- 11. Ascospores brown, protoplasts rounded to 1епїїсїаг....................--------—- 'THELOPSIS THELopsis Nyl., Mém. Soc. I. Sci. Nat. Cherbourg 3:194. 1855. Thelocarpon sect. T belopsis Stzbgr., Ber. Thütigk. St. Gall. Naturw. Ges. 149. Pyrenula Ach. Belonia Koerb. 10 № № Porina Müll. Arg. Blastodesmia Mass. Sycbnogonia Kórb., Syst. Lich. Germ. 332. 1855. Holotbelis Clements, Gen. Fung. 40. 1909. Dithelopsis Clements, Gen. Fung. 40. 1909. Type: Thelopsis rubella Nyl. Sycbnogonia, published the same year, was based on S. Bayrboefferi Zw., usually regarded as a synonym of T. rubella. Holo- Фев was based on T. flaveola Arn., as а segregate for the only species with uni- cellular spores, which may belong in Thelocarpon Nyl. Ditbelopsis was based on T. subporinella Nyl. as a segregate with uniseptate spores. Thelopsis inordinata from India might equally well have been segregated as it was described with dwarf muriform spores. Thallus thin, crustose, ecorticate, with Trentepohlia algae, often poorly devel- oped. Perithecia sessile, naked or immersed in a thalline wart, wall relatively soft, light-colored to blackening with a central ostiole; paraphyses slender, free, un- branched (disappearing in T. selemospora Dodge); asci fusiform to oblong, the wall soon disappearing; ascospores usually long-ellipsoid, unicellular to 6-celled or dwarf-muriform, often surrounded with a thin halo. | Pyrenulaceae with polysporous asci have been assembled in this genus without much regard to other morphologic characters. Thelocarpon Nyl. and Thelococcus [Vor. 40 274 ANNALS OF THE MISSOURI BOTANICAL GARDEN Nyl. of the Acarosporaceae have the perithecia immersed in thalline warts with very little development of the perithecial wall, branched paraphyses and proto- coccoid algae. In other characters they seem more closely related to Thelopsis than to other members of the Acarosporaceae. Thelopsis has been reported mostly from temperate Europe and the Mediterranean basin, with one species in southern California and one in India. THELopsis selenospora Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on bark of Amphimas pterocarpoides, Е. C. Deighton M4334C. Thallus hypophloeodes, filamentis Trentepohliae 5—6 u diametro, cellulis 10- 12 р longis et hyphis tenuibus. Perithecia sphaerica, solitaria vel subaggregata, sessilia in disco thallino stramineo, aspera, 0.4—0.5 mm. diametro, dimidiata; para- thecium 70 y crassitudine, pseudoparenchymaticum, cellulis 6-8 м diametro sub- cubicis, obscure brunneis sed non carbonaceis, ostiolo minuto umbilicato, cum cellulis algarum inter hypothecium et cellulas suberaceas corticis; hypothecium lenticulare, 25—30 p crassitudine, hyphis dense contextum; paraphyses evanescentes; asci oblongi, apicibus non incrassati, polyspori, 65-70 X 14—16 р; ascosporae fusiformes vel seleniformes, apicibus acutis, 6-loculares, septis tenuibus, 24 X 3 p. Thallus hypophloeodal except at the base of the perithecia, mostly between the cork layer and the underlying cells of the bark, consisting of Trentepohlia fila- ments and isolated cylindric cells, 10-12 X 5—6 p, somewhat deformed by mutual pressure, and slender hyphae. Perithecia solitary to somewhat aggregated, each sessile on a thin disc of stramineous thallus, surface rough with a minute, incon- spicuous ostiole about 0.4—0.5 mm. in diameter, dimidiate (appearing entire in thick sections since the walls of the cork cells at the base are darkened); wall about 70 y thick, pseudoparenchymatous, cells 6-8 p in diameter, appearing almost cubical, very dark brown but not brittle and carbonaceous, base with algal cells between the hypothecium and the cork cells of the bark; hypothecium lenticular, about 25-30 p thick, of densely woven hyphae; paraphyses early disappearing; asci oblong, tip not thickened, polysporous, 65-70 Х 14—16 p; ascospores fusi- orm to more commonly crescent-shaped, helically twisted in the ascus as in Bacidia subg. Scoliciosporum, ends very acute, predominantly 6-locular, septa very thin, protoplasts nearly cylindric, surrounded by a thin halo, about 24 X 3 № Often one end of the ascospore is prolonged as a short filamentous appendage 4s in the ascospores of the Ashbyaceae. Except for the polysporous asci and evanescent paraphyses, this species might easily be mistaken for a member of Arthopyrenia sect. Pseudosagedia. The much darker perithecial wall, evanescent paraphyses, and long-fusiform to crescent- shaped spores easily separate this species from other described species of T helopsis. : POLYBLASTIOPSIS PorvsLasrioPsis Zahlbr., in Engler & Prantl, Die nat. Pflanzenfam. I. 1*:67. 1903. Polyblastia Müll. Arg., Нога 65:401. 1882, non Lónnr. 1858. Mycoglaena Höhnel, Sitzungsber. К. Akad. Wiss. Wien, math. naturw. Kl 118:1210. 1903. 1953] DODGE—LICHENS OF TROPICAL AFRICA 275 Type: Thirteen species listed, none designated as type. Mycoglaena was based on Verrucaria subcaerulescens Nyl. allus crustose, endo- or epiphloeodal; perithecia solitary, nude or more or les covered by the thallus, hemispheric or spherical, ostiole central; paraphyses branched and anastomosing; asci 1—8-spored; ascospores ellipsoidal to fusiform, muriform, septa thin, with or without a halo, hyaline. Polyblastiopsis differs from Clathroporina only in the branched and anasto- mosing paraphyses instead of unbranched and free. It may also be confused with Laurera of the Trypetheliaceae when the perithecia are crowded. A mig of the whole group is badly nee The systematic position of de species referred here is uncertain. While, in general, the perithecia occur singly, occasionally 2—3 are concrescent, often of quite unequal age, as if a new perithecium started to expand near the base of an older perithecium and the outer portions of the “walls” grew together. A strict morphologic interpretation of the perithecial wall would limit it to the inner carbonaceous wall, and the rest true stromatal tissue, especially since spermogonia occur in it. We have essentially monoperithecial stromata. Such an interpretation would place these species in Laurera in a special section, since the species previously described have several perithecia in each stroma. We have the same situation in Pseudopyrenula Deightoni Dodge in relation to Bathelium. On the other hand, most lichenologists, not being specialists in the Pyrenomycetes, would probably overlook the distinction and look for these species where I have placed them. It is hoped that comparative studies of the development of many more species in both the Pyrenulaceae and тре will eventually place our classification on а sounder basis. 1. Ascospores T P eet 190—220 X 30 п; perithecium entire; Angola......P. fulva (Vainio) De . ^Y spores à Ascpspores 20-34 C49) X 11-13 p, Um Po 6-locellate; due e entire, rring singly in a thick stroma; Sierra Leo P. ф опей. Dofse occu 2. Asc 25—28 10 д, 6—8-locular, кы. ози Soc " Eo . tropica eee Ard ne - Ascospores more than 75 p long; perithecium entir : cnp (75—)8 ees ka 14-16 p, ило, т S ATOS perithecium thin, e Be br locul lurilocellat ‘hates pog Ascospores 305) 100—120 (-126 X 18-26 p, 52-36-loc ar, Pius ocellate, peritheci 120—140 в thick on i чаб) 20—30 м thick at the base; Congo.............. P. baematocbroa Hue x Ascospores 133—160 X 27 m, 24-locular, en c te; оли 4g ш thick, Occurring singly in a spherical stroma; Sierra . sphaerica Dodge v we D tw PoLvsLasrioPsr fulva Dodge, comb. nov. Thelenella (Euthelenella sect. Microglaena) fulva Vainio, Cat. Welwitsch Afric. Pl. 2:451. 1901. Clatbroporina fulva Zahlbr., Cat. Lich. Univ. 1:418. 1922. Type: Angola, Golungo Alto, near Mata Quisuculo, on wild Citrus medica, Welwitsch 230. ahlbruckner was in error in transferring this species to Clathroporina, as Vainio clearly states that the paraphyses are branched and anastomosing, the main character differentiating Polyblastiopsis from Clathroporina. [Vor. 40 276 ANNALS OF THE MISSOURI BOTANICAL GARDEN PorvaLasTIoPsi pyriformis Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on trunk of Anisophyllea laurina, Е. C. Deighton M4404. Thallus epiphloeodes, olivaceus, nigro-marginatus, 160 д crassitudine; cortex 55 p crassitudine, gelifactus, hyphis tenuibus periclinalibus; stratum algarum 80 p crassitudine filamentis Trentepobliae verticalibus, densis, 6-8 p diametro, cellulis subrotundatis; medulla 25 д crassitudine, hyphis tenuissimis. Stromata sessilia, subpyriformia, 0.5 mm. diametro, 0.6 mm. altitudine, basi constricta; cortex stromaticus 15 и crassitudine, hyphis periclinalibus brunneis, stratum interius pseudoparenchymaticum, 30 м crassitudine; parathecium carbonaceum, integrum, 30 р crassitudine; hypothecium lenticulare, centro 40 p crassitudine; paraphyses tenues, dichotome ramosae anastomosantesque; asci cylindrici, dein subfusiformes, 135 X 25 p; ascosporae imbricatim monostichae dein subdistichae, fusiformes, murales, hyalinae, 14—16-loculares, 6-locellatae, 30-34 (—40) X 11-13 u. Spermatia bacillares, 5—6 X 1.5 p. Thallus epiphloeodal, about 160 и thick, dark olive buff with a narrow black margin; cortex 55 и thick, gelified from slender periclinal hyphae; algal layer 80 р thick, of vertical, closely packed filaments of Trentepohlia 6-8 ш in diam- eter, cells somewhat rounded; medulla about 25 p. thick, of very slender, densely woven hyphae, extending some distance farther into the bark cells. Stromata sessile, subpyriform, about 0.5 mm. in diameter, 0.6 mm. tall, constricted at the base; cortex 15 и thick, brownish, of relatively large periclinal hyphae, interior pseudoparenchymatous, about 30 y thick; parathecium carbonaceous, entire, 30 p thick; hypothecium lenticular, about 40 p thick in the center; paraphyses slender, dichotomously branched and anastomosing in the nuclear gel; asci cylindric at first, becoming subfusiform with a rounded tip, about 135 >< 25 р; ascospores imbricately monostichous, becoming subdistichous, fusiform, muriform, hyaline, about 14—16-locular, 6-locellate, protoplasts very slightly rounded, 30—34 (740) нь Spermogonia too old for satisfactory description, arising in the base of the young stroma near the perithecial initial, becoming distorted and flattened against the stromatal cortex as the perithecium expands; wall hyaline or nearly so; sperma- tiophores not clearly seen; spermatia bacilliform, 5—6 X 1.5 p. Sierra Leone: Njala (Kori), on Anisophyllea laurina, Е. C. Deighton M4404, type; on Dialium guineense, Е. C. Deighton M4794. PorvnLasrioPss linearis Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on Citrus aurantifolius, F. C. Deighton M4634. Thallus epiphloeodes, olivaceus, subverrucosus, non nigro-marginatus, 200 р crassitudine; cortex 40 и crassitudine, gelifactus; stratum algarum 90-100 p crassitudine filamentis Trentepobliae verticalibus 8 p. diametro, cellulis ad 16 # longitudine; medulla 65 р crassitudine, hyphis tenuibus. Stromata linealiter dis- posita sed rare concrescentia, sessilia, nuda, subsphaerica, basi constricta, nigra, 1.227 1953] DODGE—LICHENS OF TROPICAL AFRICA 277 1,3 mm. diametro; cortex stromaticus 15 p crassitudine, hyphis pachydermeis inter- textis, obscure brunneis; stratum interior 90—100 p crassitudine, pseudoparenchy- matica ex hyphis periclinalibus 5 д diametro; parathecium 20—25 y crassitudine, integrum, carbonaceum; nucleus sphaericus vel subpyriformis, circa 1000 p diametro; hypothecium 30 д crassitudine; paraphyses tenues ramosae anastomos- antesque; asci cylindrici, pachydermei (ad 18 u crassitudine), dein leptodermei (3—4 м), stipes ad 130 д longitudine, tenuis, venter 510 X 40 y; ascosporae octonae, imbricatim distichae, hyalinae, murales, fusiformes, 14-loculares, 2—3- locellati, (75—) 80 (—85) X 14—16 p, tenui cum halone juventute indutae. Thallus epiphloeodal, citrine drab to deep olive, somewhat verrucose, not black- margined, about 200 д thick; cortex 40 p thick, gelified from subvertical hyphae; algal layer 90-100 д thick, of vertical filaments of Trentepohlia 8 и in diameter, cells about 16 м long, the upper surface of the layer quite uneven; medulla 65 р thick, of densely woven slender hyphae with occasional bark cells; some of the algal filaments penetrating into the bark and forming 2—3 thin layers of thallus between the layers of bark cells. Stromata often closely aggregated in lines but rarely concrescent, sessile, subspherical, nude, very constricted at the base, black, 1.25—1.3 mm. in diameter; stromatal cortex 15 р thick, of dark brown, thick- walled interwoven hyphae with lacunae (probably from old spermogonia); in- terior 90-100 y thick, of brown, relatively thin-walled pseudoparenchyma from periclinal hyphae about 5 и in diameter; parathecium 20-25 р thick, entire, car- bonaceous; nucleus spherical to subpyriform, about 1000 p in diameter; hypothecium about 30 р thick, covering the base of the parathecium; paraphyses slender, branching and anastomosing; asci cylindric and very thick-walled (18 р) when young, becoming more fusiform and thin-walled (3—4 и) at maturity, 8- spored, stipe about 130 p long, slender, venter about 510 Х 40 p; ascospores imbricately subdistichous, hyaline, muriform, fusiform, about 14-locular, 2-3- locellate, with a thin halo when young, (75-) 80 (-85) Ж 14-16 p. Рог үвт.лѕт1орѕ1ѕ sphaerica Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on bark of Orthocosmus africanus, F. C. Deighton M4795. Thallus epiphloeodes, ochraceo-olivaceus, margine tenui, nigro, 100-110 p crassitudine; cortex 40 д crassitudine, gelifactus hyphis periclinalibus; stratum algarum 25—30 p. crassitudine, filamentis verticalibus Trentepohliae, 10-11 p diametro; medulla 40—45 p. crassitudine, hyphis tenuibus dense contexta. Stromata 1-1.5 mm. diametro, subsphaerica, sessilia, basi constricta, ostiolo subumbilicato; Cortex 30 p crassitudine, hyphis periclinalibus obscure runneis; stratum interius 90—95 р crassitudine, pseudoparenchymaticum obscure brunneum; parathecium 50-55 в crassitudine, integrum, carbonaceum; nucleus subpyriformis, gelifactus; hypothecium 40 ш crassitudine hyphis tenuibus dense contextum; paraphyses tenues, ramosae anastomosantesque; asci pachydermei, cylindrici juventute, dein clavati, leptodermei, stipite tenui, 50 р longitudine, venter 300 X 50—55 p; asco- Sporae octonae, subdistichae, hyalinae, murales, 24-loculares, 8-locellatae, halone tenui indutae, 133-160 X 27 р. [Vor. 40 278 ANNALS OF THE MISSOURI BOTANICAL GARDEN Thallus epiphloeodal, between olive ochre and ecru-olive, with a very narrow black margin, 100—110 м thick; cortex about 40 и thick, gelified from predom- inantly periclinal hyphae; algal layer about 25—30 д thick, of vertical filaments of Trentepoblia 10—11 р in diameter, upper surface very uneven; medulla 40—45 p thick, of densely woven slender hyphae, including many disorganized bark cells. Stromata 1-1.5 mm. in diameter, subspherical, sessile, constricted at the base, with a slight depression about the ostiole; cortex about 30 д thick, of periclinal dark brown hyphae; interior 90-95 p thick, of dark brown thin-walled pseudoparen- chyma (black in thick sections with lacunae probably from old spermogonia); parathecium 50—55 м thick, entire, carbonaceous; nucleus highly gelified, sub- pyriform; hypothecium 40 » thick, of slender, densely woven hyphae covering the base of the parathecium; paraphyses slender, branched and anastomosing; asci 8- spored, very thick-walled and cylindric at first, becoming thin-walled and clavate as the spores mature, stipe slender, about 50 u long, venter about 300 X 50-55 р; ascospores subdistichous, hyaline, muriform, about 24-locular, 8-locellate, with a thin halo, 133-160 X 27 p. Deighton M4649 has smaller ascospores (93—) 133 (-146) X (19-) 21 p and a grayer thallus, ie. citrine-drab to deep olive, but is otherwise similar in structure. Sierra Leone: Njala (Kori), on bark of Orthocosmus africanus, F. C. Deighton M4795, type; on bark of Dialium Dinklagii, Е. C. Deighton M4649. PSEUDOPYRENULA PsEUDOPYRENULA Müll. Arg., Flora 66:247. 1883. Type: none designated, 17 species listed. Pyrenula pupula Ach. should be chosen as the type, since it belongs in the section with the largr number of species. Thallus crustose, epi- or endophloeodal with Trentepohlia algae. Perithecia entire or dimidiate, wall carbonaceous, nude or partly covered by the thallus; paraphyses branched and sometimes anastomosing; asci 8-spored; ascospores hyaline, transversely septate, protoplasts rounded, not cylindric. his genus is analogous to Pyrenula, being segregated for its hyaline spores. It differs from Porina Müll. Arg. in its branched paraphyses and the rounded or lenticular protoplasts of its ascospores. In a few species, the perithecia tend to be aggregated but not assembled in a definite pseudostroma as in Bathelium and Trypetbelium. It is widely distributed in the tropics but less common than yrenula in most floras. 1. Perithecium айне nao ый l. Peritheciurà dimüdiste, waospores 4 4 4; cospores 4-locular SPP euin nates ылайы DRC A a не ы OE И E Eo ICE PIE ДЕ КЫ уч o Т ы, „мы X €f м perite hemispheric, black, . ; ^ M perithecial warts emip ГЕЛ к Vainio mm. in diameter i 7 X 7-9 и; perithecial wart 0.5 mm. in diameter; Sio Thome... .3—0.35 3. Ascospores 22—2 3. Ascospores 18-21 X 6-8 u; peritheca very thin i i П ; гу -walled, solitary in a brown stroma, ostiole surrounded by a white disc; Sierra Leone Ке нев P. Deigbtoni Dodge 4. Asc 6-20 X 4—6 д; thallus whitish; Angola... P. bemgoana Vainio 4. Ascospores 23—24 X 5.5-6 д; thallus olivaceous; Angola .P. conica Müll. Arg. р Р 1953] | DODGE—LICHENS OF TROPICAL AFRICA 279 Sect, HOMALOTHECIUM PsEUDOPYRENULA sect. HoMALoTHECIUM Müll. Arg., Bot. Jahrb. [Engler] 6:408. 9 June, 1885. Pseudopyrenula sect. Holotbecium Müll. Arg., Flora 68:331. 11 June 1885. Section Homalotbecium was formally described, limiting the group to species with immersed, entire perithecia, listing Pyrenula annularis Fée, P. neglecta Müll. Arg., P. discolor and P. discolorella. Section Holotbecium was proposed without formal description, treating P. annularis Fée, P. porinoides Müll. Arg., P. Pupula Ach., and P. neglecta Müll. Arg., evidently intended for species with entire peri- thecia. In 1888 (Mém. Soc. Phys. Hist. Nat. Genéve 30:3:28—29), Müller Argau did not use section names, although he treated two species under the heading "Perithecium completum," P. Pupula (about one-third emersed) and P. ceratina (Fée) Müll. Arg. (more innate). Zahlbruckner (in Engler, and Die nat. Pflanzen- fam. 8:78. 1926) limits the section with 4-celled spores, listing P. Pupula (Ach.) Müll. Arg. and P. annularis (Fée) Müll. Arg. Ps—EUDOPYRENULA Deightoni Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on bark of Peltophorum africanum var. speciosum, Е. С. Deighton M4340. Thallus epiphloeodes, isabellinus vel brunneo-olivaceus, margine nigro; cortex 55 n crassitudine, decompositus; stratum algarum 30—60 р crassitudine, filamentis verticalibus Trentepohliae, 5-6 р diametro, cellulis 8 p longitudine; medulla 80— 160 u crassitudine, hyphis tenuibus dense contexta. Stromata 0.5 mm. diametro, 0.3 mm. altitudine, sessilia, nuda, nigra, ostiolo albo-marginato, 70—80 p crassi- tudine, pseudoparenchymatica, brunnea, cellulis 5—6 р diametro; parathecium 5—6 p crassitudine, integrum, carbonaceum; hypothecium 10 р crassitudine, hyalinum, hyphis tenuibus pachydermeis dense contextum; paraphyses 130—150 p altitudine, dichotome ramosae; asci cylindrici, apice non incrassati, 100 X 14 p; ascosporae octonae, imbricatim distichae, ellipsoideae, 4-loculares, protoplastis rotundatis, 5-6 р diametro, juventute cum halone, 18-21 X 6-8 p. Thallus epiphloeodal, isabelline to light brownish olive or darker, with a con- spicuous black margin; cortex about 55 p thick, decomposed; algal layer 30—60 p thick, of vertical filaments of Trentepohlia 5—6 р in diameter, cells about à и long; medulla 80-160 p thick, of slender, densely woven hyphae containing disintegrat- ing bark cells and an occasional algal cell. Stromata about 0.5 mm. in diameter and 0.3 mm. tall, each containing a single perithecium, sessile, nude, black except for the whitish disc about the ostiole; wall about 70 р thick on top and sides of the perithecium, 80 p thick below it, of brown, coarse hyphae forming a pseudo- parenchyma with isodiametric cells 5—6 p in diameter; parathecium very thin, 5-6 u, black, carbonaceous, entire; hypothecium about 10 p thick, hyaline, of densely woven, thick-walled hyphae; paraphyses 130—150 р tall, sparingly dicho- tomously branched, imbedded in the nuclear gel; asci cylindric, 8-spored, about [Vor. 40 280 ANNALS OF THE MISSOURI BOTANICAL GARDEN 100 Х 14 p, tip not thickened; ascospores imbricately distichous, ellipsoidal, 4- locular, protoplasts spherical, 5—6 и in diameter, with a conspicuous halo 3 u thick when young, 18—21 X 6-8 p. This species is related to P. infossa (Nyl.) Zahlbr. from Sáo Thomé, from which it is distinguished by the white disc about the ostiole, the more emergent and smaller "perithecia," and smaller spores. The systematic position of this species is uncertain. I have placed it in Pseudopyrenula because of the hyaline spores and the perithecia solitary in the stroma, but a strictly morphologic in- terpretation would place it in Bathelium, which usually has several perithecia рег stroma. The distinction between the parathecium and stromatal tissue is clear only in very thin sections. It is analogous to the species which I have placed in Polyblastiopsis in their relation to Laurera. Sierra Leone: Njala (Kori), on bark of Peltophorum africanum var. speciosum, Е. С. Deighton M4340, type; on Citrus aurantifolius, Е. С. Deighton M4627. SECT. HEMITHECIUM PsEUDOPYRENULA sect. HEMITHECIUM Müll. Arg., Bot. Jahrb. [Engler] 6:407. Туре: not designated; P. flavicans Müll. Arg., P. diremta (Nyl.) Müll. Arg., P. diluta (Nyl.) Müll. Arg., P. subgregaria Müll. Arg., P. superans Müll. Arg., and P. elliptica were listed. Perithecia dimidiate or base very thin, sessile or slightly immersed at the base, nude, conico-hemispheric; spores 4-locular. PsEUDOPYRENULA BENGOANA Vainio, Cat. Welwitsch Afric. Pl. 2:457. 1901. Type: Angola, Bengo near Quifandongo, on twigs, Welwitsch 437. Thallus epiphloeodal, thin, whitish, smooth, surface dull, without a distinct hypothallus or partly with a black margin, K—. Perithecia solitary, hemispheric, 0.2—0.25 mm. in diameter, nude, black or with a slight thalline covering at the base, ostiole slightly impressed; wall dimidiate, fuliginous; paraphyses branched and anastomosing; asci 8-spored, subcylindric, 40 X 12 p; ascospores distichous, ob- long, hyaline, 4-locular, protoplasts lenticular, equal, 16-20 X 4-6 p. Our specimen has a deep olive-buff thallus with a complete and conspicuous black margin; the asci are about 50—55 X 10-12 y, ascospores 20-22 X 5—6 p with spherical protoplasts 5 » in diameter. In other characters it agrees well with Vainio's description translated above. Since I have not seen the type, I have preferred to refer my material to Vainio’s species rather than describe it as new. Sierra Leone: Njala (Kori), on Peltophorum africanum var. speciosum, Е. С. Deighton M4341, М4343 PYRENULA PyRENULA Ach., Lichenogr. Univ. 64—66, 314—318, 676. 1810, pro parte minore; Mass., Ricerche Autonom. Lich. Crost. 162. 1852. Bunodea Mass., Symmicta, 74. 1855. Type: The selection of the type is very difficult. Of the nine species treated 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 281 by Acharius in 1810, seven have been transferred elsewhere and the two remaining species, P. ocellata and P. subaperta, have not been studied microscopically, hence their systematic position is uncertain. Massalongo, in 1852, was the first to use microscopic characters in the modern sense, and in 1855 he designated P. Alni Mass. as the type and Р. nitida (Weig.) Ach. (1814) as the type of his segregate Bunodea. Koerber (Syst. Lich. Germ. 359. 1855) suggested that P. nitida dif- fered from the other species which he included in the genus and might be taken as the type of a new genus, although he did not propose one. He abandoned the idea later, reducing Bunodea to synonymy in his Parerga Lichenologica 333—334. 1863. Either species would conserve the generic name in its present sense. Thallus crustose, epi- or endophloeodal, ecorticate, with Trentepoblia algae; perithecia solitary or aggregate, nude or immersed in the thallus; wall black, en- tire or dimidiate; paraphyses simple, free; asci 8-spored; ascospores usually 4-locular (2-locular in sect. Pseudacrocordia and 6—10-locular in sect. Fusidiospora) , brown, with lenticular, round or polyhedral protoplasts. This genus differs from Pseudopyrenula in its brown ascospores, and from Melanotbeca by the absence of a clearly developed pseudostroma, in which the perithecia are imbedded. The genus is widely distributed in tropical regions with a few species in the temperate zone. — . Perithecia entire, wholly immersed at maturity except for the ostiolar disc ог papilla................ 2 Perithecia emergent to nearly sessile at maturity. 5 2, нен. flat above, forming a black disc abo ut the ostiole below the level of the € the | lower half of the perithecium sunken in the bark; spore size e Р ocellata Ach. 2. Perthecis Thiet а black; thallus greenish ashy becoming T ‚ @ рейды Т Stein t — * 2 ong n o аты мыс шр си эрес 2. клем зага ascospores 23 X 10—12 p, 4- locular; Socotra....P. obscurata rags red - Base of ice Yer thinner than the sides, sapin 0.7-0.8 mm. in diameter; spo 22-34 5 м; си е Е =t 3 ма Vainio . Base of Е — as thic ade. зы леш у EUN не 4. Spores 25—34 Tor и; y Pede olive: Kenys— eon P. weston ees Müll. Arg 4. POM (17-)23-24 (-29) X 10-13 д; Ee dra mineous; Mozambique....P. ambica Vainio 4. Spores 17-24 X 9-10 u; thallus pale olive; perithecia 0.15—0.3 mm. P diame ter Angola abbr Te & than 20 lose аца Цеа иы ст С [79 w мл 1 a о FE a Ф et g: D в to a t E: et E E: c wn ks] mm E. o “ nw E 8 ritheci meter; ascospores 36—42 X 16—18 ш; Angola... 6. ^W де, т аы thinner than sides; iat celu of ascospores smaller than tke aidie dam Л уу. О ое Perithecial warts 0.5—0.6 mm.; perithecium oblate-spheroid but not pen чт о base; ascospores (25—)36 = (11)15 р; Angola ....... ertet — Vainio . Perithecial warts 0.5-0.7 mm.; perithecium oblate-spheroid, winged at гаф T en spores 30—37 14-16 и; Angola E E тА CE P, сауни Vainio м ч d Perithecial warts 0.7—0.8 mm.; perithecium spherical; ascospores 30—38 X 39 . thallus white; Sierra Leone............ senis parva Vainio m SE deis cb enna инее SAP SEER" wo ed 8 5. 8 p © b со o 8 E 8 Du IN х i * РА > E Е A E Б л 2 MÀ —— [Vor. 40 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN 10. Thallus white or pale ashy; perithecia conic-hemispheric; apical cells of ascospores smaller than middle cells 11 10. Thallus ashy olive; perithecia 0.15-2 mm., flattened-conic, base thinner, winged; ascospores 15— 4.5-5 м; apical cells not smaller; Guinea Р. heteroclita Ach. 10. Thallus green or yellow-green 12 10. Thallus drab to hair brown; perithecia conic, base winged, 0.6—0.7 mm.; nucleus conic; ostiole umbilicate; ascospores 13-14 X 8 и, apical cells not conspicuously Пег; Nigeria Melanotheca nigeriensis Dodge _ — i я. > a [£] - e хл 8 8 Е ase, not winged; ascospores 11-16 X 7-9 ди; thallus whitish, partly black-margined; São Thomé P. glabriuscula (Nyl.) Vainio —0.8 mm., nucleus spherical; ascospores 13—14 X 5-6 ш; thallus pale ashy; Mozambique P. Limae Vainio 12. Perithecia 0.5—0.6 mm., hemispheric with flat base as thick as the sides; ascospores X 8—11 џи, apical cells smaller; thallus yellowish-greenish; Sierra Leone P. aspistea Afz. in Ach. 12. Perithecia 0.6-0.7 mm., flattened-conic with papilla; ascospores 17-21 X 7-8 p, apical cells not smaller; thallus green; Sierra Leone P. тат ава (Ach.) Trev. — м 2 M la] =. et ш о a - PYRENULA TROMBETANA Vainio, Cat. Welwitsch Afric. Pl. 2:454. 1901. Type: Angola, Golungo Alto, near Trombeta, 330—660 m., on bark of Legumi- nosae, Welwitscb 124. Thallus epiphloeodal, pale olive buff with a paler inconspicuous margin, about 125 p thick; cortex 100 д thick, of very slender, periclinal, conglutinate hyphae, the outer 25 и somewhat decomposed and granular; algal layer up to 25 p thick, lying on the outermost bark cells, somewhat discontinuous, of T'rentepoblia fila- ments about 6 и in diameter; medulla not differentiated but fungus hyphae pene- trate between the cork cells and disorganize them. Perithecia 0.6-0.8 mm. in diameter, spherical, nearly innate in the bark until the spores mature, then emersed about one half and appearing hemispheric; wall 55 p thick at the ostiole, expanding to 80 u at the base, then thinning to about 25 и under the thecium, carbonaceous, covered with a layer of thalline cortex about 15 p. thick; ostiole somewhat excen- tric (but not as much so as in Paratbelium) , somewhat umbilicate and paler in old perithecia before the upper half cracks away, first exposing the nuclear remains, then leaving a cupuliform depression; hypothecium scarcely developed; paraphyses slender, dichotomously branched in the thecial gel; asci cylindric, thin-walled, about 135 X 20 y, the wall disappearing before the ascospores mature; ascospores monostichous, broadly ellipsoidal, fuscous, 4-locular, apical protoplasts hemispheric, about 5 р in diameter, central spherical to somewhat angled, 8-9 p in diameter, the whole ascospore 27-32 X 14—16 p. The above description is based on our material. Vainio reports the thallus wholly endophlocodal, the perithecia only 0.5—0.6 mm. in diameter, the ascospores distichous and slightly larger (25-) 36 X (11—) 15 yp, with the apical protoplasts only a little smaller. The thalline characters of P. oculifera Vainio from Angola are closer to our material but the thallus is abruptly thinned leaving an area 0.5—0.4 mm. about the ostiole and the ascospores are somewhat longer and narrower. Nigeria: Ondo Province, Owena near Akure, on Theobroma, С. A. Thorold 171. 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 283 PyRENULA EUCALYPTA Vainio, Cat. Welwitsch Afric. Pl. 2:453. 1901. Type: Angola, Golungo Alto, near Luinha, on Ficus Quebeba Welw., Welwitsch 213 p.p. Thallus epiphloeodal, relatively thick, about 250 u, pale olive buff; cortex 65 п thick, gelified, of densely woven, predominantly periclinal, very slender hyphae; algal layer about 30 џи thick, cells short-cylindric, 5—6 и in diameter, mostly not united in filaments, Trentepohlia; medulla about 155 p thick, of densely woven hyphae, containing occasional disintegrating bark cells and algal cells. Perithecia solitary, immersed in the thallus at first, becoming hemispheric, the lower half covered by thallus and the upper portion by a thin layer of cortex thinning to 10 м about the ostiole through which the black perithecium shows, and lining the ostiolar depression (about 40 p thick) which is about 180 p in diameter, thus appearing white and conspicuous; parathecium about 230 р thick in the middle of the sides, tapering toward the ostiole and toward the base, not produced into a wing at the base in our material, base thinner, about 125 y thick, somewhat flattened (but not as flat as in P. mamillana), carbonaceous and brittle; hypo- thecium scarcely developed; paraphyses slender, dichotomously branched, without conspicuous oil droplets in the thecial gel; asci cylindric, 8-spored; ascospores fuscous, 4-locular, ellipsoid, protoplasts square in optical section, the sides of the middle two about 6-8 р, of the apical ones 5—6 p, the whole ascospore 30—37 K 14-16 р. Cameroons: Kumba, on Theobroma, С. A. Thorold 102. PYRENULA FUSCOLURIDA Vainio, Cat. Welwitsch Afric. Pl. 2:453. 1901. Type: Sierra Leone, near Freetown, on Spondias Mombin L., Welwitsch 247 p.p. Thallus endophloeodal, indicated by a fuscous or testaceo-fuscous area sur- rounded by a black line; perithecia solitary or rarely 2—3-confluent, flattened- hemispheric and only slightly emergent until the ascospores mature then becoming hemispheric and empty as the nucleus disintegrates, 0.5 mm. or more in diameter, dimidiate, base angled but not winged; ostiole minute, not surrounded by a papilla; parathecium about 100 р thick at the base, tapering to about 40 р thick about the ostiole, carbonaceous, brittle; hypothecium very thin so that the asci and paraphyses appear to arise directly from the somewhat darker bark cells; paraphyses branched, apparently somewhat anastomosed above the asci; asci 8-spored, narrowly cylindric; ascospores obliquely monostichous, 4-locular, protoplasts of about equal size, rounded, fuscous, 14—17 X 6-7 p. Our Nigerian material is very old and the spores are shrunken so that no measurements could be obtained. Our material from Sierra Leone is very young and shows an epiphloeodal citrine thallus 80 р thick, cortex 25—50 р thick, of decomposed periclinal hyphae; algal layer 50-75 p thick, somewhat Seno, of more or less disorganized Trentepohlia filaments; medulla not differentiated, but fungus hyphae extend between the bark cells and disorganize them. Young peri- thecia show a thin black wall about 25 р thick, of very slender periclinal hyphae, [Vor. 40 284 ANNALS OF THE MISSOURI BOTANICAL GARDEN surrounded by a brown pseudoparenchymatous layer 50—70 u thick, then an algal layer about 50 м thick, and a cortex 40 u thick of brownish more slender hyphae with granules. As the perithecium matures, the algae die and disintegrate and the whole darkens into the thick parathecium. Mature ascospores agree in size with those described by Vainio. Sierra Leone, Njala (Kori), on Cassia siamea, F. C. Deighton M4793. Nigeria: Ondo Province, Aponmu near Akure, on Theobroma, C. A. Thorold 1722; PvRENULA HETEROCLITA Ach., Syn. Lich. 127. 1814. Type: Guinea, corticole, Afzelius. Thallus epiphloeodal, about 60 р thick, ashy olive; cortex 30 р thick, of slender, conglutinate periclinal hyphae, the outer 10 р decomposed and granular; algal layer about 30 и thick with some of the Trentepohlia filaments penetrating the bark cells beneath; medulla not differentiated. Perithecia 150-250 p in diameter, about 150 y tall, lenticular in opical section, about half emersed at maturity, nude or with remnants of the decomposed cortex, about 5-8 и thick; parathecium about 50 р thick above, winged at the base another 50 и, thinning under the thecium to 25 р, carbonaceous; hypothecium 10 y thick, of very slender, densely woven hyphae; paraphyses slender, dichotomous; asci evanescent; ascospores 4-locular, fusiform to ellipsoid, fuscous, apical protoplasts subconic, only slightly smaller than the subspheric central ones, 15—16 Х 4.5-5.5 The Sierra Leone material has very old perithecia with the thecia disintegrated and only a few shrunken brown ascospores were seen, but such characters as were observable would place it here. The Cameroons material is very scant, only a portion of a thallus growing on Theobroma with P. mamillana (Ach.) Trev., Sarcographa labyrinthica (Ach.) Müll. Arg., and Phacographis lynceodes (ЇЧу!.) Zahlbr. After a study of the type, Müller Argau concluded that the two varieties were growth stages, v. minuscula Ach. being the juvenile stage of the mature v. denigrata Ach. Sierra Leone: Sugar Loaf Mt., 650-750 m., on twigs, F. C. Deighton М4441А. Cameroons: Tombel on Theobroma, C. A. Thorold 136, 138. PYRENULA MAMILLANA (Ach.) Trev., Conspect. Verruc. 13. 1860. Е Verrucaria mamillana Ach., Meth. Lich. 120. 1803. Type: Sierra Leone, corticole, Afzelius. Thallus epiphloeodal, deep olive buff, about 65 p. thick; cortex 15 p thick, de- composed, filled with minute granules; medulla 50 џи thick, of slender, compact, periclinal hyphae with Trentepohlia filaments and cylindric cells 4—5 и in diameter, not clearly aggregated as an algal layer. Perithecia solitary, up to 1 mm. in diam- eter and 0.6 mm. tall, hemispheric with a slight papilla about the ostiole, base flat, surface dull and very minutely and shallowly pitted (seen only under 36X magnification) ; parathecium 50 y thick at the ostiole, rather abruptly thickening to 100 р and then tapering to 130 y at the base and prolonged another 130 p as 1953] DODGE—LICHENS OF TROPICAL AFRICA 285 wings, thinning to 40-50 и under the hypothecium, the whole carbonaceous and brittle; hypothecium scarcely differentiated, groups of asci radiating from various points as if produced by different ascogonia; asci cylindric, 8-spored, about 110 Х 8-9 р; paraphyses slender and dichotomously branched, the thecial gel filled with oil droplets; ascospores ellipsoid, fuscous, 4-locular, 17-21 7-8 p, protoplasts rounded, central ones about 3 и in diameter, apical ones somewhat smaller. Sierra Leone: Kanema (Nongowa), on Copaifera copallifera, F. C. Deighton M5013. Nigeria: Ojo Province, Iseyin, on Theobroma, C. A. Thorold 103. Cameroons: Tombel, on Theobroma, C. A. Thorold 136. ANTHRACOTHECIUM ANTHRACOTHECIUM Hampe in Mass., Atti I. R. Ist. Veneto III, 5:330. 1860. Bottaria sect. Anthracothecium Vainio, Etude Lich. Brésil 2:196. 1890. Type: A. Doleschalii Mass. Thallus uniform, crustose, endo- or epiphloeodal, with Trentepohlia algae. Perithecia solitary or sometimes aggregated but not forming a pseudostroma, usually nearly covered by the thallus; parathecium carbonaceous (sect. Euanthraco- thecium) or light-colored with a darkened area about the ostiole (sect. Porin- astrum) , usually entire; paraphyses unbranched and free; asci 1-8 -spored; ascospores ellipsoidal, muriform, with rounded protoplasts, brown; spermatia acicular, often curved, Since А. euthelium (Nyl.) Zahlbr. and A. lugescens (Nyl.) Zahlbr. were described as having hyaline, muriform spores, they may belong in Clathroporina rather than in Anthracothecium where Zahlbruckner transferred them. 1. Spores 7-12 X 5-7 и, dwarf-muriform, 4-locular with the two middle cells divided by a longitudinal septum; Usambara ........—.————7 l. Spores less than 1 on Leo ARCHOS UU ANUS MEN Cu IRIS d - : 1. Spores over 200 u long, hyaline; Sao Thomé (perhaps belong in Clatbroporina) ............. m 2. Thallus white; perithecia about 0.1 сас Фатен c eoo A. punctuliforme Müll. Arg. 2. Thallus orange-yellow; perithecia 0.7 mm. in diametet.......----------------- X 22-25 д; thallus ashy, black-margined; A. cinerosu: es 80-95 Х 32-35 д; thallus pale olive buff to smoke gray; perithecia 2.5 mm. in diameter; Ilha Principe ................. rre tnm A. guineense (Nyl.) Zahlbr. 4. Spores 210—230 X 45-75 и; asci monosporous; perithecia 0.5 mm. diameter........ оон що оони ноте от оне nn f м ANTHRACOTHECIUM GUINEENSE Zahlbr., Cat. Lich. Univ. 1:462. 1922. "Verrucaria guineensis Nyl., Lich. Insul. Guineens. 56. 1889. Type: Ilha Principe, at sea-shore, corticole, Quintas. Thallus epiphloeodal, pale olive buff to smoke gray, about 65 p thick; cortex 40 p thick, of slender, conglutinate, periclinal hyphae; algal layer discontinuous, up to 25 y thick, cells about 6 р in diameter, not in distinct filaments and some- What angular from mutual pressure, Trentepoblia; medulla not developed, but the fungus hyphae penetrating deeply into the bark. Perithecia solitary, about 2.5 [Vor. 40 286 ANNALS OF THE MISSOURI BOTANICAL GARDEN mm. in diameter, 0.9 mm. tall, covered with a thin layer of thalline cortex to near the ostiole; parathecium 125 y thick near the ostiole, expanding to 600 p thick at the base, 125 и thick under the hypothecium, carbonaceous, with a very dense inner layer about 125 p thick next the nucleus, the rest of thick-walled pseudoparenchyma with occasional small cavities (remains of old spermogonia?) ; nucleus subspherical, becoming conic in very old perithecia, 1200—1300 џ in diam- eter; hypothecium 50 и thick, of densely woven, deeply staining, slender hyphae; paraphyses slender, dichotomously branched in the thecial gel; asci cylindric, dis- appearing before the spores mature; ascospores brown, broadly ellipsoid, 14-locular, 8-locellate, 80-95 Ж 32-35 y. The relation of this species to А. стетозит (Ach.) Müll. Arg. is not clear. It agrees in most characters with Müller Argau's description of the Acharian type except in spore size. Азсозрогез, already free of the ascus and just beginning to assume the smoky color of immature brown spores, fall within the measurements given for А. cimerosum. They apparently increase in size at the expense of the thecial gel as they mature and become dark brown. If Müller Argau measured such spores and failed to find the very dark brown mature spores, А. guineensis may be a synonym of A. cinerosum (Ach.) Müll. Arg. I referred my material to the latter species before I found the very dark brown spores in another perithecium on the same thallus. Only a developmental study of these species can settle the synonymy. Sections of one “perithecium” of Thorold 104 show two perithecia concrescent with a common wall and ostiole, and might be mistaken for Par- mentaria Cbevalieri Bouly de Lesdain. Nigeria: on Theobroma, С. A. Thorold 104; Aponmu near Akure, on Theo- broma, C. A. Thorold 172. TRYPETHELIACEAE Thallus crustose, uniform, epi- or endophloeodal; ecorticate or corticate (never pseudoparenchymatous) ; algae Trentepohlia; several perithecia (rarely only 1 or 2) immersed in each well-developed pseudostroma, erect, with central, individual ostiole; spermatia exobasidial. 1. Азсозрогез rata Tade eo 2 a 1. Акорда BOW ме а ИЙ : 2. Ascospores unicellular .................. -Riddles Dew T Asc — ез septate, usually 4- or more celled........... TS серое а o Laurera aci. 3. pies thin-walled, "n cylindric or nearly so. Tomasellia E 3. Ascospores thick-walled, protoplasts rounded or lenticular 4. Ascospores rag а. быт, E 4. Ascospores 6—many-locular, large . 5. Ascospores septate, usually 4 or more called: protoplasts rounded or lenticular...... ati z e" ee er rrr rere rere rrr 1953]: DODGE—LICHENS OF TROPICAL AFRICA 287 RIDDLEA Riddlea Dodge, gen. nov. Type: R. papillosa Dodge. Thallus crustosus, epiphloeodes; cortex decompositus; algae Trentepoblia. Pseudostromata carnea; perithecia immersa, integra; hypothecia in pulvinulis lenticularibus, plura in quovis perithecio; paraphyses dichotome ramosae anasto- mosantesque; asci longe stipitati; ascosporae octonae, hyalinae, fusiformes, uniloculares; spermogonia in verrucis thallinis immersa; perifulcrum nigrum; spermatiophorae ampullaceae; spermatia acicularia, recta. Thallus epiphloeodal, sometimes separating from the bark and appearing sub- foliose, but without a lower cortex; cortex decomposed; algae Trentepohlia. Pseudo- stromata soft and fleshy, thalline when young, the algae finally dying and leaving lacunae, variable in shape; perithecia immersed in the pseudostroma, ostioles cen- tral; parathecium black; several hypothecia per perithecium, lenticular; paraphyses dichotomously branched and anastomosing; asci long-stipitate, very thick-walled when young, the wall thinning as the ascospores mature, 8-spored; ascospores hyaline, fusiform, unilocular; spermogonia immersed in thalline warts, wall black; spermatothecium folded into labyrinthiform cavities; spermatiophores flask-shaped; spermatia acicular, straight, relatively short. e fungus component of this genus does not seem closely related to other lichen-forming fungi. The placenta-like cushions from which the long-stalked asci radiate are suggestive of the Coronophorales, but true paraphyses are present and the pseudostroma is quite different. Although the pseudostroma is essentially thalline (i.e. containing algae), it shows little relation to Coccotrema, which per- haps belongs in the Pertusariaceae (close to or the same as Perforaria) rather than in the Pyrenulaceae. For the present we include Riddlea in the Trypetheliaceae, analogous to Monoblastia of the Pyrenulaceae, although the relationship does not seem close. I take pleasure in dedicating this genus to my late friend, Professor Lincoln W. Riddle of Harvard University, who first described Monoblastia. Rwoea papillosa Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on bark of Anisophyllea laurina, F. C. Deighton M4624. Thallus epiphloeodes, pallide flavo-olivaceus, 75 p crassitudine; cortex 10 p crassitudine, decompositus, minutis cum granulis obscuris; stratum algarum 65 р crassitudine, filamentis verticalibus Trentepobliae, 6.5 p diametro, cellulis cylindricis isodiametricis, dense compactis; medulla non evoluta. Pseudostromata carnea, thallina cum cellulis algarum inter perithecia morientibus, dein lacunata, alia sub- Sphaerica ca. 1 mm. diametro, alia in pulvinulos applanatos 2—3 mm. diametro, alia sublinearia, hyphis hyalinis 2 р diametro, cortice thallino tecta; perithecia 370 р diametro, 250 y altitudine, oblate sphaeroidea, cervice 100 р longitudine Superne tenuescente ad ostiolum parvum in papilla applanata; parathecium in- [Vor. 40 288 ANNALS OF THE MISSOURI BOTANICAL GARDEN tegrum, nigrum, 15 y crassitudine; hypothecia plura in quovis perithecio, pulvini lenticulares, 135 и diametro, centro 40 y crassitudine; paraphyses tenues, dicho- tome ramosae anastomosantesque, apicibus liberis; asci 200 Ж 30 y, juventute pachydermei, stipitibus 100 X 5 и, venter clavatus vel fusiformis, 100 X 30 y; ascosporae octonae, distichae, hyalinae, fusiformes, uniloculares, 24-32 X 8-11 p, tenui cum halone. Spermogonia in verrucis thallinis immersa, sphaerica, 250 p diametro; perifulcrum nigrum, 15 y crassitudine, hyphis tenuibus periclinalibus; spermatothecium plicatum; spermatiophorae ampullaceae, 10 p longitudine, venter 2 р diametro, cervice elongato tenuescente; spermatia acicularia recta, ca. 10 X 1 p. Thallus crustose, epiphloeodal, light yellowish olive, about 75 p thick, some- times separating from the bark and appearing subfoliose but lacking a lower cortex; upper cortex about 10 u thick, decomposed, with minute dark granules (appearing black in thick sections) ; algal layer 65 м thick, of Trentepohlia filaments, vertical, closely packed, cells cylindric, more or less isodiametric, about 6.5 р in diameter; medulla not differentiated but hyphae penetrate some distance into the bark cells. Pseudostromata soft, fleshy, of thalline tissue, ie. containing masses of algal cells between the young perithecia, dying and leaving lacunae, very variable in size and shape, some subspherical, about 1 mm. in diameter, others in flattened cushions 2—3 mm. in diameter while others are elongate and sublinear, composed of closely woven hyaline hyphae about 2 и in diameter, with masses of algal cells between the young perithecia, covered by the thin thalline cortex; perithecia 370 р in diameter, 250 р tall, oblate-spheroidal, with a short neck (about 100 p long), tapering upward to a small ostiole surrounded by a low, broad papilla; parathecium entire, black, 15 р thick; hypothecia several per perithecium, as lenticular, placenta- like cushions about 135 и in diameter and 40 р thick in the center, from whic tufts of asci and paraphyses radiate into the thecial gel; paraphyses slender, dicho- tomously branched and anastomosing, tips free; asci 200 X 30 p, very thick- walled when young, the lower half a stipe about 5 р in diameter, the upper half clavate to fusiform, the wall thinning as the ascospores mature, 8-spored; asco- spores distichous, hyaline, fusiform, unilocular, 24—32 X 8-11 p, with a thin halo. Spermogonia immersed in thalline warts, about 250 p. in diameter, wall black, 15 в thick, of slender periclinal hyphae; spermatothecium folded into labyrinthiform cavities; spermatiophores about 10 p long, flask-shaped with a long tapering neck, venter about 2 y in diameter; spermatia acicular, straight, about 10 X 1 p- In habit Riddlea papillosa suggests BATHELIUM papillosum (Ach.) Dodge comb. nov. (Trypethelium papillosum Ach., Syn. Lich. 104. 1814), and I had ten- tatively referred my material to that species until I found the mature unicell ascospores, since the ascospores while still in the ascus are about the size of those of B. papillosum before the septa are visible. BATHELIUM BaTHELIUM Ach., Meth. Lich. 111. 1803, non Mass. 1860. Leightonia Trev., Flora 44:19. 1861. Trypethelium sect. Bathelium Müll. Arg., Bot. Jahrb. [Engler] 6:589. 1885. 1953] DODGE—LICHENS OF TROPICAL AFRICA 289 Pseudopyrenula subg. Trypetbelium sect. Batbelium Vainio, Etude Lich. Brésil 2:206, 1890, Type: B. mastoideum Afzelius in Ach. Leightonia was based on Trypethelium porosum Ach. Thallus crustose, epi- or endophloeodal, cortex thin, gelified, algae Trentepohlia. Pseudostromata hemispheric or more flattened and irregular; perithecia ellipsoid to spherical, wall entire; asci 8-spored; ascospores hyaline, ellipsoid, 2—4-locular, protoplasts rounded. l. Pseudostromata often containing a single perithecium; ascospores 20 X 8 и, apical cells larger than middle cells; Mozambique B. duplex f. simplicius (Vainio) Роби 1. Pseudostromata normally containing several perithecia : 2. Papillate about the ostiole [papilla sometimes breaking away in very old perithecia]; thallus ashy olive to fuscous 3 2. Not papillate about the ostiole; thallus white or nearly so. 4 2. Upper part of the pseudostroma breaking away forming ап ostiole 250 д in diameter; thallus deep olive buff; ascospores 19-21 X 8-9 и, long remaining hyaline but nally brown when fully mature; Sierra Геопе........................... Melanotbeca porosa Dodge 3. Ascospores 18—22 X 7-8 ш; Sierra Leone mastoideum Ach. 3. Азсозрогез 20-27 Х 8-9 ш; Guinea В. papillosum (Ach.) Dodge 4. Ascospores 20 X 8 д, with halo; МоғатЫдие.................--------- B. compositum (Vainio) Dodge 4. Ascospores 28—33 X 9-10 ш; São Thomé. B. subalbens (Nyl.) Dodge BATHELIUM MASTOIDEUM Afzelius in Ach., Meth. Lich. 111. 1803. Trypetbelium mastoideum Ach., Lichenogr. Univ. 307. 1810. Type: Sierra Leone, corticle, Afzelius. Thallus epiphloeodal, 100 ш thick, deep olive buff, conspicuously black-mar- gined; cortex 55 и thick, of vertical, slender interwoven hyphae in a gel; algal layer 45 р thick, of predominantly periclinal hyphae and disorganized filaments of Trentepoblia in a gel; medulla not differentiated but hyphae penetrating the under- lying bark cells. Pseudostromata hemispheric, confluent into irregular masses containing 1—10 perithecia, fuscous, yellow within at first, blackening and car- bonaceous in age, with a low papilla about the ostiole which soon cracks off; parathecium fusing with the stroma; nucleus rounded at first, becoming subconic, 80 u in diameter at the base, 100 p at the ostiole and 185 p tall, the pseudostroma extending 80 и below the nucleus and resting on the bark cells; hypothecium 12-14 р thick, of slender, closely woven, predominantly periclinal hyphae; para- physes dichotomously branched and anastomosing in the nuclear gel, about 150 p tall; asci fusiform with rounded tips, 8-spored, about 80 X 14—16 р; ascospores distichous, hyaline, ellipsoid, 4-locular, with large, subequal, rounded protoplasts, 18-22 X 7-8 p. Sierra Leone: Njala (Kori), on bark of Anisopbyllea laurina, F. C. Deighton 4625. BarHELIUM compositum Dodge, comb. nov. Pseudopyrenula composita Vainio, Bol. Soc. Broter. II, 6:177. 1929. Trypetbelium compositum Zahlbr., Cat. Lich. Univ. 10:101. 1938. [Vor. 40 290 ANNALS OF THE MISSOURI BOTANICAL GARDEN BATHELIUM DUPLEX f. simplicius Dodge, comb. nov. Pseudopyrenula duplex £. simplicior Vainio, Bol. Soc. Broter. II, 6:177. 1929. Trypethelium duplex f. simplicius Zahlbr., Cat. Lich. Univ. 10:101. 1938. BaTHELIUM papillosum Dodge, comb. nov. Trypetbelium papillosum Ach., Syn. Lich. 104. 1814. BATHELIUM porosum Dodge, comb. nov. Trypetbelium porosum Ach., Syn. Lich. 106. 1814. Verrucaria porosa Eschw. in Martius, Fl. Brasil. 1:135. 1833. Leigbtonia porosa Trev., Flora 44:19. 1861. Trypetbelium Sprengelii v. porosa Nyl. in Hue, Nouv. Arch. Mus. [Paris] Ш, 29. 1892 BaTHELIUM subalbens Dodge, comb. nov. Trypetbelium subalbens Nyl., Flora 69:178. 1886. TRYPETHELIUM TRYPETHELIUM Sprengel, Einleitung in das Studium der kryptogamischen Gewáchse, 350. 1804 [often cited as Anleitung zur Kenntniss der Gewichse 3]. Trypetbelium sect. Eutrypetbelium Müll. Arg., Bot. Jahrb. [Engler] 6:393. 1885. Pseudopyrenula subg. Trypethelium sect. Eutrypethelium Vainio, Etude Lich. Brésil 2:204. 1890. Type: T. Eluteriae Sprgl. Thallus crustose, epi- or endophloeodal, cortex gelified, thin; algae Trentepoblia. Pseudostromata hemispheric, or flattened and irregular, usually of a different color than the thallus, containing 2 to many perithecia, which are ellipsoidal or spherical, ostiole central, parathecium entire, carbonaceous; paraphyses branched and anas- tomosing; asci 8-spored; ascospores long-fusiform, hyaline, 6—22-locular, proto- plasts rounded. i2 ideni i blood-red, surface pruinose; thallus ashy; ascospores unknown, so perhaps Bottaria or Melanotbe hich h н са which have species with red чт rie y Seber. — à Peeudortromata ; not blood-red ..... Ascospores acicular, 18—22-locular, 56 X 2.5 B Мужа на... T. “aciculare Е * Ascospores much broader, ellipsoid to fusiform rri у e PA etr 188 X 42 д; thallus and ун. olive-glaucescent; 4 v 3 Я 5 HE д^ Т 4 оѕрогеѕ нра 55 X 14 п; thallus olive ochre анааан y ow ochre, dark fus within, 0.5 mm. " e and duh Guinea....T. anomalum Ach. 4. Ascospores 14-locular 80-85 X 14- thallus n aristis reds qe stromata 1 mm. tall, subspheric, finally ‘black with 2—6 perithecia; Angola......-.---- ны E енене Күш) Zahlbr. 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 291 TRYPETHELIUM aciculare Dodge, sp. nov. Type: Nyasaland, Kasungu Hill, 1100 m., corticole, L. J. Brass 17458а. Thallus epiphloeodes, glaber, subrimulosus, 135 u crassitudine, olivaceus, mar- gine plumoso, 3 mm. latitudine, homoeomerus; filamentis Trenfepobliae plus minusve verticalibus, 6-8 д diametro. Pseudostromata thallina, pulvinata, irreg- ulariter rotundata, 1 mm. diametro, cum 12-20 peritheciis; perithecia ellipsoidea; parathecium integrum, 12-15 д crassitudine, pseudoparenchymaticum, ex hyphis periclinalibus superne obscurum, inferne hyalinum; hypothecium non bene evo- lutum; paraphyses dichotome ramosae anastomosantesque, apicibus liberis, 1.5 p diametro; asci cylindrici pachydermei juventute, 60 X 8 p; ascosporae octonae, fasciculatim dispositae, hyalinae, 18—22-loculares, protoplastis rotundatis, aciculares, subcurvatae, apicibus obtusis, 56 Х 2.5 p. Thallus epiphloeodal, somewhat rimulose, 135 p thick, citrine drab to deep olive, margin 3 mm. wide, of radiating, plumose strands; homoeomerous, the fila- ments of Trentepohlia more or less vertical, 6-8 и in diameter. Pseudostromata thalline, pulvinate, irregularly rounded, 1 mm. in diameter, containing 12-20 ellipsoidal perithecia; parathecium entire, 12-15 и thick, pseudoparenchymatous from periclinal hyphae, the upper portion dark brown shading to hyaline below; hypothecium scarcely differentiated; paraphyses dichotomous and anastomosing, 1.5 р in diameter, tips free in the thecial gel; asci cylindric, thick-walled when young, 8-spored, about 60 X 8 y; ascospores fascicled, hyaline, 18—22-locular, protoplasts rounded, acicular, slightly curved, ends obtuse, 56 X 2.5 p. TRYPETHELIUM ANOMALUM Ach., Syn. Lich. 105. 1814. Type: Specimens cited from the West Indies and Guinea, Afzelius. Müller Argau borrowed the Guinea specimen and found it a true Trypetbelium, reducing it to synonymy with the later T. platystomum Mont. Vainio, presumably study- ing the West Indian material, referred it to Melanotheca Acbariana Fée. Zahl- bruckner followed the Müller Argau tradition and recognized the species as a true Trypetbelium. Acharius’ choice of the specific name is unfortunate as it is not an anomalous species in either Trypetbelium or Melanotbeca. Thallus epiphloeodal, olive ochre, about 100 р thick; cortex 30 р thick, of predominantly periclinal, interwoven hyphae in a gel; algal layer 70 p thick, of short, more or less vertical filaments of Trentepohlia. Pseudostromata yellow ochre, rounded, about 2 mm. in diameter or elongate and irregular from con- fluence, slightly constricted at the base, with many black papillae which fall away, leaving small pits; perithecia ellipsoid, about 360 p in diameter and 500 p tall; parathecium 30 р thick, carbonaceous, surrounded by brown pseudostromatic tissue and covered by thalline cortex; hypothecium about 15 и thick, of slender, densely woven hyphae; paraphyses dichotomous and anastomosing, tips free in the thecial gel; asci fusiform, 8-spored; ascospores imbricately distichous, hyaline, fusi- form, one end obtuse, the other acute, 10—14-locular, protoplasts slightly rounded, 55 X 14 p [Vor. 40 292 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN T. Perrotetii Fée (Ann. Sci. Nat. 23:432. 1831) may be a synonym, as Miller Argau reports that the type from Senegal, Cap Vert, Perrotet, is old and contains no spores. Т. leucostomum (Nyl.) Dodge has white ostioles and much larger spores (85—110 X 15-18 д). Sierra Leone: Njala (Kori), on bark of Anisophyllea laurina, Е. С. Deighton M4407. TRYPETHELIUM leucostomum Dodge, comb. nov. Trypethelium platystomum f. leucostomum Nyl., Flora 69:178. 1886. Trypetbelium anomalum f. leucostomum Zahlbr., Cat. Lich. Univ. 1:487. 1922. MELANOTHECA MELANOTHECA Fée, Suppl. Essai Crypt. Ecorces Officin. 70. 1837. Arthopyrenia sect. Melanotbeca Stzbgr., Ber. Thatigk. St. Gall. Naturw. Ges. 147. Segestrella sect. Melanotbeca Branth & Rostr., Bot. Tidsskr. 3:257. 1869. Pyrenula subg. Melanotheca Vainio, Cat. Welwitsch Afric. Pl. 2:465. 1901. Porotbelium Eschw., Syst. Lich. 18. 1824, non Fr. 1818, 1821. Porodotbion Fries, Syst. Orb. Veg. 262. 1825. Type: М. Achariana Fée (based on Trypetbelium anomalum Ach., West Indian plants cited, not Afzelius’ Guinea plant). Porothelium Eschw. and Porodothion Fr. were based on P. arthonioides, Porina compuncta Ach., Try petbelium anomalum Ach., and T. conglobatum Ach., all except P. arthonioides and part of T. anomalum belonging in Trypetbelium as now understood. Since Melanotheca has been used in its present sense, either as a genus or subgenus since it was proposed, it should be conserved. It would be unwise to select P. arthonioides as the type of Poro- dotbion, although it was figured by Eschweiler as the invalid Рото Бейит (renamed Porodotbion by Fries) as all species of the present Melanotbeca would have to be transferred to Porodotbion. Thallus crustose, epi- or endophloeodal; cortex often poorly developed or ab- sent; algae Trentepoblia. Stroma or pseudostroma usually with several perithecia, irregularly rounded or sublinear; perithecia immersed; parathecium carbonaceous, ostiole central; paraphyses either unbranched or branched and anastomosing; asci usually 8-spored; ascospores brown to black, ellipsoidal or fusiform, 4- or more celled, with rounded protoplasts; spermatiophores simple; spermatia filiform, straight or curved. 1. : Pseudostromata purple; ascospores 4-locular, 24 X 12 p; Usambara....M. purpurascens Mall. Arg. Pseudostromata cinnabar red to red; ascospores 4-locular, 20 X 8 и; Ken 1. Pseudostromata not purple or red, usually dark-colored 2. Ascospores 10-locular; pseudostromata small; Abyssinia M. pusilla (Jatta) рей А res 4-locular; pseudostromata larger : 3. Ascospores 15 Ш or less long 3. Ascospores more than 17 p long oases E N E REDE e EÊ sesatasonssaassetn 1953] DODGE—LICHENS OF TROPICAL AFRICA 293 4. Ascospores narrowly ellipsoid, 15 X 4—5 ш; thallus fuscous olive; Mozambique........ M. obs 4. Ascospores broadly ellipsoid, 13-14 X 8 д; thallus drab to hair brown; Nigeria........ nigeriensis Dodge 5. Perithecia entire; ascospores 19—21 X 8—9 и, apical cells much smaller than middle ones; top of perithecium cracking away to leave an opening about 250 д in diameter; M. porosa Dodge Sierra Leone 5. Perithecia dimidiate or nearly so 6. Ostiole white-annulate; ascospores 17—21 X 6-7 д; Cameroons.......... M. cameroonensis Dodge 6. Ostiole small, not white-annulate; ascospores 17—19 X 5.5-8 u; Angola M. angolensis (Vainio) Dodge MELANOTHECA angolensis (Vainio) Dodge, comb. nov. Melanotheca Achariana var. angolensis Vainio, Cat. Welwitsch Afric. Pl. 2:453. 1901 Type: Angola, Golungo Alto, near Sange, on Entandrophragma angolensis, Welwitsch 205. Pseudostromata thin, ostiole small, without a white annulus; nucleus almost hemispheric or depressed conoid-subspherical; paraphyses unbranched; asci 8- spored; ascospores brown, 4-locular, protoplasts lenticular, 17-19 X 5.5-8 p. Welwitsch 187, from the type locality and on the same species of tree, is reported to have dimidiate perithecia and perhaps belongs in M. cameroonensis Dodge, but I have not seen this specimen. The whole group of species centering about Melanotheca Achariana Fée needs a thorough revision based on the types of all the species and varieties proposed. MELANOTHECA nigeriensis Dodge, sp. nov. Type: Nigeria, Ondo Province, Owena near Akure, on Theobroma, C. A. Thorold 170. Thallus epiphloeodes, brunneus, 90—110 p crassitudine; cortex 27—55 p crassi- tudine, decompositus, hyphis tenuissimis, verticalibus, dense intertextis, gelifactis; stratum algarum 55—65 p crassitudine, filamentis subverticalibus T'rentepobliae 7-8 и diametro. Perithecia (aut stromata cum peritheciis singulis aut binis) solitaria vel confluentia, hemispherica, nucleo conico, 0.6 mm. diametro, 0.4 mm. altitudine; parathecium (aut stroma) 125 p crassitudine ad ostiolum centralem, ad basem 240 и grandescens, 65 p crassitudine sub hypothecio, carbonaceum, cortice thallino 10 p crassitudine usque ad ostiolum tectum; ostiolum subumbili- catum; hypothecium 13-14 y crassitudine, hyphis dense intertextis; asci cylindrici, evanescentes; ascosporae octonae, late ellipsoideae, obscure brunneae, 4-loculares, protoplastis rotundatis, subaequalibus, 13—14 8 p. Thallus epiphloeodal, drab to hair brown, 90—110 р thick; cortex 27-55 p thick, decomposed, of densely woven predominantly vertical, very slender, gelified hyphae. Perithecia (or stromata with one or two perithecia) solitary or aggregated into small groups, lentiform, becoming hemispheric with a conic nucleus about 0.6 mm. in diameter, 0.4 mm. tall; parathecium (or stroma) 125 p thick at the ostiole, expanding to 240 џи thick at the base, about 65 p thick under the hypo- thecium, covered by a thin layer of thalline cortex 10 р thick all the way to the [Vor. 40, 294 ANNALS OF THE MISSOURI BOTANICAL GARDEN slightly umbilicate ostiole, carbonaceous; hypothecium 13—14 р thick, of densely woven hyphae; asci cylindric, 8-spored, soon disappearing; ascospores broadly ellipsoid, dark brown, 4-locular, protoplasts rounded, subequal, 13-14 X 8 p. The systematic position of this species is intermediate between Pyrenula and Melanotbeca. When occurring in pairs, the perithecia appear innate in a carbona- ceous stroma from the complete fusion of the parathecia and the stromatal tissue. Occasionally I have found abortive perithecia in the angles at the base when the perithecia appear solitary. Unfortunately, I have had no young material to study development. I have therefore included this species also in the key to the tropical African species of Pyrenula. МЕГАМОТНЕСА porosa Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on bark of Anisophyllea laurina, F. C. Deighton M4408. Thallus epiphloeodes, glaber, obscure olivaceo-alutaceus; cortex 120—130 p crassitudine, hyphis periclinalibus tenuibus dense intertextis, gelifactus, paucis cum cellulis suberosis hyalinis; stratum algarum 55 p crassitudine, filamentis disinte- gratis Trentepohliae inter cellulos suberosos decompositos; medulla non evoluta. Pseudostromata parva, irregulariter rotundata aut frequenter confluentia, lenticu- laria, 0.5 mm. crassitudine, sublinearia curvataque; perithecia immersa, ostiolis ad 250 р diametro; nucleus са. 250 p diametro, 300 д altitudine, pyriformis; para- thecium integrum, carbonaceum, 15 y crassitudine; hypothecium 25 p crassitudine, yphis periclinalibus gelifactis; paraphyses tenues, dichotome ramosae, compactae; asci 65 X 13-14 p; ascosporae octonae, imbricatim monostichae, hyalinae dein brunneae, late ellipsoideae, 4-loculares, protoplasti lenticulares, apicales conspicue minores, 19—21 X 8-9 y. Thallus epiphloeodal, smooth, deep olive buff; cortex 120—130 p thick, of slender, interwoven, mostly periclinal hyphae in a gel, with hyaline remains of bark cells in the lower portion; algal layer 55 p thick, of disorganized filaments of Trentepoblia among the remains of bark cells; medulla not differentiated but hyphae penetrating deeply into the brownish bark cells. Pseudostromata sma”; irregularly rounded or more frequently confluent into sublinear or curved masses, about 0.5 mm. thick, lenticular in cross-section; perithecia immersed, the over- lying stroma cracking away, exposing the white top of the nucleus, thus making a pseudo-ostiole about 250 in diameter; nucleus pyriform, about 250 р in diam- eter, 300 p tall; parathecium fusing with the stroma, entire, about 15 p thick below the hypothecium, the bark cells below blackened to a depth of about 65 А (the distinction visible in very thin sections) ; hypothecium 25 р thick, of gelified, predominantly periclinal hyphae; paraphyses slender, dichotomous, very closely packed in the nuclear gel and then appearing simple; asci 8-spored, about 65 X 13—14 p; ascospores imbricately monostichous, hyaline, finally brownish, broadly ellipsoidal, 4-locular, protoplasts lenticular, the apical ones conspicuously smaller, 19-21 X 8-9 p while still smoky gray, shrinking slightly when fully brown. 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 295 At first sight, this species might be mistaken for Bathelium porosum (Ach.) Dodge, from the West Indies, but it lacks a papilla at any stage and has smaller ascospores. Although the spores remain hyaline until late, they become brown when fully mature. MELANOTHECA cameroonensis Dodge, sp. nov. Type: Cameroons, Tombel, on Theobroma, C. A. Thorold 142. Thallus endophloeodes, obscure olivaceo-alutaceus, glaber, nigro-marginatus; pseudocortex 10 u crassitudine, cellulis suberosis ni gricantibus, decompositis, tenuibus cum hyphis dense contextis; filamenta Trentepobliae periclinalia, 5-6 p diametro inter cellulas suberosas penetrantia. Pseudostromata irregularia, lateribus abruptis, 250-270 p altitudine, nigra, ostiolis depressis, albis cum annulis, hyphis et cellulis suberosis decompositis; perithecia dimidiata, subsphaerica, 300 р diametro, 220 p altitudine; parathecium 40 p crassitudine, carbonaceum; hypothecium 10 p crassitudine, hyphis tenuibus, periclinalibus, dense intertextis; paraphyses tenuis- simae, dichotome ramosae; asci cylindrici, evanescentes; ascosporae octonae, im- ricatim monostichae, brunneae, 4-loculares, protoplastis rotundatis, 17-21 X 6—7 p. Thallus endophloeodal, deep olive buff, black-margined; pseudocortex about 10 р thick, of disintegrated, blackened cork cells with partly decomposed, densely woven, slender hyphae; periclinal filaments of Trentepoblia 5—6 р in diameter, penetrating deeply into the bark. Pseudostromata of disintegrated bark cells and hyphae, quite irregular in size and shape, black with abrupt sides, about 250-270 p tall, ostioles slightly depressed, surrounded by white rings; perithecia dimidiate, subspherical, 300 м in diameter, 220 р tall, immersed in the pseudostromata; para- thecium 40 p thick, carbonaceous, finally indistinguishable from the blackened pseudostroma in old perithecia which have lost their nuclei and which usually have a thin layer of blackened bark cells at their base and so appear entire; hypothecium about 10 р thick, of slender, densely woven, predominantly periclinal hyphae, nearly disappearing as the asci mature; paraphyses very slender, dichotomous in the nuclear gel; asci cylindric, 8-spored, wall early disappearing; ascospores im- bricately monostichous, brown, ellipsoid, 4-locular, protoplasts round, apical ones only slightly smaller, 17-21 Ж 6-7 р. This species differs from М. Acbariana Fée in having dimidiate, papillate peri- thecia with a white annulus and umbilicate ostiole and slightly longer and slenderer ascospores. MELANoTHECa obscurascens (Vainio) Dodge, comb. nov. Pyrenula (Melanotheca) obscurascens Vainio, Bol. Soc. Broteriana II, 6:35. 1929. Trypethelium anomalum v. obscurascens Zahlbr., Cat. Lich. Univ. 10:101. 1938. Melanotheca Acbariana v. obscurascens Zahlbr., Cat. Lich. Univ. 10:100. 1938. Type: Mozambique, Tungue Bay, Palma, 10? 5' S. lat., Americo Pires de Lima. Thallus fuscous olive; perithecia partly solitary or confluent into pseudo- Stromata; spores fuscous, 4-locular, 15 X 4-5 p [Vor. 40, 296 ANNALS OF THE MISSOURI BOTANICAL GARDEN The dark thallus, more solitary perithecia, and smaller ascospores, should sep- arate this species from all other members of the M. Achariana group. MELANOTHECA pusilla Dodge, comb. nov. Trypetbelium pusillum Jatta, Nuovo Giorn. Bot. Ital. 14:174. 1882. LAURERA LAURERA Reichb., Der Deutsche Botaniker seu Repert. Herbar. 15. 1841. Meissneria Fée, Suppl. Essai Crypt. Ecorc. Officin. 65. 1837, non DC. Meristos porum Mass., Atti I. К. Ist. Veneto III, 5:327. 1860. T belenella sect. Meristosporum Vainio, Etude Lich. Brésil 2:215. Batbelium Trev., Flora 44:21. 1861 p. p.; Müll. Arg., Bot. Jahrb. ripe 6:394. Type: M. varia Fée. Meristosporum Mass. was based on Trypetbelium meristo- sporum Mont. & v. d. Bosch. Batbelium Trev. was intended to be the same as Batbelium Ach., but included species with muriform spores, citing Meissneria varia Fée among others. Müller Argau treated Bathelium Ach. as a section of Try- betbelium and retained Batbelium Trev. (citing Meristosporum Mass. as a synonym) for the species with hyaline, muriform spores. Thallus crustose, epi- or endophloeodal, ecorticate, with Trentepoblia algae. Pseudostromata containing two or more perithecia with carbonaceous parathecia; paraphyses branched and anastomosing; asci 2—8-spored; ascospores muriform, hyaline, with more or less cubical cells. This genus is the stromatic analogue of Polyblastiopsis and Clatbroporina. Although widespread in the tropics, it has hitherto been reported only from Mozambique and Socotra Island, except for Trypethelium marginatum Fée from Senegal, Cap Vert, formerly considered a synonym of L. madreporiformis (Eschw.) Riddle. l. Ascoipotes under 40 м lo ЩЕ ри ME 2 1. Ascospores over 40 д long а оон е e е RE НИЕ 4 2. Stromata 1.5-2 mm. in diameter; — 25-38 X 10-15 и, 6—8-locul locellate; perithecia E e L. vela is (Mall. “е; dine 2. Stromata less th eter; ascospores less than 30 p long................————— 3. Ascospores вава бә туннин 25 X 4—10 ци; perithecia dimidiate; 8 ле а сы ыз аш, Lote uperrime (Mall, Arg) Zahlbr. 3 meats dioe 1—3-locellate, 30 X 11 u; Мога fec IU, oidella (Vainio) Zahlbr. Ascospores 12-20-locular, 40-75 Ж 12-15 д Peut dime D reddish brown with 2-5 оне each; — marginata (Fée) Doris Ascos Hine Мерари 2—2 a о 4: ес 8—10-1ос xp кеи 42-45 X 14-16 ш; pseudostromata immersed e bark, black; Nigeria L. — Dodge 5. ае dork uscous, nude, constricted at the base, 0.8 mm. in diamet spores: 63 М L. elegans (Vainio) Zahlbr. 5 ee substramineous to white, covered by thallus, not constricted at th iameter with 2—5 perithecia; ascospores 50-75 X 19—22 џ............-------------- т - I ИЕ: (Vainio) Zahlbr. LaunERA marginata Dodge, comb. nov. Trypetbelium marginatum Fée, Ann. Sci. Nat. 23:433. 1831. 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 297 LAURERA nigeriensis Dodge, sp. nov. Type: Nigeria, Moor plantation near Ibadan, on Theobroma, С. A. Thorold 165. Thallus epiphloeodes, roseo-alutaceus, 30 p crassitudine; cortex 10 и crassi- tudine, decompositus; stratum algarum 20 y crassitudine cellulis Trentepobliae et suberosis disintegratis. Pseudostromata 370 р altitudine in cortice arboris immersa, irregularia, ostiolis inconspicuis; parathecium 10—12 м crassitudine, integrum; nucleus pyriformis, 175 и diametro, 360 y altitudine, ostiolo 40 р diametro; hypo- thecium 16-18 y crassitudine, hyphis tenuibus dense contextum; paraphyses tenues, dichotome ramosae; asci clavati, 110 X 28 pu, evanescentes; ascosporae octonae, irregulariter distichae, hyalinae, murales, 8—10-loculares, 2—4-locellatae, protoplastis rotundatis, ellipsoideae vel subfusiformes, 42—45 Ж 14—16 p. allus epiphloeodal, cinnamon buff to pinkish buff, about 30 р thick; cortex about 10 р thick, the outer half decomposed, the inner half with structure ob- scured by dark granules; algal layer not sharply differentiated, about 20 м thick, of intermingled cells of Trentepoblia and disintegrating cork cells. Pseudostromata black, about 370 p thick, immersed in the bark, protruding about 30 р, nude, very irregular in shape and size, ostioles very inconspicuous; parathecium entire, 10—12 p thick at the base, fused with the pseudostroma above, later the underlying bark cells blackening to a depth of 50 и; nucleus pyriform, about 175 p in diameter, 360 p tall, ostiole about 40 p in diameter; hypothecium 16—18 p thick, of densely woven, slender hyphae; paraphyses slender, dichotomously branched; asci clavate, 8-spored, about 110 X 28 p, wall disappearing early; ascospores irregularly distichous, hyaline, muriform, 8—10-locular, 2—4-locellate, protoplasts slightly rounded, ellipsoid to subfusiform, 42—45 Ж 14—16 p. ASTROTHELIACEAE Thallus crustose, uniform, epi- or endophloeodal (epilithic in Lithothelium) ; cortex absent or poorly developed; algae Trentepoblia; perithecia pyriform with long necks, usually radially arranged, nearly free or more often immersed in a stroma or pseudostroma, the necks often confluent, opening into a common ostiolar canal, rarely remaining separate, each with its own ostiole; spermatia exobasidial. — . Ascospores. septate, 3—8-locular siiryi rira E ТТТ Ascospores muriform - 2. Ascospore protoplasts cylindric or neary so; preme блк кн aie EUM ое Аято Бейит Eschew. Pyrenastrum Eschew. ospores brown 1 Ascospores hyaline ian saley entnehmen Cryptothelium Mass. Deco EE .Parmentaria Fée ия — H PYRENASTRUM PvnENAsTRUM Eschw., Syst. Lich. 16. 1824. Type: P. septicolare Eschw. may be chosen as the суре, since P. plicatum described at the same time apparently has not been recognized since. SUN Thallus crustose, endo- or epiphloeodal, ecorticate or with a cartilaginous [Vor. 40, 298 ANNALS OF THE MISSOURI BOTANICAL GARDEN almost amorphous cortex; algae Trentepohlia. Регісһесіа innate in the bark or nearly so, usually radially arranged with long concrescent necks opening in a com- mon ostiole; parathecium entire; paraphyses branched and anastomosing; asci 4—8- spored; ascospores ellipsoid to fusiform, brown, 4—8-locular with rounded or lentiform protoplasts. 1. Perithecia spherical, 370 м in diameter; ascospores 16-27 X 8—14 м; Sierra Leone........ P. pruinosum Dodge 2 1, аураны ellipsoidal ospores 13-14 X 5-6 p; perithecia erect, 190 м in diameter, 240 p pe 1 Sere s Dodge 2: эн» e X 5-6 в, perithecia recumbent, 100 д in diameter, اجا‎ Sierra Leo P. ae м Dodge PYRENASTRUM pruinosum Dodge, sp. nov. Type: et Leone, Njala (Kori), on bark of Anisophyllea laurina, Е. C. Deighton M440 Thallus оа superficie olivaceo-alutacea, 250 p crassitudine, hyphis dense intertextis et cellulis Trentepohliae. Pseudostromata elongata, ca. 2 mm. latitudine, in reticulo concrescentia, thallo pallidiora et plus pruinosa, verrucis hemisphaericis ad 1 mm. diametro ostiolo circumdantibus; ectostroma carbonaceum in cortice arboris 600 p penetrans; endostroma pallidum, perithecia continens; perithecia subsphaerica, longis cum cervicibus, circum ostiolum commune ad 375 p diametro radiantia; parathecium 15 y crassitudine, carbonaceum, integrum; hypo- thecium 25 u crassitudine, hyphis tenuibus dense contextum; paraphyses tenues, dichotome ramosae anastomosantesque, apicibus liberis; asci cylindrici, ca. 80 12 p; ascosporae octonae, ellipsoideae, brunneae, 4-loculares, protoplastis rotund- atis, subaequalibus, 16-27 8-14 и. Thallus endophloeodal, surface deep olive buff to olive buff, completely dis- organizing the bark cells to a depth of 250 p, filling them with slender, densely tangled hyphae and including disorganized filaments of Trentepoblia, not in a definite layer. Pseudostromata elongate, about 2 mm. wide, concrescent into а network, paler than the thallus and more pruinose, with small hemispheric verrucae about the ostioles up to 1 mm. in diameter; ectostroma carbonaceous, extending about 600 р deep into the bark; endostroma not darkened, containing the peri- thecia, up to 375 и in diameter, subspherical with long necks, radially сеи about the common ostiole; parathecium entire, 15 p thick, carbonaceous; hypo- thecium 25 и thick, of slender, densely woven hyphae; paraphyses dichotomously branched and anastomosing, tips free; asci cylindric, 8-spored, about 80 X 12 в; ascospores ries brown, 4-locular with rounded, nearly equal protoplasts, 16—27 Х 8-14 PYRENASTRUM erumpens Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on bark of Phyllanthus discoideus, Е. С. Deighton M4397. Thallus epiphloeodes, alutaceus, ca. 80 p. crassitudine; cortex 25 p crassitudine, hyphis pachydermeis, periclinalibus, gelifactus; stratum algarum 55 ۾‎ crassitudine, 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 299 cellulis cylindricis Trentepohliae, 8 р diametro; medulla non evoluta. Stromata erumpentia, carbonacea, orbicularia, 2 mm. diametro vel elongata, 1.5 X 4 mm., vel irregularia, parva, duobus cum ostiolis depressis, majora cum 15—20 ostiolis; ectostroma 100 y crassitudine ab endostromate faciliter separans; perithecia ellip- soidea, 190 p diametro, 240 y altitudine, cervicibus excentricis brevibus in cervice communi aperientibus, 60 р diametro ad ostiolum, inferne latiore, 150 p altitudine; parathecium ab endostromate carbonaceo non distinctum; hypothecium 15 y crassi- tudine, hyphis tenuibus dense contextum; paraphyses parce ramosae, apicibus liberis; ascosporae octonae, imbricatim monostichae, brunneae, 4-loculares, proto- plastis rotundatis, subaequalibus, 13—14 Ж 5-6 p. Thallus epiphloeodal, deep colonial buff, about 80 p thick; cortex 25 р thick, of densely woven, mostly periclinal hyphae, highly gelified; algal layer 55 р thick, of disoriented, subcylindric cells of Trentepohlia, 8 р in diameter; medulla not differentiated. Stromata erumpent, carbonaceous, circular, about 2 mm. in diam- eter, or elongate, about 1.5 Ж 4 mm., or quite irregular, the smaller ones with only 2 depressed ostioles, the larger with 15—20 ostioles; the ectostroma about 100 p thick, tending to crack away from the equally carbonaceous endostroma in sectioning; perithecia ellipsoidal, about 190 p in diameter, 240 р tall, neck excentric, rather short, opening into the common neck of the group, about 60 p in diameter at the ostiole, wider below, about 150 д tall; parathecium not distinct from the endo- stroma; hypothecium about 15 p thick, of slender, densely woven hyphae; para- physes little-branched, tips free; asci 8-spored; ascospores imbricately monostichous, brown, 4-locular, protoplasts rounded, subequal, 13—14 X 5-6 р. PYRENASTRUM parathelioides Dodge, sp. nov. Type: Nigeria, Moor plantation near Ibadan, on Theobroma, С. А. Thorold 169. Thallus epiphloeodes, olivaceo-alutaceus, 65 p crassitudine; cortex 55 p crassi- tudine, gelifactus, hyphis tenuissimis periclinalibus; stratum algarum 10 p crassi- tudine, cellulis Trentepobliae compactis; medulla non evoluta. Stromata 0.6—0.7 mm. diametro, 0.2 mm. altitudine, carbonacea, lenticularia, thallo tecta, senectute dimidia parte supera frangente ut in speciebus Pyrenulae; perithecia solitaria vel geminatim disposita, cervicibus brevibus in ostiolo communi aperientibus, ellipsoidea, 100 д diametro, 180 р longitudine; parathecium integrum, 12-15 y crassitudine, carbonaceum; hypothecium 20 p crassitudine, hyphis tenuibus dense contextum; asci cylindrici, 85 Х 12 р; ascosporae octonae, ellipsoideae, brunneae, 4-loculares, protoplastis rotundatis, apicalibus minoribus, 17-19 Х 5—6 p. Thallus epiphloeodal, olive buff, about 65 р thick; cortex 55 p thick, gelified, of very slender periclinal hyphae; algal layer compact, 10 p thick, of Trentepohlia cells, which occasionally penetrate deeply into the bark; medulla not differentiated but hyphae penetrate deeply and disorganize the bark cells. Stromata 0.6—0.7 mm. in diamater, 0.2 mm. tall, lenticular, carbonaceous, covered by the thallus which finally cracks off along with the upper half of the stroma, exposing cup-shaped depressions as seen in some species of Pyrenula; perithecia solitary or mostly in [Vor. 40, 300 ANNALS OF THE MISSOURI BOTANICAL GARDEN pairs, necks short, emptying into a common ostiole, ellipsoidal with the long axis parallel to the substrate, up to 100 p in diameter and 180 p long; parathecium carbonaceous, entire, usually concrescent with the stroma, rarely cracking apart; hypothecium 20 y thick, of densely woven, slender hyphae; asci cylindric, 8-spored, about 85 X 12 р; ascospores ellipsoid, brown, 4-locular, protoplasts rounded, the apical ones finally much smaller than the middle ones, 17-19 Ж 5-6 Sections of a stroma with a solitary perithecium closely Pipini those of Parathelium with a short, lateral neck, whence the name. Old material, after the upper portion of the stroma has cracked off, might be mistaken for an old thallus of Pyrenula. CYPHELIACEAE Thallus crustose, uniform or effigurate, ecorticate, with protococcoid or Trentepoblia algae; mazedia sessile or immersed in subcylindric thalline warts, with or without a parathecium. 1. Thallus with protococcoid algae dud 1; NM en EM PUR ; bu 2 sn Hrem ich 3. Mise Кемен ascospores hyalin arriola 3. Mazedia lecanorine; ascospores finally brown -Carlosia mp. "à Asci polysporous ERE HETE ив 4 As Френ i 0 а са Е хл . Mazedium with only parathecium, ascospores 2—4-locular. .Pyrgillus Nyl. . Mazedium with both РГ еси. and amphithecium; ascospores mostly 2-locular........ T»ylopboron Nyl. Mazedium with both parathecium and amphithecium; ascospores 4-locular, 2 2-locellate Schistophoron Stirton v мл h TYLOPHORON TYLOPHORON Nyl., Bot. Zeit. 20:279. 1862. Type: not designated, based on T. protrudens Nyl. and T. moderatum Nyl. Thallus crustose or disappearing; with Trentepohlia algae. Mazedium im- mersed in a subspherical thalline wart at first, then sessile, subcylindric with an open disc; both parathecium and ampithecium present; hypothecium hyaline or brownish; asci cylindric, 8-spored; ascospores 2-(rarely 3-)locular, subspherical to ellipsoid or subfusiform with a thick epispore; spermogonia immersed in the thallus with a hyaline wall; spermatiophores cylindric, slightly branched; spermatia acicular, straight. Ascospores 8 Х 4 u; mazedia 0.6-1 mm. in diameter, 0.2—0.4 mm. tall; quem moe qx. ascidioides Vainio Ascospores 9—11 X 5-7 p; mazedia 0.2-0.3 mm. in diameter; a А E E eG id moderatum v. modestius Zahlbr. TYLOPHORON AsCIDIOIDES Vainio, Bol. Soc. Broter. II, 6:173. 1929. Type: Mozambique, Ponta Vermelha, lignicole, Pires de Lima 25. Thallus epiphloeodal, white (slightly greenish when moist), homoeomerouss 40—50 p thick, of disorganized filaments of Trentepoblia, with very slender hyaline hyphae, a few Ке of Trentepoblia penetrating between layers of bark cells. 1953] DODGE—LICHENS OF TROPICAL AFRICA 301 Mazedium cylindric ог constricted at the base, about 0.5 mm. in diameter, 0.7 mm. tall, disc urceolate at first, becoming slightly convex at maturity; amphithecium about 180 д thick, of slender, densely woven hyphae containing large crystals, but no algal cells seen; parathecium slightly developed; hypothecium of slender vertical hyphae, slightly brownish, 250 p tall; thecium 120 y tall; asci cylindric to clavate, 8-spored; paraphyses slender, forming the capillitium; ascospores monostichous, brown, nearly cylindric, 2-locular, sometimes slightly constricted at the septum, wall relatively thick, 8-11 X 4 y. Our specimen has smaller mazedia and slightly longer spores than in the type, but we have preferred to refer it here in the absence of more specimens to show the amount of variability to be expected in the genus. Nigeria: Ondo Province, Owena near Akure, on Theobroma, C. A. Thorold 168. CRYPTOTHECIACEAE Thallus crustose, epiphloeodal, of very loosely woven hyphae, a true cortex not developed, but the algal layer covered by a layer of hyphae, somewhat more densely woven than those of the medulla; algae Trentepohlia; asci 1—8-spored, borne singly throughout the thallus usually in the medulla, rarely in the algal layer, bitunicate and pushing up to the surface for spore discharge; ascospores septate to muriform. The thallus resembles that of a sterile Crocynia or of Chiodecton subg. Bysso- phorum, and it is possible that some species referred to these genera may be found to belong in this family. In some species, the brownish tips of the resting asci may be seen at the surface under relatively high magnifications. Santesson (Foliicolous Lichens I. Symb. Bot. Upsal 12:1:57-68. 1952) transferred the family to the Arthoniaceae and included a group of species from Arthonia and Arthothelium in Stirtonia and Cryptothecia respectively. In this group, the asci are borne in groups in a sort of primitive (?) ardella, without clearly developed paraphyses, often showing a disc at the surface of a slightly different color (fertile areas) from that of the rest of the thallus. While we can- not be certain until we have more data of their development, it is probable that they should be segregated as different genera. For completeness I have included species of this group in the keys. Miss A. L. Smith mentions a thin, hyaline or. slightly brownish peridium about each ascus, but I have found no such differ- entiation in the numerous specimens I have studied. Apparently, the expanding ascus pushes aside the medullary hyphae, so that they are more compact about the asci. The *paraphyses" of some descriptions аге only the medullary hyphae between the asci, and are neither true paraphyses nor paraphysoids. .........Sfirtoniopsis Groenh. Ascospores bilocular, small iain ere iiie ere р Ascospores septate, 8—14-locular 7 Vot uM onm е A edis Ascospores muriform EAEE E AEA ST AKEE a parT ry [Vor. 40, 302 ANNALS OF THE MISSOURI BOTANICAL GARDEN CRYPTOTHECIA CrYPTOTHECIA Stirton, Proc. Phil. Soc. Glasgow 10:164. 1876. Myriostigma Krmplh., Lich. Foliic. quos legit О. Beccari annis 1866—1867 in insula Borneo, 22. 1874; Nuovo Giorn. Bot. Ital. 7:44. 1875. Myxotheca Ferd. & Winge, Bot. Tidsskr. 30:212. 1910. Type: C. submidulans Stirton. Myriostigma was based on M. candidum Krmplh. Although Myriostigma antedates Cryptothecia, Santesson has proposed the latter as а nomen conservandum. Myxotheca was based оп М. bypocreoides Ferd. & Winge. Thallus spreading, smooth or minutely furfuraceous, whitish to pale glaucous, margins sometimes fimbriate; cortex scarcely differentiated, intricate, slightly more compact than the medulla; algal layer of Trentepohlia filaments, more or less dis- organized; medulla relatively thick, loosely woven, of slender hyphae; asci widely scattered throughout the medulla, rarely in the algal layer, pyriform to subspherical, thick-walled when young, 1—8-spored; ascospores hyaline, muriform, ellipsoid to subspherical. 1. Ascospores small, 20-38 X 8-14 р, asci 4-5рогеф eee LLL eec coco eec наннан" 2 Ascospores much larger 2. Thallus lacerate in long, irregular lobes, pseudo-ardella with a pale flesh-colored disc; n. ascospores 20-38 X 8—14 р; foliicole; Angola........ Arthothelium laceratum (Vain.) Zahlbr. 3. Thallus continuous; asci very scattered, no disc present; ascospores 28 X 14 №; co eria C i H icole; Nig nigeriensis Dodge 3. Asci monosporous, rarely 2-spored "uq d cS Rod а iab eec te ae ee 4 3. Asci 8-spored NO UOCE A eA Mer 3 4. Ascospores 50—75 Х 20-32 u; Cameroons C. subnidulans Stirton 4. Ascospores 40 X 35 и; Nigeria C. Thoroldi Dodge . Ascospores 20-locular, 4—5-locellate, 70-125 X 25-30 p, sharply bent; Usambara........ ^ C. geniflexum (Müll. Arg.) Sant. 5. Ascospores 12—14-locular, 4—6-locellate, 70-95 X 25-35 u, curved but not sharply bent; Usambara Arthothelium dictyophorum Mill. Arg. Ascospores about 10-locular, 4—6-locellate I E 6. Ascospores 70-100 X 25-50 и; Cameroons С. Stirtoni A. L. Smith 6. Ascospores 60—80 X 25-35 д; Usambara C. caesioalba (Müll. Arg.) Sant. мл мл . CRYPTOTHECIA nigeriensis Dodge, sp. nov. Type: Nigeria, Ina near Ibadan, on Theobroma, C. A. Thorold 151a. Thallus subarachnoideus, pallide griseus, epiphloeodes, 50—200 p crassitudine, superficie inaequali; ecorticatus; stratum algarum 25—40 y crassitudine, mentis verticalibus T'rentepobliae, cellulis 5—6 р diametro, cylindricis vel subsphaericis; medulla hyphis tenuibus, 1 и diametro, granulis inspersis, laxe intertextis; asci solitarii in medulla sparsi, subsphaerici, 40 р diametro, pachydermei; ascosporae quaternae, hyalinae, ellipsoideae, murales, ca. 28 X 14 p. Thallus subarachnoid, light mineral gray, epiphloeodal, 50-200 p thick, surface irregular; ecorticate; algal layer 25—40 p, of more or less vertical filaments of Trentepoblia 5-6 и in diameter, cells cylindric to subspheric; medulla of loosely woven hyphae about 1 y in diameter, more or less inspersed with minute granules; asci borne singly in the medulla, subspherical, about 40 p. in diameter, very thick- walled, 4-spored; ascospores hyaline, ellipsoidal, muriform, about 28 X 14 p 1953] DODGE—LICHENS OF TROPICAL AFRICA 303 CRYPTOTHECIA Thoroldi Dodge, sp. nov. Type: Nigeria, Ondo Province, Owena near Akure, on Theobroma, C. А. Thorold 163. Thallus epiphloeodes, 135 p crassitudine, byssoideus, pallide griseus, margine albo, byssoideo; pseudocortex 25 ш crassitudine, hyphis erectis 2-3 р diametro, laxe intertextis; stratum algarum 15—25 y crassitudine, filamentis Trentepobliae, plus minusve periclinalibus, 4 и diametro; medulla 85 p crassitudine, hyphis tenui- bus laxe intertextis; asci solitarii, in strato algarum sparsi, pyriformes vel sub- sphaerici, 43 X 27-35 y, juventute pachydermei, monospori; ascosporae hyalinae, muriformes, 40 X 26-34 p. Thallus epiphloeodal, 135 р thick, byssoid, spongy, light mineral gray with a white byssoid margin; pseudocortex 25 р thick, of erect hyphae 2-3 y in diameter, forming a velvety surface; algal layer 15-25 p thick, of more or less periclinal filaments of Trentepoblia about 4 p in diameter; medulla 85 р thick, of very slender, loosely woven, hyaline hyphae; asci monosporous, borne singly in the algal layer, pyriform to subspheric, 43 X 27-35 р, thick-walled when young; ascospores hyaline, muriform, 40 X 26-34 р. . | ARTHONIACEAE Thallus crustose, uniform, homeo- or heteromerous, epi- or endophloeodal, ecorticate, with Palmella, Trentepoblia or Phyllactidium algae; ardella without parathecium, variable in shape from rounded, elongate, or radially branched, usually single (in a stroma in Зума" Рота); paraphyses branched and anastomosing to form a thick epithecium; asci subspheric to broadly pyriform; ascospores septate or muriform. 1. : 5 ; becoming brown; algae Trentepoblia Ardellae immersed in a stroma; ascospores bec 5 ga Synarthonia Müll. Arg Rr АУР meme crean pen E ти ИМИ 2 1. Ае Tusltbill и : 3 CAU Pei 5 И tice e TT 4 2. Algae Pbyllactidium; foliicole .... mmm ^ == 3. ospores septate ен tiet e AA te 2 3. res О рЫ ЯНЕШЕ SRO роз Nyl. ......................... мл М P. eec Rs VS DENM НИЕ ЫНАН нн a аанча ает о „АІ Artboniopsis Müll. Arg. і Trichophyma Rehm мл D ARTHONIA ARTHONIA Ach., Neues Jour. f. d. Bot. 1:3:3. 1806. Ustalia Fr., Syst. Orb. Veg. 274. 1825. : Type: not designated; eight species included in the original treatment. The foliose species are now placed in ja. Of the six remaining, all belong in Artbonia as now recognized. Opegrapha radiata Pers. may be selected as the type. Ustalia Fr. was based on Graphis caribaea Ach. For а discussion of the numerous [Vor. 40, 304 ANNALS OF THE MISSOURI BOTANICAL GARDEN other possible synonyms and their types, see Santesson, Foliicolous Lichens I. Symb. Bot. Upsal. 12:1:68—75. А Thallus crustose, uniform or Ms epi- or endophloeodal, or saxicolous; ecorticate or with a pseudocortex of disintegrated bark cells; algae Trentepoblia; medulla often scarcely developed; ardellae rounded, elongate or star-shaped, more or less immersed in the thallus; parathecium absent ог the outermost paraphyses - blackened forming a pseudoparathecium; paraphyses branched and anastomosing above, forming a thick epithecium; asci subspheric, pyriform, rarely ellipsoid, thick-walled when young, especially above, 8-spored; ascospores oblong-ellipsoid to clavate when the apical cell is much larger than the others, septate, 2— pluri- locular, hyaline or brownish; spermogonia superficial, wall dark, spermatiophores subcylindric; spermatia cylindric, straight or curved. Stylospores sometimes present, terminal, ovoid to ellipsoid, unicellular or septate, hyaline or brownish. 1. Ardella white or very pale 2 1. Ardella bright red to s 3 1. Ardella dark fuscous to 7 lac isc white, more or less confluent like a minute pertusarial stroma —— in Stirtonia, fide Sant.) ; ascospores 8—10-locular, 50-60 X 20-22 ш; Usambara........ А. pertus НЫЕ Mill. Arg. $ — fee flesh-color, circular or angular Sten in Stirtonia, т Sant.); asco- AN 38-46 Х 10-16 ш; Usam carneoalbens Müll. Arg. 2 Dic. estote: white-pruinose, pale brown ve che PM 0.2 mm. =>. diameter; ascospores 4—6-locular, 18—22 X 5.5—6 ш;.Ѕіегга Leon su gat Dodge 2. Disc yellowish me lobulsta or angular; ascospores аи 17-20 T 3-5 EIC A A d Mall. 22 Disc = ascospores а 15-20 X 6-7 и; Mozambique Disc cinnabar to viole Disc ow AD ascospores 6-locular, 30—32 X 10 u; Mozambique ا . w . Е НА A. lept — arr: Zahlbr. 4 Thallus white SS s arpa Vainio 4. еи Pale rose, or rose spotted on white A throcar а is TOR гора ы Vainio Ardella round, disc caesio- o-prudaoes, mi margin black; Mozambi а: rhe dus nw Nyl. Ardella chicas: disc red-pruinose, margin black; со 17-22 X 5. sae E NAE CC perme pobiti у. rimata (ушы Zahlbr. Ardella long and narrow, disc blackish, margin rd “ascospores a ES и сс: nnabarina v. EE (Vainio) -— ww *- мл he L RUN camels Een nate Seen te em ee ee ee TT ose, then and nmn Кебу А. cinnabarina v. es zii. Я рис cinnabar purple; 1 net 0 X 8 u; Usambara........A. cinnabarina v. speciosa M 6. Disc fuscous, pruinose; recense THE EF Angels s eee v ер of РЕ Ке ‘Ss 3t gs PE چ‎ © T i a иза an area 4—5 mm., very dendroid-brancheds ascospores d ccc nabarina v. dendritica "е u X 5—6 и; ardella elliptic, 0.2 mm. wide, disc red en moist, black when s not pruinose; Somaliland........... „А. somaliensis Ml. =: Ha Hem. E: "A fuscous; nhi dendepid-branched, branch ong, disc dark violet when d pruinose; sp to 3 mn. long, dic dark viole when dry, noe к азаа v. dendritica Steiner Ascospores 5—6-locular г Ascospores 7—12-locula 8. Ardella round, 0.4—9. 4-loc pa Ardi 8. Aul mM k К 0.3—0.5 0.1—0 ound, 0 eter; ascospores 12—16 X 3—45 џ, ries ы г; Mozambic En ا ا مهه جه‎ аа т - А. palmensis (Vainio) Zahlbr. чч „ры: EE 3 à ч > о 9 3 8 PER Pg 8. & ^ È === === мм еее еее еен "Pe e ea ea CT тавобеи, Re “10 5 (-0.7) mm. in diameter; ascospores 17-20 Х би, with halo, ar; eus ER SU IERE S er ЫП modeste 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 305 Ascospores 18—20 X 4-5 п; ardella 0.5 X 1 mm., disc black; Sierra Leone........ A. elevata Dodge Ascospores 28 X 11-12 ш; ardella 0.5-0.8 X 0.12—0.15 mm., disc fuscous black; Angola A. leptogramma Müll. Arg. 9. Ascospores 30—32 X 10 u; ardella 0.2-0.5 X 0.1—0.15 mm., disc rufescent; Mozambique 1 u А. leptographoidea (Vainio) Zahlbr. 10. Ascospores 22—30 X 8-9.5 u; ardella 0.5 X 0.25 mm.; Kenya A. ilicinodes Steiner 10. Ascospores 24 X 8-9 u; ardella 0.5-1 X 0.1-0.5 mm.; Mozambique........................ A. microcarpella (Vainio) Zahlbr. 10. Ascospores 24—27 X 10.5-13.5 и, 8-locular; ardella round, 0.5 mm. in diameter; Sierra Leone A. leptogrammodes Dodge 10. Ascospores 50—60 X 18-20 и, 10—12-locular; ardella round, 0.5-1 mm. in diam- eter; Socotra A. calospora Müll. Arg. ARTHONIA modesta Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on bark of Peltophorum africanum у. speciosum, Е. С. Deighton M4342. Thallus endophloeodes, superficie fumoso-griseus; pseudocortex ad 25 y crassi- tudine, cellulis suberosis decompositis et hyphis tenuibus hyalinis; stratum algarum ad 20 y crassitudine, filamentis periclinalibus Trentepobliae inter cellulas suberosas decompositas. Ardella orbicularis, modice convexa, 0.4-0.5 (—0.7) mm. diametro, disco nigro; hypothecium non bene evolutum; thecium 80 y altitudine; paraphyses tenues, superne ramosae anastomosantesque, epithecium brunneum, ca. 20 y crassi- tudine super ascos formantes; asci ellipsoidei vel subpyriformes, 30 X 20 р, apicibus incrassatis; ascosporae octonae, hyalinae, halone circumdatae, (3-) 4-loculares, late clavatae, cellula apicali majore, 17—20 X 6 p. allus pale smoke gray to smoke gray; endophloeodal; pseudocortex up to 25 м thick, of disintegrated bark cells and very slender, hyaline hyphae; algal layer up to 20 д thick, of periclinal filaments of Trentepohlia in and between disinte- grated bark cells, with a few medullary hyphae penetrating more deeply into the bark. Ardella circular, 0.4-0.5 (-0.7) mm. in diameter, moderately convex, disc black; hypothecium not differentiated, the asci and paraphyses appearing to arise from disintegrated bark cells; thecium 80 р tall in the center, slightly lower toward the margin; paraphyses, slender, branched and anastomosing above the asci, form- ing a brownish epithecium about 20 p thick; asci ellipsoid to subpyriform, 8- spored, about 30 X 20 р, tip thickened, protoplast broadly mamillate; ascospores hyaline, broadly clavate, with a thin halo, 3—4-locular, terminal cell somewhat larger, 17-20 X 6 p. ARTHONIA elevata Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on bark of Anisophyllea laurina, F. С Deighton М4406. И Thallus endophloeodes, albidus, pruinosus; pseudocortex 25 р crassitudine, cellülis suberosis decompositis et hyphis hyalinis tenuissimis; stratum algarum 50 م‎ crassitudine, filamentis verticalibus Trentepohliae et cellulis suberosis decompositis. Ardella ad 1 mm. diametro, orbicularis vel oblonga (dein 0.5 Х 1 mm.), disco nigro, pseudolecideina; pseudoparathecium paraphysibus nigricantibus; thecium 65 м altitudine; hypothecium 10 p crassitudine hyphis periclinalibus pallide brunneis; asci pyriformes vel subsphaerici, pachydermei, 32 X 25 p; ascosporae [Vor. 40, 306 ANNALS OF THE MISSOURI BOTANICAL GARDEN octonae, fasciculatim dispositae, clavatae, hyalinae, 5—6-loculares, 18—20 X 5-6 р. Thallus endophloeodal, nearly white, pruinose, subrimose on very rimose bark; pseudocortex about 25 p thick, of disintegrated bark cells and very slender hyphae; algal layer about 50 и thick, of short, vertical, partly disorganized filaments of Trentepoblia and some disintegrated bark cells; medulla not differentiated but medullar hyphae penetrating some distance into the bark and disorganizing the cells. Ardella round, up to 1 mm. in diameter or oblong, then about 0.5 X 1 mm., disc black, convex, seated on bark cells less disintegrated than in the surrounding thallus, thus appearing elevated and lecideoid (rarely lecanoroid); pseudopara- thecium of blackened paraphyses, progressively paler within; thecium 65 y tall in the center, somewhat lower toward the margin with younger asci; hypothecium 10 р thick, of pale brownish periclinal hyphae; asci pyriform to subspheric, 8- spored, 32 X 25 u when nearly mature, thick-walled; ascospores fascicled, clavate, hyaline, 5—6-locular, 18—20 X 5-6 y. ARTHONIA leptogrammodes Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on twigs of Lagerstroemia speciosa, F. C. Deighton M4347. Thallus epiphloeodes, albidus, tenuissimus, tenuissime nigromarginatus; ecorti- catus; filamentis T'rentepobliae, 5-6 р diametro, hyphisque tenuissimis, crystallis minutis nubilatis. Ardella orbicularis, 0.5 mm. diametro, rare 2—3-confluentia, disco nigro, immarginata; hypothecium hyalinum, tenue; thecium 55 y altitudine; paraphyses tenues, ramosae anastomosantesque, superne brunneae; asci pyriformes, 35 X 25 p, juventute pachydermei protoplastis mamillatis, dein leptodermei; asco- sporae octonae, hyalinae, polystichae, halone tenui circumdatae, ad 8-loculares, cellula apicali 9 п longitudine, basali 6 p, intermedii 2 и longitudine, 24—27 x 10.5—13.5 р. Thallus epiphoeodal, white, very thin, with a narrow black margin; algae Trentepohlia, filaments 5-6 p in diameter, partly disorganized and surrounded by very slender hyphae covered with minute crystals. Ardella nearly circular, 0.5 mm. in diameter, rarely 2—3 confluent, disc black, immarginate; hypothecium very thin and hyaline; thecium 55 р tall; paraphyses slender, branched and anastomosing, slightly brownish above; asci pyriform, 8-spored, thick-walled when young, espe- cially above, protoplast mamillate, becoming quite thin-walled at maturity; asco- spores hyaline (a few slightly smoky as if finally becoming brown) with a thin halo, up to 8-locular, the apical cell 9 џ, the basal cell 6 р, the intermediate cells about 2 p long, 24—27 X 10.5—13.5 и. Perhaps closest to A. leptogramma Müll. Arg. in size and shape of the asco- spores, although both terminal cells are much larger than the middle cells, which is very unusual in the whole genus Arthonia. ‘The ardella is completely different in size and shape. 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 307 ARTHONIA (PACHNOLEPIA) subcaesia Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on twigs of Bauhinia tomentosa, Е. C. Deighton M4307H. Thallus epiphloeodes, tenuissimus, albidus, tenuiter nigro-marginatus, cellulis cylindricis Trentepobliae et hyphis tenuissimis, superne crystallis minutis nubilatis. Ardella orbicularis vel suboblonga, erumpens, disco dense caesio-pruinoso, 0.2 mm. diametro; hypothecium 50 д crassitudine, pallide brunneum, hyphis tenuibus cum crystallis; thecium 50 и altitudine; paraphyses tenues, ramosae anastomosantesque; asci pyriformes, 35—40 X 18—20 р, juventute pachydermei; ascosporae octonae, polystichae, clavatae, hyalinae, 4—6-loculares, 18—22 X 5.5-6 p. Thallus epiphloeodal, very thin, white to pearl gray with a narrow black mar- gin, nearly homoeomerous; algal cells of Trentepohlia, cylindric, from disorganized filaments, surrounded by very slender, tangled hyphae, the outer portions covered by minute crystals. Ardella round to slightly oblong, erumpent, disc densely white-pruinose (pale brownish when the pruina is rubbed off), about 0.2 mm. in diameter; parathecium not differentiated; hypothecium about 50 p thick, pale brownish, of slender hyphae covered with crystals; thecium 50 p tall; paraphyses slender, branched and anastomosing to form the epithecium above the asci; asci 8-spored, pyriform, 35—40 Ж 18—20 р, very thick-walled, especially above, when young; ascospores polystichous, clavate, hyaline, 4—6-locular, 18—22 X 5.5-6 p. OPEGRAPHACEAE Thallus crustose, cortex poorly developed or absent, algae Palmella (Xylo- grapheae), Trentepohlia (Opegrapheae), ог Phyllactidium (Micrographeae) ; medulla poorly developed or absent. Lirellae simple or branched, usually sessile, rarely immersed, single (with two parallel thecia per lirella in Ptychographa and Diplogramma) ; parathecium usually thick, black, and carbonaceous (rudimentary in Gymnograpba) ; ascus more or less clavate with 8 or less ascospores; ascospores unicellular, septate, or muriform, walls and septa thin, protoplasts cylindric in septate spores, appearing cubical in muriform spores. 1. Algae Palmella (Xylographeae) ......————————7 2 1. Algae Trentepoblia, ascospores septate ог muriform (Оревгарһеае)...........—-------------6 1. Phyllactidium (Micrographeae) ==" i ч ja res urnicelluldf LL ЕА а aee rin ud со5рогез e dE fi E A миноре 2. As а етуй Я дона .Aulaxina Fée 3. Lirellae with a single thecium each ......——— ba NI 3. Lirellae with two parallel thecia each ..............——————————870000887 hographa Nyl. 4. Hypothecium pale; on w rarely on bark... Xy rapba Fr. 4. Hypotheci dark; saxicolous .............. nnt Lithographa Nyl. 5. Ascospores bilocular, lirellae caespitose иене alographa М 5. Ascospores 4-locular, 2 parallel thecia per lirella........--------- scs Diplogramma Müll. Arg. г oe bs E М БЕЛИНЕН ЕНЕ oen ы ae * соѕрогеѕ а а a ыы 7 Арна septate * uou qe ure ре Омь Pers. 7. Азсозрогез muriform ee ee Н.Г. Fée 8. Lirellae immersed; parathecium rudimentary...———77777 Gymnograpba Müll. Arg 8. Lirellae sessile; parathecium well developed......—————————- $6 br 9. Ascospores hyaline с М Тош. 9. Ascospores brown и ecco Micrograpbba Müll. Arg. [Vor. 40, 308 ANNALS OF THE MISSOURI BOTANICAL GARDEN OPEGRAPHA OPEGRAPHA Pers., Neue Ann. Bot. [Usteri] 1:23, 29-32. 1794, non Humb., Fl. Friberg. Specim. 57. 1793 Alyxoria S. Gray, Nat. Arr. Brit. Pl. 1:101. 1821. Hysterina S. Gray, Nat. Arr. Brit. Pl. 1:504. 1821. Scapbis Eschw., Syst. Lich. 14. 1824. Oxystoma Eschw., Syst. Lich. 14. 1824. Zwackhia Koerb., Syst. Lich. Germ. 284. 1854. Xylastra Mass., Framm. Lich. 10. 1855. Type: Opegrapha Humb. was based on Lichen scriptus L. and its varieties, which species has been the type of Graphis since that genus was founded. How- ever, Opegrapha Pers. 1794 has been universally recognized in the present sense since its publication and being based wholly on species now included in the genus, it should be conserved against Opegrapba Humb. 1793; otherwise all the species now in Graphis would have to be transferred to Opegrapha and all those now in Opegrapha would have to be transferred to Alyxoria or Hysterina. In his generic description, Persoon cites Lichen scriptus var. a. L. rugosus L., Hoffm., which later in the discussion of the species he cites as a synonym of his O. faginea. This entity is now considered a fungus; hence in selecting a type from the other 13 species, it would be better to select another species, such as O. licbenoides Pers. to conserve the genus in its present sense. Alyxoria Ach. in S. Gray was based on Opegrapha notba Ach. and О. diaphora Ach. Hysterina S. Gray was based on 12 species still included in Opegrapha. Scapbis Eschw. was based on О. notha Ach. (figured) and on $. anfractuosa Eschw. and $. compressa Eschw. Oxystoma Eschw. was based on О. cylindrica Raddi. Zwackhia Koerb. was based on Z. involuta (Wallr.) Koerb. Xylastra Mass. was based on Arthonia fuscescens Fée. Thallus crustose, uniform, ecorticate or nearly so, either saxicole or corticole; algae Trentepohlia; lirellae immersed to sessile, lips somewhat connivent, nearly closed to open, especially when wet; parathecium entire or dimidiate (in sect. Pleurothecium) ; paraphyses branched and anastomosing above the asci; asci clavate, 8-spored; ascospores ellipsoid to fusiform, straight or slightly curved, hyaline, septate, 2-18-locular, protoplasts cylindric; spermatia exobasidial, filiform, straight or curved. Stylospores sometimes present, terminal, ovoid to long-ellipsoid, straight or slightly curved, hyaline. In the following key, the species are corticole unless otherwise stated: Га" . Perithecium entire (EUOPEGRAPHA 1. Perithecium dimidiate (PLEUROTHECIUM) 2. Ascospores 4-locular 2. cospores 4—9-locular a A 10—14-locular —— a cp О { alus and disc ochraceo-pruinose; lirellae 0.3-2 X 0.25—0.3 mm., simple or rarely orked; ascospores 14 X 4 p, without halo; Angola... O. Vainioi Zahlbr. ( L о 3. Thallus and disc not w 1953] DODGE—LICHENS OF TROPICAL AFRICA 309 с ced more than 18 д long; thallus white 5 cospores less than 18 д long 5. iuda seule 0.3-0.5 Х 0.2 mm.; پرا‎ 35—40 u long; São Thomé......0. subnotbella NL 5. Lirellae semi-immersed, 0.2—0.8 X 0.2—0.3 mm.; ascospores 18—23 X 79 м; Angola albocinerea Vainio 7 6. Thallus white 6. Thallus re Lis d eiis X 4—5 p, with halo; pu 1.5—1.7 0.2—0 4 n Zambesi enybartii Mal Arg. 6. oos ‘pale у sl. prunis ascospores 16 X 6 pu; wg 15-25 3$ * uU К, Mill Arg. 5 Te а white; saxicole; ascospores 16—18 X 4 п; Senegal О. alboatra Nyl. 7. Thallus whitish-granulate; corticole; ascospores 14-15 X 4.5 и; São Thomé........ О. lepidelle Nyl. 7. Thallus white, i: corticole; ascospores 13-15 X 3-3,5 А И Island O. aterula Müll. E 8. Ascospores more than 27 yu un thallus ashy to glaucescent............. нения 8. Ascospores less than 27 p lon; ie 9. Ascospores 26—30 X 6—6.5 iut v resa cells subequal, no halo; thallus ashy; lirellae 0.2-0.5 Х 0.1 mm.; Ken О. pees Steiner 9. астар: 30-34 X 4-5 p, eder cells subequal; lirellae angular ог renifor: alcicole; Socotra О. cretacea Mal. Arg. 9, м 32-45 X, 5—6 u, with halo, 6—8-locular, one or two А. cells much longe than the others; i w pale glaucescent; lirellae 0.3-1 X 0.1 ; Angol seplemseptat Vainio Азсозрогез 27.5—40 X 4—5 iw piden en 7—9-locular; thallus ca to o glavcecen lirellae 0 Х 0.1 mm . dei Ties үнө 12 ashy 10. Thallus pale наь or glauc 10. Thallus V Toa asc n 16-21 X 5-6 и, 4—8-locular; lote D ight; Ilha cipe leptogrephe Nyl. ll. Ascospores "- x45 6-locular; auem re lirellae rounded, diameter, or elongate, 0. 8-15 X 0 base of parathecium 7 hinner agis the sides; Usambar rufa Müll. Arg. ll. Ascospores 20—25 М 4—5 p, — thallus pale; lirellae (0. — x 0.3—0.5 mm., straight or flexuous; Mozam vermelbana Vainio ll. Ascospores 15—19 X 4—5 д днн thallus white; Angola..................... ‚ lomdensis Vainio 11. Ascospores 15 X 4 m, эон thallus white; lirellae VR 0.2 mm. ids flexuous tra m Mall Arg. 11. Алы — 6—7 de 8-locular; lirellae simple or with lateral Foo 1- vas ми. Arg. 20-28 X mm. long; lignicole; Soc "e"! x. 0.2 11. С дды lirellae simple ог branc 1-1.5 ние 18-27 X 4 p, 8-9 risu irellae simp’ ids "p ur нар Arg. i X д ; lirellae immersed, simple or usd i m scospores 20—25 5 p, 4-6-locular; lire О. ереен сунам мап. Arg. broad; calcicole; Socotra ........... лает нииииииитинти 12. = 19-21 X 3-4 p 4—6-locular; thallus pale yellowish-glaucous; lir gi adially branched, ан BUGeria. онен rade ge 12. Free 11-13 (-15) × 3-3.5 d н pale yellow, ri p ui nally cracking away; lirella 2 X 0.2 ف‎ Тлар M 8. 13. Argon 47-37 FS e Ч 13 Амара 30—46 Х 5—8 и, 8—10-locular; lirellae 2-3 X 0.2 mm.; Angola....O. grapbidiza — 14. Ascospores 6-locular ............. nennen nnne tnnt ttnnnnnnn ntn 3 14. Ascospores 8—14-locular ............---------- 20 ив 15. T 5 И ; : pet velia PEE viens т а О. loandensis Nyl. sec. Vsioio 15. Ascospores 22 X 6—7 и, apical cells Же lirellae 1-1.5 X pu 3 mm.; mei ABS pd thallus pale greenish; Sierra Leoné 222 coi depen 8. 15. Ascospores 23-35 X 3.5 ш; lirellae radiately Lane P whi aes — мі 0-56 X -locular 16. Thallus olive to glaucous; ascospores 30-56 X 6-8 №, with halo, ou lirella ме X 0. 2 mm., simple, Е or curved; Angola.......... Rutas] A Vainio [Vor. 40 310 ANNALS OF THE MISSOURI BOTANICAL GARDEN 16. Thallus white or pale glaucous 17 17. Lirellae radially branched, 5 X 0.2-0.3 mm.; ascospores 9-locular, 32 X 3 pu; i O. mozambica Vainio 18 Mozambique 17. Lirellae simple or forked 18. Ascospores 32-37 X 3—4 д, 8-9-locular; lirellae white, 0.3-2.5 X 0.2-0.4 mm., K—; Mozambique О. frustulosa Vainio 18. Ascospores 30-46 X 5-8 д, 8—10-locular; lirellae 2-3 X 0.2 mm., straight or flexuous; Angol O. graphidiza Nyl. 18. АЕ 27—40 X 5—6 u, 9—12-locular; lirellae 1-4 X 0.25 mm., simple or forked; thallus рае glaucous to white, К orange; Тапрапу{Ка.....О. subgrapbidiza Zahlbr. ОРЕСВАРНА nigeriensis Dodge, sp. nov. Type: Nigeria, Ondo Province, Owena near Akure, on Theobroma, C. A. Thorold 164. Thallus pallide flavo-glaucus, subpruinosus, margine albo-byssino, angusto, 240 p crassitudine; cortex vix evolutus; stratum algarum 50 y crassitudine, cellulis cylindricis Trentepobliae isodiametricis; medulla 190 p crassitudine, duobus stratis sistens, superum 135 y crassitudine hyphis tenuibus hyalinis laxe intertextis, in- ferum 55 р crassitudine hyphis periclinalibus dense contextum. Lirellae radiato- ramosae, ad 1.5 mm. diametro, thallo immersae vel usque ad 30 д super thallum emergentes; parathecium integrum, 80 p altitudine, carbonaceum, labiis conni- ventibus; hypothecium vix evolutum; thecium rotundatum, 40 y altitudine; para- physes tenues, ramosae anastomosantesque, epithecium obscure brunneum 8-10 p crassitudine formantes; asci ellipsoidei, 24 X 8 p, leptodermei; ascosporae octonae, hyalinae, 4—6-loculares, cellulis cylindricis, 19-21 X 3-4 p. Thallus pale yellowish-glaucous, subpruinose, with a very narrow white byssine margin, about 240 y thick; cortex scarcely differentiated, consisting of very slender hyphae with minute crystals extending a few micra above the algal cells; algal layer about 50 и thick, of Trentepohlia cells closely packed, cylindric, nearly iso- diametric, with scattered cells deeper in the medulla; medulla 190 p thick, in two layers, the upper about 135 p thick, of slender, hyaline hyphae, very loosely woven, e lower about 55 д thick, of compactly woven periclinal hyphae, slightly brownish next the substratum. Lireilae radiately branched groups about 1.5 mm. in diameter, immersed in the thallus or protruding about 30 д above the surface of the thallus; parathecium entire, 80 џ tall, 40 д thick, carbonaceous, lips con- nivent; hypothecium scarcely differentiated; thecium 40 р tall, rounded; para- physes slender, branched and anastomosing, forming a dark brown epithecium 8-10 р thick; asci 8-spored, ellipsoid, about 24 X 8 p, wall thin, only slightly thickened at the tip when young; ascospores hyaline, 4—6-locular, wall rather thick but protoplasts cylindric, 19—21 Х 3—4 p. ' The texture and the color of the thallus resemble those of Chiodecton at first sight, while the radially branched lirellae suggest a Pbaeograpbis. The lirellae are distinct, without thickened and coalescent bases as in the Chiodectonaceae. They are immersed in the algal layer and the upper part of the loose medulla, not reach- ing the substrate as in the Graphidaceae and most other Opegraphaceae. 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 311 OPEGRAPHA PROSODEA Ach., Meth. Lich. 22. 1803. Type: Sierra Leone, corticole, Afzelius. Thallus epiphloeodal, between grape green and vetiver green, minutely verru- cose and somewhat rimulose, without a black margin, 35—55 и thick, almost homoeomerous; algae short vertical filaments of Trentepohlia, cells short-cylindric, 5-6 ш in diameter, terminal cells subspheric and slightly larger. Lirellae black, elevated, linear, straight, about 135 u wide and up to 1.5 mm long, covered by a thin hyaline layer 5-6 м thick, of decomposed hyphae; parathecium 25-30 p thick below and on the vertical sides, thinning to 12-15 p at the somewhat conni- vent lips, carbonaceous; hypothecium 12-15 p thick, of slender, deeply staining hyphae; paraphyses slender, somewhat branched, tips clavate, covered by minute brownish crystals, gelifying as the asci mature; asci 8-spored, narrowly ellipsoid, 80 X 20 д, thick-walled at first, then thin-walled except at the somewhat thick- ened tip; ascospores hyaline to slightly brownish when moribund, long-fusiform, 40 X 5-6 u, 12—14-locular, walls and septa rather thick, but protoplasts cylindric. If Nylander (Ann. Sci. Nat. Bot. IV, 11:229. 1859) was reporting the spores of the type (“ех hb.”), our spores are somewhat smaller. In other respects our specimens agree well with the Acharian description, although it lacks any fuscous shade in the thallus. O. graphidiza Nyl. from Angola differs in a less well-devel- oped thallus and perhaps dimidiate lirellae. Nylander states, “hypothecium fuscescens" in the formal description, using hypothecium in the sense of base of lirella, as he often did, thus implying that the parathecium is entire. He com- pares it, however, with the dimidiate Graphis scripta in the notes at the end of his description, implying that it is dimidiate. Vainio, who presumably studied the type, places it in the section Pleurothecium with dimidiate lirellae. Nigeria: Ina near Ibadan, on Theobroma, C. A. Thorold 150. GRAPHIDACEAE Thallus crustose, cortex amorphous or absent, epi- or endophloeodal, with Trentepohlia algae (Palmella in Xyloschistes); lirellae usually more or less im- mersed in the thallus, or, if sessile, the sides covered with thallus until late, roundish, oblong, simple or branched, single; parathecium entire or dimidiate, carbonaceous, colored or hyaline (very rudimentary to practically absent in Graphina sect. Platygrammodes) ; paraphyses slender, unbranched, tips thickened and warted in Acanthographis; asci clavate when young becoming oblong to cylindric when mature with 8 or fewer ascospores; hypothecium hyaline or pale; ascospores hyaline or brown, septate or muriform, protoplasts rounded to lenticular. 1, Ascospores Зербар сене теле НОО Е реа TS 1. Азсоврогез. muriform а rte einn ccena обес ieiuno el eê line 2. Ascospores brown at maturit ария - : 3. Paraphyses without warted and conspicuously thickened clavate tips........... т ес > 3. Paraphyses with conspicuously clavate and warted tips--...-..---- se: g Loc e 4. Protoplasts of ascospores cylindric; ascospores mostly Е ке _.......Phacographis Müll. Arg. [Vor. 40, 312 ANNALS OF THE MISSOURI BOTANICAL GARDEN 5. Ascospores hyaline 6 5. Ascospores brown : z 6. Paraphyses without thickened and warted tips Grapbina Müll. Arg. hyses with conspicuously clavate and warted tips... Acanthotheciopsis Zahlbr. E a Palmella; lignicole Xyloschistes Vainio Algae Trentepohlia Pbaeograpbina Müll. Arg. GRAPHIS GRAPHIS Adans., Fam. Pl. 2:11. 1763 [uninomial nomenclature]; Ach., K. Vetensk. Akad. Nya Handl. 145. 1809. Fissurina Fée, Essai Crypt. Ecorces Officin. 59. 1824. Aulacographa Leight., Ann. Mag. Nat. Hist. II, 13:389. 1854. Dyplolabia Mass., Neagenea Lich. 6. 1854. Diplograpbis Mass., Atti 1. В. Inst. Veneto Ш, 5:273. 1860. ? Limboria Trev., Conspect. Verruc. 15. 1860, non Ach. Anomorpha Nyl., Lich. Ins. Guineens. 50. 1889. Digraphis Clements, Gen. Fung. 59. 1909. NN Type: Lichen scriptus L. G. pulverulenta, G. Cerasi, G. betuligna, and G. serpentina were described as new at the same time, all now often considered vari- eties of G. scripta (L.) Ach. Fissurina Fée was based оп Е. Dumastii Fée, now placed in Graphis, and F. incrustans Fée, now in Graphina Müll. Arg. Aulaco- grapha Leight. was based on Gra phis elegans Ach. Dyplolabia Mass. was based on Graphis Afzelii Ach. Diplographis Mass. was based on G. rufula Ach., now in Graphis, and G. chlorocarpa Fée, now in Graphina. Limboria Trev. non Ach. belongs here according to the description, although the species cited all belong elsewhere, i.e. L. constellata (Ach.) Trev. is now placed in Diploschistes and Г. tridens (Eschw.) Trev. in Pbaeograpbis. Anomorpha Nyl. and Digraphis Clements were both based on A. £urbulenta Nyl., now usually placed in Graphis. Thallus crustose, epi- or endophloeodal; ecorticate or with cortex of periclinal hyphae; algae Trentepohlia. Lirellae immersed to sessile, elongate or branched, very rarely short and rounded, lips nearly closed (connivent) to open, entire or sulcate; parathecium entire or dimidiate, carbonaceous or subhyaline; hypothecium thin, hyaline; paraphyses slender, unbranched, not thicker at the tips; asci clavate to subcylindric, septate, 2-many-locular, protoplasts lenticulate to subspherical; spermatia exobasidial, cylindric, relatively long (rarely observed). 1. Lirellae wholly densely жерше ВЫ Rah ER E : 1. Lirellae not densely white-pruinose, disc sometimes slightly pruinose. —À 2. al u$ rose color or partly paler; lirellae 0.2-7 mm. long, immersed; ascospores not e seen, ПеПСе genus uncertain; Angola С. roseotincta Vainio - Thallus scarcely visible except at bases of lirellae; São Thomé — allus powdery, bluish white; lirellae long, subflexuous, dimidiate; ascospores 35 X 9—10 ш (as in С. scripta Dd lu a G. caesia Müll. Arg. . Thallus yellowish; lirellae 2—3 оона Ge o ы y 2. Thallus pale ellow; 1 а E РИА N aa n N -4 ; ; 6 X 8 n, 4- (-4) X 1 mm., emersed; ascospores 16 X С. АБАЙ Ach. Thom n С. lynceodes Nyl. 3. Ascospores 30-36 Х 13-15 и, 4-locular; lirellae 1 mm. or less ТВ timidula dis 3. Ascospores 40-70 X 11-18 и; lirellae 1-3 X 0.5 mm., lips sulcate ш С. albonotata Ny 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 313 4. Parathecium dint ua or n 5 4. Parathecium black, 6 4. Parathecium black, dimidiate 14 4. Parathe -— structure unknown; ascospores 24—27 X 6-7 pu, 8-locular; thallus pipe ue infida Nyl. Ma . Ascospores 12-13 X бы, 4-locular; lirellae 0.5-1.3 mm. long; Usambara 5 G fopra Müll. Arg. 5. Ascospores 16 X 8 p, 4-locular; lirellae up to 2 X $ Nigeria...) G. nigeriensis Dodge 5. Ascospores 20 : 5 p, 8-locular, with halo; thalles je poer ire white; Jil im mersed, 0.4—2.5 x о. 15—0.5 mm., ellipsoid; МогатЬдие.............................. С. pallescens Vainio 6. Азсозрогез 4-loc 7 6. Азсозрогез С ocular; thallus white to glaucous 8 Ascospores 12—18-1осшаг 12 і Peesthecium ннан thin; ascospores 17-18 X 5-7.8 и; Мосина РЕСИН ОО Vainio 7. Parathecium emersed, thick on sides, very thin at base; ascospores 14-17 x Mozambique G ме Vainio 7. Parathecium emersed, thin but not thinner at base; ascospores 14—15 X 5-6 ш; Nigeria G Thoroldi Dodge 9 8. Vie Lirellae immersed or subimmer 8. Lirellae semi- to completely наа тн sessile odd unbranched, flexuous; ascospores 23—40 X 7-9 ш, 8—10-locular; Ilha Principe С мия Nyl. еы и" а 0.5—5 X 0.15—0.2 mm.; ascospores 2 K ph loc ры Vainio Lirellae adl branche (0.7—) 2—4 X 0.1 mm.; ascospores 27.5-32 X 7. 5 My, loc 10 hs ^w $ i Vainio 10. E fus at ee: ool pruinose; lirellae mostly unbranched, some s forked; asco- spores 50—56 X 10-12 u, 8-locular; French Tropical Africa....G. aperiens v. pruinosa "ms connivent; thallus whitis Lips н. „тезе 37—49 X 6-9 p, 8— licen lirellae short, unbranched; Guinea............... a (Ach.) Sprgl. 11. Ascospores 25—35 X 6-7.5 u, Siomin lirellae Paraoa, ылы ы or rarely forked, 0.6-0.8 X 0.1—0.15 mm.; us K—; Mozambique................ G. G. subintican Vainio и; — pores 24—26 t-loculars “irs ao чане ет 0.7—2.5 A е po Vainio X thallus K an gena the en 12. хона 32—45 -8. n" и; авсат very thin at the base; Un. emer: Кей пуа С ды Mit. Arg. Х 0.3 mm 12. Ascospores 74—80 X 12-1 4 12. Же, mi x 15:355 и; asci only 2—4-spored; lirellae оо, 1-3 "prem Mall 13. Nile 2. 5 mm. pen везни Ъу ellos, fns etis ИТ НИЕ Schroederi | Zihlbr 13. Lirellae 4 mm. long, emersed, lips sulcat G. ondensis езе 4 14. Ascospores 4(-5)-locular; Модове i 14. Ascospores 8—14-locular 17 15. Ascospore конч connected by plasmodesmata (as in the Кенин. ascospores 18-20 X 7 G. palmensis Vainio 16 16. Ascospores 16. Ascospore $ 17.5-20 X 6-7 u 17. Lips striate-sulcate 17. Lips entire, not striate-sulcate 18. Ascospores 22-24 X 7 и, without halo; Mozambique.-..----------------++-+----+--- oj 18. Ascospores 32—42 X 8-9 и, 8—12-locular; Guinea ы С. ОД rio Pm 18. Asc —11 11—14-locular; lirellae кон curved, cuspidate оп опе diy. EET ыы е z ; strisisla a v. cuspidata Müll. E $ 19. Lips connivent 2 19. Lips open, at least when mo FE T 20. Thallus ; 25-38 7-8 u, 10-14-locular; Angola ochraceous; ascospores x №, ; ими Mall. Ar 15. Ascos toplasts not connected 2 pore pro ê asts no un G. triticella Vainio С. Limae Vainio 20. Thallus white to Me Eco аан 21. виь 0.3-0.7 (-1) X 0.3-0.5 mm., straight, unbranched; vet 20-27 X 7 ar; Socotra С. br еби Mall Arg. [Vor. 40 314 ANNALS OF THE MISSOURI BOTANICAL GARDEN Lirellae 0.5(—1.8) X 0.25 mm., straight, unbranched; ascospores 30 X 8 и, 8—10- locular; Sierra Leone С. Deigbtoni Dodge 21. Lirellae very long and slender; ascospores 21-30 X 7-8 #, 8—10-locular; Guinea....G. tenella Ach. irellae 4-6 X 0.3 ar; Kenya ......... G. oxyclada Mill. Arg. 21. Lirellae 2 X 0.13 mm., curved, somewhat branched; ascospores 27 Х 7-8 р, 10-locular; ш С. guineensis Dodge 22. Ascospores 40—50 X 7-9 и, 8—10-locular; thallus greenish white; lirellae 1-3 Х 0.25 mm.; Usambara ...... 7. aterrima Mill. Arg. 22. Ascospores 55 X 8 и, 11-14-locular; thallus white, parathecium reddish black; Usambara ... G. erythrocardia Mill. Arg. GRAPHIS AFZELII Ach., Syn. Lich. 85. 1814. Dyplolabia Afzelii Mass., Neagenea Lich. 6. 1854. Type: Guinea, corticole, Afzelius. Thallus epiphloeodal, 75 и thick, light brownish olive, without a black margin; cortex 25 y thick, gelified, of periclinal, slender, thick-walled hyphae about 2 y in diameter; algal layer 50 u thick, of subvertical filaments of Trentepohlia about 4 p in diameter; medulla not differentiated, but hyphae penetrating deeply between the bark cells. Lirellae straight, curved or flexuous, rarely once-forked, 2-3 (—4) X 1 mm., about 0.3 mm. tall, densely white-pruinose, lips connivent, thalline cortex with an occasional algal cell extending halfway up the parathecium where it is replaced by a layer of slender vertical hyphae 4 д in diameter, closely septate, somewhat gelified and decomposed, forming a layer about 80 p thick; parathecium 80 р thick above the thecium, expanding to 180 д thick on the sides, narrowing abruptly at the level of the base of the thecium then expanding rapidly to 135 р, . making an acute angle with the bark and extending only slightly under the margin of the hypothecium, thus dimidiate, carbonaceous; hypothecium 25 и thick, the lower half gelified, the upper half of deeply staining, densely woven, slender hyphae; thecium 90—105 p tall; paraphyses slender, simple, tips not thickened; asci 8-spored, clavate, tips not thickened, 80-90 X 8—11 р; ascospores ellipsoidal, 4-locular, 16 X 8 y, terminal protoplasts spherical, central ones lenticular, con- nected by a very slender isthmus which is finally obliterated. The shape of the parathecium is quite variable in cross-section in this species. The above description was based on Deighton M4337. In Deighton M4408, the lips are not so closely connivent, the thecium is wider, and the gelified layer of the hypothecium is 55—65 p thick, slightly yellowish, resting on a thin, discontinuous layer of blackened bark cells. In Deighton M4629, the outer margin of the para- thecium is not so conspicuously angled at the level of the base of the thecium. АП collections cited agree closely in all other characters. G. Afzelii differs from G. nivea Fée, Essai Crypt. Ecorces Officin. 47. 1824 (type from Peru on Cinchona oblongifolia Mutis), to which tropical American material should be referred instead of to G. Afzelii, in the thinner cortex and thallus, relatively thicker algal layer, lirellae dimidiate instead of entire and with a thinner base, lips not sulcate, broader and thinner parathecium, lower thecium and smaller asci and ascospores. 1953] Т DODGE—LICHENS OF TROPICAL AFRICA 315 Sierra Leone: Njala (Kori), on Peltophorum africanum у. speciosum, Е. C. Deighton M4337, on Anisophyllea laurina, Е. C. Deighton M4408, on Citrus aurantifolia, F. C. Deighton M4629. СВАРНБ nigeriensis Dodge, sp. nov. Type: Nigeria, Owena near Ondo, on Theobroma, C. A. Thorold 124. Thallus epiphloeodes, ravus, 55 и crassitudine, homoeomerus, filamentis ver- ticalibus Trenfepobliae 6 и diametro et hyphis tenuibus intertextis cum crystallis ad 10 р diametro. Lirellae flexuosae, usque ad 2 X 0.5 mm., simplices, labiis elevatis, subpruinosis, conniventibus mox delabentibus et discum pruinosum ex- ponentibus; parathecium dimidiatum, superne 18—20 u ad 25 y in lateribus dilatatum, hyalinum vel pallide brunneum, hyphis tenuibus, septatis, 3—4 p diametro, pseudoparenchyma formantibus; hypothecium 15 p crassitudine, hyphis tenuibus dense contextum; thecium 65-70 y altitudine, ad 360 y latitudine; para- physes tenues, simplices, apicibus non incrassatae; asci cylindrici, 65 Х 12 y; ascosporae octonae, imbricatim monostichae, hyalinae, Linie protoplastis terminalibus subconicis, centralibus rotundatis, ellipsoideae, 16 X 8 Thallus epiphloeodal, 55 и thick, drab, cortex not differentiated; ag filaments of Trentepoblia about 6 p in diameter, vertical and penetrating deeply between the bark cells; medulla not differentiated, but of slender, interwoven hyphae in- closing many hyaline crystals up to 10 и in diameter between the algal filaments. Lirellae flexuous, up to 2 X 0.5 mm., unbranched, lips elevated, slightly pruinose, connivent, soon breaking away and exposing the pruinose disc; parathecium dimidiate, 18—20 y thick above the thecium, up to 25 p thick on the sides of the thecium, hyaline or pale brownish, of slender, septate hyphae 3—4 p in diameter, orming a pseudoparenchyma; hypothecium 15 u thick, of densely woven, deeply staining, slender hyphae; thecium 65-70 p tall, 360 џи wide; paraphyses slender, unbranched, tips not thickened; asci cylindric, 8-spored, 65 X 12 p; ascospores imbricately monostichous, hyaline, 4-locular, terminal protoplasts subconical somewhat smaller than the rounded central ones, ellipsoidal, 16 X 8 p Grapuis Thoroldi Dodge, sp. nov. Type: Nigeria, Ina near Ibadan, on Theobroma, C. A. Thorold 151b. Thallus epiphloeodes, laevis, 15-20 р crassitudine, obscure olivaceo-griseus, homoeomerus, filamentis subverticalibus Trentepobliae 6-7 ш diametro. Lirellae rectae, 0.5—0.6 X 0.2 mm., emersae, nigrae, labiis conniventibus; parathecium integrum, carbonaceum, superne 15 ad 35 p in lateribus et sub hypothecio dilatatum; hypothecium 15 ш crassitudine, hyphis tenuibus dense contextum; thecium 80 и altitudine, 125 в latitudine; paraphyses tenues apicibus subclavatis, brunneis; asci clavati, dein ellipsoideae, 55 X 16 p; ascosporae octonae, hyalinae, 4-loculares, anguste ellipsoideae, 14-15 X 5-6 p. Thallus epiphloeodal, smooth, 15-20 p thick, dark olive gray, homoeomerous, of subvertical, partly disorganized filaments of Trentepohlia, 6-7 p in diameter, and slender hyphae, somewhat decomposed above, somewhat thicker near the | ] [Vor. 40, 316 ANNALS OF THE MISSOURI BOTANICAL GARDEN lirellae and disorganizing the cork cells to a depth of 80 u, but the algae not penetrating between them. Lirellae straight, 0.5—0.6 X 0.2 mm., emersed, black, lips connivent; parathecium carbonaceous, entire, 15 u thick at the lips, expanding to 35 р on the sides and under the hypothecium, and blackening the cork cells to a depth of 55 p below it; hypothecium 15 д thick, of slender, densely woven hyphae; thecium 80 д tall, 125 u wide; paraphyses slender, tips slightly clavate, brownish in the epithecial gel, which is about 18 д thick; asci clavate, becoming ellipsoidal, 8-spored, 55 X 16 p; ascospores hyaline, 4-locular, 14—15 Ж би, narrowly ellipsoidal. СкАрніѕ ondensis Dodge, sp. nov. Type: Nigeria, Ondo Province, Ipetu, on Theobroma, C. A. Thorold 120. Thallus epiphloeodes, ad 55 y crassitudine, laevis vel minute verrucosus, opacus, albidus; cortex 8 р crassitudine, hyphis septatis periclinalibus, gelifactus, granulis minutis inspersus; stratum algarum 25 и crassitudine, filamentis verticalibus Trentepobliae 5—6 и diametro; medulla 22 p crassitudine, hyphis tenuibus, magnis cum crystallis hyalinis. Lirellae emersae, 4 X 0.6 mm., curvatae vel flexuosae, margine sulcato, labiis conniventibus, thallo 40 д crassitudine sine crystallis in medulla tectae; parathecium integrum, ad labias 15 p, lateribus 230 p, basi 80 p crassitudine, сит 6—7 sulcis 60 и altitudine, carbonaceum; hypothecium 25 p crassitudine, hyphis tenuibus dense contextum; thecium cordiforme, 105 y alti- tudine latitudineque; paraphyses tenues, pachydermeae, apicibus brunneis non incrassatis; asci cylindrici, 90 X 30 p; ascosporae binae vel quaternae, hyalinae, 12-loculares, protoplastis rotundatis, pachydermeae, 80 X 14 p. Thallus epiphloeodal, up to 55 ш thick, smooth to minutely verrucose, surface dull, whitish; cortex 8 д thick, gelified, of septate, periclinal hyphae inspersed with minute granules; algal layer about 25 p thick, of loosely packed, vertical filaments of T'rentepoblia 5—6 i. in diameter; medulla about 22 p thick, of slender yphae inclosing large hyaline crystals, penetrating about 35 p into the cork cells but not disorganizing them. Lirellae emersed, about 4 X 0.6 mm., curve or flexuous, margin sulcate, lips connivent, covered by a layer of thallus 40 p thick, but with few or no crystals in the medulla; parathecium entire, 15 p thick | at the lips, expanding to 230 p thick on the sides and narrowing to 80 p under the hypothecium, with 6-7 grooves extending about 60 м deep, carbonaceous; hypothecium 25 y thick, of slender, closely interwoven hyphae; thecium cordi- form, 105 м tall and broad; paraphyses slender, thick-walled, unbranched, tips brownish but not thickened; asci 2—4-spored, cylindric, 90 Ж 30 p; ascospores about 12-locular, hyaline, protoplasts rounded, thick-walled, about 80 X 14 р. Grapuis Deightoni Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on twigs of Bauhinia tomentosa, F. С. Deighton M4307E, M4307G Thallus epiphloeodes, albidus, anguste nigro-marginatus, 100 p crassitudine; cortex decompositus, 8—10 p crassitudine; stratum algarum 40 p crassitudine, . 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 317 filamentis verticalibus Trentepohliae 5—6 и diametro; medulla 50 ш crassitudine, hyphis tenuibus laxe intertextis, magnis cum crystallis, ad cellulas suberosas magis compactis periclinalibusque. Lirellae rectae aut subcurvatae, 0.5 (1.8) mm. longi- tudine, semi-emersae, labiis conniventibus; parathecium dimidiatum, superne 15 p, inferne usque ad 55 y dilatatum, lateribus verticalibus non alatis, carbonaceum; hypothecium 15—20 u crassitudine, hyphis tenuibus dense contextum; thecium 120 ш altitudine, 150 м latitudine; paraphyses tenues, semel vel bis dichotome ramosae, apicibus subclavatis, cellula terminali 5—6 и diametro; asci clavati dein subcylindrici, ca. 60 X 16 p; ascosporae octonae, subclavatae, apice uno obtuso, altero magis acuto, 8—10-loculares, protoplastis lenticularibus, 30 X 8 p. Thallus epiphloeodal, whitish, narrowly black-margined, at least in contact with the thalli of other lichens, about 100 и thick; cortex decomposed, 8—10 y thick; algal layer 40 р thick, of vertical filaments of Trentepohlia 5—6 р in diameter; medulla 50 и thick, of slender, loosely interwoven hyphae inclosing large, hyaline crystals, more compact and periclinal next the cork cells. Lirellae straight or slightly curved, 0.5 (—1.8) Х 0.26 mm., semi-emersed, lips black, connivent; parathecium dimidiate (sometimes the outermost bark cells are blackened, then appearing entire but thinner at the base in thick sections), 15 u thick above, ex- panding to 55 y at the base, sides vertical, not winged at the base, carbonaceous; hypothecium 15—20 y thick, seated on the yellowish to blackened bark cells, of slender, densely woven hyphae; thecium 120 м tall, 150 м broad; paraphyses slender, once or twice dichotomously branched, tips clavate, terminal cells brown- ish, 5—6 и in diameter; asci clavate, becoming subcylindric, about 60 X 16 p, 8- spored; ascospores hyaline, 8—10-locular, subclavate, one end obtuse, the other more acute, protoplasts lenticular, 30 X 8 p. Graruis guineensis Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on twigs of Lagerstroemia speciosa, F. C Deighton M4345. Thallus epiphloeodes, ca. 25 p crassitudine, laevis, pallide cinereus vel albidus; Cortex 15 p crassitudine, gelifactus, hyphis periclinalibus; stratum algarum ca. 10 ш crassitudine, filamentis periclinalibus Trentepobliae 5—6 р diametro. Lirellae curvatae, 2 X 0.13 mm., rare furcatae, emersae, labiis conniventibus; parathecium dimidatum, carbonaceum, superné 15 u, ad 35 y inferne dilatum, basi subalatum; hypothecium 20 p crassitudine, hyphis tenuibus dense contextum; thecium sub- cordiforme, 55 м altitudine, 95 y latitudine; paraphyses tenues, simplices; asci cylindrici; ascosporae octonae, curvatae, 10-loculares, 27 X 7-8 p. Thallus epiphloeodal, about 25 p thick, margin indefinite, smooth, pale ashy to white; cortex 15 и thick, gelified, of periclinal hyphae; algal layer about 10 р thick, of periclinal filaments of Trentepohlia 5—6 p in diameter, some penetrating between the bark cells; medulla not differentiated. Lirellae curved, up to 2 X 0.16 mm., some forked, emersed, lips connivent, black, sides covered with thallus; parathecium dimidiate, carbonaceous, lips 15 p thick, sides 35 p thick, very slightly [Vor. 40 318 ANNALS OF THE MISSOURI BOTANICAL GARDEN angled at the base; hypothecium 20 д thick, of slender, densely woven hyphae; thecium subcordate, 55 џ tall, 95 и wide; paraphyses slender, unbranched; asci 8-spored, cylindric; ascospores curved, 10-locular, 27 X 7-8 p. PHAEOGRAPHIS PHAEOGRAPHIS Müll. Arg., Flora 65:336. 1882. Leiorreuma Eschw., Syst. Lich. 14. 1824. Pyrochroa Eschw., Syst. Lich. 15. 1824. Platygramma Meyer, Nebenstudien, 332. 1825. Leiogramma Eschw. in Martius, Icon. Pl. Crypt. Brasil. 2:11. 1828. Grapbidula Norm., Nyt Mag. Naturvidensk. 7:240. 1853. Hymenodecton Leight., Ann. Mag. Nat. Hist. II, 13:387. 1854. Chiographa Leight., Ann. Mag. Nat. Hist. II, 13:388. 1854. Solenograpba Mass., Atti I. R. Ist. Veneto III, 5:270. 1860. Pyrrbograpba Fée in Mass., Atti I. R. Ist. Veneto III, 5:272. 1860. T beloscbisma Trev., Conspect. Verruc. 14. 1860. Type: none designated for Phaeographis Müll Arg. Теюттеита Eschw. was based on Opegrapha Lyellii Sowerby now in Pbaeograpbis. When Eschweiler in- cluded Leiorreuma as a section of his Leiogramma in 1828, he added Graphis scalpturata Ach., now in Phaeographina, and Г. tartareum Eschw., now in Ope- grapba. In his section Lecanactis of Leiogramma, he included L. sericeum and L. punctiforme (both now in Phaeographis) and L. pruinosum (now in Helmintho- carpon). Pyrochroa Eschw. was based оп Graphis coccinea Holl and Р. flammula Eschw., both now in Phaeographis, and Graphis caribaea Ach., now in Arthonia. Platygramma Meyer included P. dendriticum, P. Lyellii (both now in Phaeo- graphis), and Р. suffultum (now in Phaeographina). Graphidula Norm. is а nomen nudum, as no type was designated, nor new combinations made. Hymeno- decton Leight. was based on Graphis dendritica Ach. Chiographa Leight. was based on Opegrapha Lyellii Sowerby. Solenographa Mass. was based on Lecanactis confluens, now in Phaeographis. Pyrrhographa Fée was based on Pyrochroa flam- mula and P. medusulina, both now in Phaeographis, and P. javanica, now in Graphina. Theloschisma was based on T. Eschweileri (Mont.) Trev. (Verrucaria as pistea Eschw. non Ach.). From the above synonymy, it is obvious that many names antedate Phacographis Müll. Arg. Since the latter name has been in constant use by most lichenologists for the last seventy years, a type species should be chosen and Phaeographis should be conserved, but without prejudice against older names if the genus is ivided. There are several groups of species, now in Phaeographis, which differ in important characters and should be segregated as genera. It would be unfortunate, how- ever, if this were done without a thorough study of the species from all regions. Until such a monograph is published, it seems wise to continue the use of Phaeo- graphis for all Graphidaceae with brown, septate ascospores except those now placed in Melaspilea. 1953] DODGE—LICHENS OF TROPICAL AFRICA 319 Thallus crustose, epi- or endophloeodal; ecorticate or with a cortex of peri- clinal hyphae; algae Trentepohlia. Lirellae immersed to sessile, elongate or branched, rarely short and rounded, lips connivent or open, entire or sulcate; parathecium entire or dimidiate, carbonaceous or subhyaline; hypothecium thin, hyaline; para- physes slender, unbranched, tips not conspicuously clavate; asci clavate to sub- cylindric, 4—8-spored; ascospores brown, fusiform to ellipsoid or subcylindric, septate, protoplasts lenticular to subspherical; spermatia exobasidial, cylindric, relatively long (rarely observed). 1. Parathecium pale; ascospores 4- ocular, 13— “ х 6 p; lirellae тич ое disc open, ack, margin densely pruinose; São Thom subnivescens (Nyl.) КИМ: 1 Parathecinm black arathecium ent ; 2. Parathecium dimidiate 11 3. Parathecium thicker below, lips connivent, sulcate 4 3. Parathecium not conspicuously thicker below 5 4. Ascospores $—12-locular, 38-46 X 9-11 п; on palm wood; чере» "EUR Жо P. palmarum M. Arg. 4. Ascospores 4-locular, 27—30 X 7-8 п; corticole; Sierra I P. Deigbtoni Dodge 5. Lips connivent, disc very narrow; lirellae prominent, caine a: or TCE branched 6 5. Lips widely divergent 8 6. Ascospores 4-locular; Mozambique 7 6. Ascospores 10-17 i 55—65 X 8-10 ш; lirellae orange-pruinose; Cameroon P тан Dodge and 6. oed bis ои 85 X 15 в; lirellae white-pruinose at first, soon nude c ck; ний Dodge 7. м Ж. 7 ет dh allus P. micrograpta (Vainio) Zahlbr. 7. Ascospores 18.2—23 X 6.5-7.5 P ib. К reddening..................... P. tigrinella (Vainio) Zahlbr. 8. Lirellae innate, branched; ascospore size not given; Liberia.............. P. dendriticella Mäll. Arg. 8. Litalle rounded, ellipsoid to oblong, not branched woos 9. Foliic аз es 6-locular, 20-22 X 6—7 p; Апдо!а.............- P. Phyllocharis (Vainio) e 9. Cortic 10. Thallus endophloeodal; lirellae prominent, 0.3—1.5 X 0.2—0.3 mm., disc black or pruinose; ascospores 6-locular, 18-28 X 6-7 и; Angola... ers T si^ Ponce Zahlbr. 10. Thallus ч verrucose, NIMM whitish; lirellae innate, us SK 15.5-27 7-7.5 и; МотатЫаче.................-..-.----—- N E FH: ITI Г) Е subdevelans (Vainio) Zahlbr. 11. Lips sulcate or striolate; ascospores 6-locular, 20 X 7.5 м; Usambara......P. duplicans Mill. Arg. ll. Lips striolate; ascospores 6-locular, 25 X 8-9 и; lirellae 0.7 mm. broad, sd, fura te; Usambara фа Mill. А, . 11. Lips not micat or — enti 12. Ascospores 12 or more locular, 100-105 X 13 A asci 4-spored; lirellae seen by thallus; 3 mm. hg unbranched; Sierra Leone... есїа 2a = j ak e AE iT a ; Азс 6-locular, 20—30 65 H res o- , 3. ‘Tie tn simple o or г sparingly "branched, 0.5-2. idee Т. neviculeris ard Zahlbr. i " 6-8-locular, 23-26 X 6.5 коте simple 0.5 5 2 Ж 0,2-0,23 mms NCOP eat: Р. Peel Mall Arg. 13. id 0.2 mm., disc open, pruinose; ascos Lirellae ЕЯ SUMMAS, осони о = ЖИ d е v. тайна МЫ. Ат. 14. ад elingle or daily branched, imm xU. abii "di и 14. Lirellae dichotomously and partly radially Windil 1+ Ж D T qu 5 оное очнь ваз зо чье чье pl ce DA E 16 msi 15. Lirell 02.2.5 X 0.1-0.5 mm.; ascospores 9-16 - js 7, M di {туп бет Дим х 0 и sop là nad Zahlbr. i 6 ; orticole Sio Thomé............ : шаш нә тен pra е ра? Iynceodes (Nyl.) Zahlbr. 16. Asc я CU LUE Р. micrograptoides (Vainio) Zahlbr. 16, M opel i * Коры ыен о a e (Vainio) Zahlbr. [Vor. 40 320 ANNALS OF THE MISSOURI BOTANICAL GARDEN PHAEOGRAPHIS Deightoni Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on bark of Ampbimas pterocarpoides, F. C. Deighton М4334А. Thallus endophloeodes, albidus. Lirellae rectae, subfusiformes, sessiles, 1 X 0.25 mm., nigrae, labiis conniventibus, sulcatis; parathecium integrum, carbona- ceum, basi 185 p crassitudine, in lateribus ad 55 р tenuescens; thecium 135 р altitudine; paraphyses tenues, apicibus non incrassatis; asci clavati; ascosporae octonae, distichae, 4-loculares, brunneae, rectae vel subcurvatae, ellipsoideae vel subfusiformes, 27-30 Ж 7-8 p. Thallus endophloeodal, whitish, consisting of a few Trentepohlia cells and hyphae between the partly disorganized bark cells, making the surface silvery-ashy. Lirellae straight, subfusiform, sessile, about 1 X 0.25 mm., black, lips connivent, somewhat sulcate-striate; parathecium entire, carbonaceous, 185 р thick below, thinning to 55 р thick on the sides; thecium 135 p tall; paraphyses slender, tips not thickened; asci 8-spored, clavate; ascospores distichous, ellipsoid to subfusi- form, brown, straight or slightly curved, 4-locular, 27-30 X 7-8 p. PHAEocRAPHIS ochracea Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on twigs of Bauhinia tomentosa, F. C. Deighton M4307G. Thallus epiphloeodes, 80 , crassitudine, minute verrucosus, nitidus, pallide olivaceo- griseus; cortex 25—30 д crassitudine, hyphis periclinalibus, 3 p diametro, gelifactis; stratum algarum 50—55 p crassitudine, filamentis verticalibus Trente- bobliae 3—4 м crassitudine, magnis cum crystallis; medulla non evoluta. Lirellae sessiles, rectae vel subcurvatae, rarissime dichotome ramosae, 1-2.5 Х 0.5 mm., labiis conniventibus, obscure sulcatis, aurantiaco-pruinosis; parathecium dimidi- atum, carbonaceum, 15—30 д crassitudine ad labias, 30—45 р ad latera, usque ad 185 р in alis dilatatum; thecium cordiforme, 80—105 p altitudine latitudineque; paraphyses tenues, apicibus non incrassatis, sed cum crystallis brunneis; asci clavati dein ellipsoidei, 100 X 25 п; ascosporae octonae, subcylindrici, brunneae, 10-12- loculares, 55—65 X 8—10 и. Thallus epiphloeodal, 80 р thick, minutely verrucose, shining, pale olive gray; cortex 25—30 p thick, of gelified periclinal hyphae 3 и in diameter; algal layer 50—55 p thick, of disorganized, vertical filaments of Trentepoblia 3—4 p in diam- eter, with very large hyaline crystals (such as one sees in the Thelotremaceae)- Lirellae sessile, straight or slightly curved, very rarely once-forked, 1—2.5 x 9. mm., lips connivent, very obscurely sulcate, covered by cortex at first, then the cortex cracking away, exposing powdery orange material about the parathecium; amphithecium consisting of thalline cortex and alcohol-soluble, orange materia which replaces the algal layer on the sides of the parathecium, but not extending into the thallus proper; parathecium dimidiate, carbonaceous, about 15—30 p thick at the lips, 30—45 м on the sides and expanding to 185 р where it makes an acute angle with the bark cells which are sometimes blackened to a depth of 55 p below 1953] DODGE—LICHENS OF TROPICAL AFRICA 321 the thecium (thus giving the appearance of an entire parathecium); thecium cordiform, 80—105 р tall and broad; paraphyses slender, tips not thickened but the epithecium covered with brownish crystals; asci 100 X 25 p, clavate, becoming ellipsoidal, 8-spored; ascospores subcylindric, 55—65 Ж 8—10 м when pale brown, 9—11-septate, shrinking to 30 X 6 p when very dark brown. After the thecium has partly disintegrated, a new thecium forms below the old hypothecium (seen in most of the lirellae sectioned in Thorold 121). As the new parathecium expands, the old parathecium breaks away at the junction with the new parathecium, the scars leaving the new parathecium shallowly sulcate when this has been repeated about four or more times, as the new parathecium is covered by a new outgrowth of the thallus to form each new parathecium. Only a single, shrunken but very dark brown spore was found in Thorold 121 after repeated sectioning of various lirellae, but the other structures agree well with those of the Sierra Leone material. Sierra Leone: Njala (Kori), on Bauhinia tomentosa, Е. C. Deighton М4307С. Cameroons: Tombel, on Theobroma, С. A. Thorold 121. PHAEOGRAPHIS sierraleonensis Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on twig of Baubinia tomentosa, F. C. Deigh- ton M4307Ga. Thallus epiphloeodes, 70-80 р crassitudine, minutisime verrucosus, pallide olivaceo-griseus; cortex 25 р crassitudine, decompositus, hyphis periclinalibus geli- factus; stratum algarum 45—55 p crassitudine, filamentis verticalibus T'rentepobliae 6-7 p diametro; medulla non evoluta. Lirellae usque ad 2 Х 0.4 mm., rectae vel flexuosae, elevatae, juventute cortice thallino tecta, dein nudae, nigrae; labiis con- niventibus; parathecium integrum, carbonaceum, superne 15 p crassitudine, in lateribus 120 p et sub hypothecio 80 р, margine subalato; hypothecium 20 p crassi- tudine, hyphis tenuibus dense contextum; thecium 160 р altitudine, 200 р lati- tudine; paraphyses tenues, super ascos dichotome ramosae, cellulis ultimis clavatis vel rotundatis, 3 р diametro, crystallis subbrunneis minutis tectis; asci clavati dein ellipsoidei, 100 X 40 п; ascosporae octonae?, 16-loculares, protoplastis rotundatis, brunneae, 85 X 15 p. Thallus epiphloeodal, 70—80 р thick, very minutely verrucose, light olive gray; cortex 25 y thick, decomposed, of predominantly periclinal, gelified hyphae; algal layer 45-55 и thick, of vertical filaments of Trentepohlia 6-7 p in diameter; medulla not differentiated. Lirellae up to 2 X 0.4 mm., straight or flexuous, un- branched, elevated, covered by a thin thalline cortex when young, finally nude and black in the upper portion; lips connivent; parathecium entire, carbonaceous, 15 р thick at the lips, expanding to 120 p on the sides and 80 p thick below the hypo- thecium, slightly winged; hypothecium 20 р thick, of slender, densely woven hyphae; thecium 160 y tall, 200 р broad; paraphyses slender, once or twice dicho- tomous above the asci, terminal cells clavate to subspherical, 3 p in diameter, covered with minute brownish crystals; asci clavate, becoming ellipsoidal, 8- > [Vor. 40 322 ANNALS OF THE MISSOURI BOTANICAL GARDEN spored?, 100 Ж 40 y; ascospores long remaining hyaline, finally becoming brown, 16-locular, protoplasts somewhat rounded, about 85 Ж 15 р. РнАЕОСВАРН!$ tecta Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), оп twigs of Bauhinia tomentosa, F. C. Deighton M4307 pro parte minore. Thallus epiphloeodes, 40 ш crassitudine, albidus; cortex gelifactus, 15 и crassi- tudine, hyphis periclinalibus; stratum algarum 25 y crassitudine, filamentis Trente- pobliae periclinalibus ca. 6 р diametro; medulla non evoluta. Lirellae ad 3 Х 0.4 mm., flexuosae, strato thallino tectae, labiis conniventibus; parathecium carbona- ceum, dimidiatum, ad labias 30 p crassitudine, lateribus ad 150 р, extus intusque alatae; hypothecium centro 25 и crassitudine ad margines tenuescens, hyphis tenuibus dense contextum; thecium cordiforme, 130 p altitudine, 180 р latitudine, dein rotundatum; paraphyses tenues, super ascos dichotome ramosae, cellulis terminalibus clavatis, brunneis, 4 и diametro; asci ellipsoidei, 105 Х 27 p; asco- sporae quaternae, brunneae, 12- aut pluri-loculares, protoplastis lenticularibus 100-105 X 13 p. Thallus epiphloeodal, whitish, about 40 p thick; cortex gelified, 15 p thick, of periclinal hyphae; algal layer 25 p thick, of loosely arranged, periclinal filaments of Trentepoblia about 6 р in diameter; medulla not differentiated. Lirellae up to 3 X 0.4 mm., flexuous, completely covered by a thin layer of thallus, lips conni- vent; parathecium carbonaceous, dimidiate, 30 u thick above the thecium expand- ing to 150 д at the base in contact with the bark cells, angled both without and within; hypothecium 25 y thick in the center, thinning toward the margin, of deeply staining, densely interwoven hyphae; thecium cordiform then rounded, 130 р tall, 180 м wide; paraphyses slender, dichotomous above the asci, terminal cells clavate, brown, 4 и in diameter; asci 4-spored, ellipsoidal, 105 X 27 p; asco- spores finally brownish, 12 or more locular, protoplasts lenticular, 100-105 X 15 p- In very old lirellae, the thallus and tops of the lips weather away, exposing the narrow disc, and the tops of the lirellae appear somewhat pruinose. Below the hypothecium, one or two layers of bark cells are blackened, giving the appearance of an entire parathecium with a very thin base in thick sections. Apparently the browning of the spore occurs late, so that nearly mature spores might be loo for in Graphis. PHAEOGRAPHIS LYNCEODES Zahlbr., Cat. Lich. Univ. 2:381. 1923. Graphis lynceodes Nyl., Flora 69:174. 1886. ; Type: Sáo Thomé, Bom Successo, 1150 m., corticole, A. Moller, comm. Henriques. Thallus epiphloeodal, about 65 и thick, deep olive buff; cortex 25 в thick, of compact, more or less vertical, slender hyphae, the outer 6 д partly decomposed, with abundant minute crystals; algal layer of Trentepoblia filaments about 6 Р in diameter, more or less periclinal, rather few; medulla of slender, loosely wove? hyphae with lacunae, in other portions more densely woven. Lirellae variable in 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 323 shape, mostly short and rounded, often confluent in somewhat curved lines, margin and open disc densely but minutely white-pruinose, immersed or nearly so; para- thecium dimidiate, about 10 д thick, not wider at the base, deep brown; hypo- thecium poorly developed, about 6 и thick; thecium 55—60 p tall; paraphyses slender, unbranched, tips not thickened; asci broadly clavate, about 40 X 8 y, walls slightly thickened when young, 8-spored; ascospores early pale brown, 4- locular, protoplasts somewhat rounded, 14-18 X 5-6 и. In the lirella sectioned, cell division of the ascospore is slower and less syn- chronous than is usual in the Graphidaceae. Even after the ascospores in the ascus have begun to turn brown and they have reached nearly the mature size, both 2-locular and 3-locular ones may be observed. The systematic position of this species is somewhat doubtful Nylander states that it is close to Graphis (Phaeographis) inusta in the С. (Phaeographis) dendritica group. Our material, though scant, agrees closely with Nylander’s description except for hyaline spores. Cameroons: Tombel, on Theobroma, C. A. Thorold 136, growing with Pyrenula heteroclita Ach., P. mamillana (Ach.) Trev., and Sarcographa labyrinthica ( Ach.) üll. Arg. GRAPHINA GrapHina Müll. Arg., Flora 63:22. 1880. Diorygma Eschw., Syst. Lich. 13. 1824. Leucogramma Meyer, Nebenstudien, 331. 1825. Hemithecium Trev., Spighe e Paglie, 12. 1853. Thalloloma Trev., Spighe е Paglie, 13. 1853. Glaucinaria Mass., Atti I. R. Ist. Veneto III, 5:319. 1860. Stenographa Mudd, Man. Brit. Lich. 235. 1861. Type: not designated. Diorygma Eschw. was based on D. tinctorum Eschw. (figured). Leucogramma Meyer was based on Г. turgidum, Г. plicatum, Г. con- fertum, L. serpentarium, L. raddacense and L. carneum, all placed as species dubiae in Graphina by Zahlbruckner; none of them have been reported since the original descriptions in Sprengel, Syst. Veg. 4:2:327. 1827. Hemitbecium 'Trev. was based on six species, of which all but Н. chrysenteron (Mont.) Trev. remain in Graphina as now used. Thalloloma Trev. and Stenographa Mudd were both based on Ustalia anguina Mont. Glaucinaria Mass. was based on Graphis Poitiaei Fée, С. holoki «са Mont., С. Jungbubnii Mont., and С. raddacensis Meyer. From the above synonymy, it is evident that we have a case parallel to that of Phaeographis with many names antedating Graphina Müll. Arg., although the latter has been in general use since its publication. In the same way, a type epecies should be chosen and Grapbina Müll. Arg. conserved without prejudice against older names, if the genus is divided. Thallus crustose, epi- or endophloeodal, ecorticate or with a cortex of longi- tudinal hyphae; algae Trentepohlia. Lirellae immersed to sessile, elongate or branched, lips connivent to open, entire or sulcate; parathecium entire or dimidiate, [Vor. 40 324 ANNALS OF THE MISSOURI BOTANICAL GARDEN carbonaceous to hyaline; hypothecium thin, hyaline; paraphyses slender, un- branched; asci cylindric to somewhat rounded; 1—8-spored; ascospores muriform, hyaline, protoplasts rounded. 1. Lirellae Seve branched, lips connivent; para athecium not des ae y op nit Hes um 6—8-locular, 2—3-locellate; asci 6—8-spored; — th G Ай poen чш. Arg, 1. {ыз "imple or very ng a£ branched 2. Lirellae immersed o к 2. Lirellae emersed to t 5 3. Asci vh, rcg ascospores 16-23 X 9-12 pw, 4—6-locular, dy вы: рага- thec e fuscous; Zanzibar coast pyrenuloidea Müll. Arg. 3. Asci «роста; elise 80 X 22 p, 20-locular, 10-locellate; Fw cibo absent; Sierra Le ©. ото Dodge 3. Asci4 іу ascospores 96-100 X 36 д, Dosen: 6—8-locellate; eer fu entire, pe connivent, derbi Sierra Le ort rr Dodge 3. Osporous; pa re m pale, is lips Fea A л E A A a 4. dam б 70-85 u mede lirellae —4 X 0.3 mm.; Abyssinia.......... р rE Arg. 4. ige vi 75-106 X 30—44 y; i 0.2-2 x 0.1 5—0.25 mm.; T thick; ере С. ulcerata үш RÀ Si SUI entire Raca ESSE и: s. ое dimidiate or base hyaline: азс 1—2-spored че ее ^ 5. Рагае m: scarcely Perge dalis Poe ха ау, ep inns er 2 pare ascospores 27-40 X 1 —10-loc 3—4-locellate; Socotr: у Müll. Arg. 6. Par rathecium green TUA k oor thallus Ry. ‘oli ve, wes iose; 8-spored; зе, ves 35 X 10 p, finally ini: Angola (probably belón ngs gp nm pbina) № sorediella Müll. Arg. 6. Pipe rini darker; thallus not sorediose; 1-4 apoted...—.. r£ ii ee 7 7. Parathecium fuscous-rufous or paler; lips open; a 0.7-2.5 x ак. 7 mm.; spores solitary, 110-150 X rn И; giers inea (Vainio) Zahlbr. 7. Parathecium fulvous, “~ connivent; lirellae 3 X 1 p; asci pues ascospores s 96—100 x gos Leo бе Dodge и; disci eum n k; аа white, wrinkled; ascospores solitary, 66—12 7 G пост, (Ny) Zahlbr. $: d ато pe above, paler below; thallus pale ochre; ascospores 0 28-34 и; Б G > Zahlbr. 8. Lips entire, cals arid white i cag л еы онаа оО НИ a 9. — 23-26 X 8 Nineta 2-3-locellate; lirellae 0.7-1 X 0.25—0.3 mm Sóc G. varians Mall. Arg. 9: ا‎ 30-65 X, 1-2 5 9. Азсозрогез 100 X 25 pu; lirellac 1-2.5 X 0.3 mm.; Usambara.............. ene Müll. Arg. 10. Lirellae 0.81.25 X dd os mund ascospores 60—68 x FETE P (90—110 —32 и when s С. heterospora Steiner 10. Lirellae 0. 27-3 X 0.25—0.5 mm.; унала жулан 30—60 x 14-22 p б ique........- на С. Pelle fieri v.m acrior (Vainio) Zahlbr. 10. Lirellae 2-6 X 0.3-0.4 mm.; ascospores 54—65 X 14—16 и; да с ое Е mbrizensis (Vainio) Zahlbr. Grapuina Deightoni Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on Citrus aurantifolia, Е. C. Deighton M4630. Thallus epiphloeodes, cartilagineus, 200 p. crassitudine (usque ad 575 p circa parathecium) з obscure griseo-olivaceus; cortex 80—95 p crassitudine, hyphis verti- calibus dense intertextis, partim decompositus et minutis cum granulis obscuris nubilatus; stratum algarum 55 y crassitudine, discontinuus, filamentis verticalibus Trentepobliae 7-8 и diametro, cellulis brevibus, subrotundatis; medulla 50-65 р crassitudine, hyphis periclinalibus, tenuibus, dense contexta. Lirellae rectae vel subcurvatae, 3 X 1 mm., labiis conniventibus, pluries sulcatis; parc zhecium in- 1953] DODGE—LICHENS OF TROPICAL AFRICA 325 tegrum, fulvum, labia intima 105 p crassitudine, fulva, pallidiori ad thecium; thecium 270 y altitudine; paraphyses tenues, apicibus non iricrassatis; asci sub- cylindrici, 245 X 32 p; ascosporae quaternae, deer ei? hyalinae, 28-loculares, 6—8-locellatae, 96-100 X 36 y. Thallus epiphloeodal, cartilaginous, margin indefinite, 200 р thick (up to 575 p thick next the parathecium), deep grayish olive; cortex 80-95 р thick, of densely woven, predominantly vertical hyphae, the outer 20 д partly decomposed and nubilated with minute, dark granules; algal layer about 55 д thick, somewhat discontinuous, of short, mostly vertical filaments of Trentepohlia 7-8 p in diam- eter, cells short and somewhat rounded; medulla 50—65 y thick, of densely woven, slender hyphae, mostly periclinal, containing some disorganized bark cells. Lirellae straight to somewhat curved, about 3 X 1 mm., filleul buff, lips connivent, several times sulcate in older lirellae; parathecium entire, innermost lip fulvous, 105 p thick, paler next the thecium; thecium 270 м tall; paraphyses slender, tips not thickened; asci subcylindric, about 4-spored, 245 X 32 y; ascospores subdistichous, hyaline, about 28-locular, 6—8-locellate, 96-100 36 u when free from the ascus. GRAPHINA ULCERATA Zahlbr., Cat. Lich. Univ. 2:429. 1923. Grapbis ulcerata Vainio, Cat. Welwitsch Afric. Pl. 2:438. 1901. Type: Angola, Golungo Alto, Cungulungulo, corticole, Welwitsch 245. Thallus epiphloeodal, rimose subareolate, tea green, 300—600 р thick (only 200 u thick in our material); cortex scarcely differentiated, decomposed, about 5 p thick; algal layer about 40 p thick, of Trentepohlia, cells about 5—6 p in diameter, filaments disorganized in a rather compact tangle of slender hyphae; medulla about 165 р thick, very loosely woven, with some disorganized bark cells in the lower portion. Lirellae immersed, rounded or elongate, mostly curved and forked, 0.2-2 X 0.15-0.25 mm., covered by the thallus when young, then lips spreading and disc open at maturity; disc chalky white, pruinose, plane, level with the thalline margin; parathecium entire, about 30 p thick, tawny, of thick-walled pan hypothecium about 30 и thick, of slender, densely woven, deeply staining hyphae; thecium 105—110 p tall; paraphyses slender, unbranched, tips not thick- ened but covered by a tawny granular material in the upper 10 р; азс! clavate at first, becoming ellipsoid, monosporous, about 100 X 25 и (immature); ascospores hyaline, about 20-locular, 6-locellate, 75-106 Ж 35-44 p when free from the ascus. Our specimens have a somewhat thinner thallus with most ascospores nearer the lower limits of the size given by Vainio. Nigeria: Ina near Ibadan, on Theobroma, C. A. Thorold 119. | Sect. Platy$rammodes Dodge, sect. nov. Type: Graphina artbotbelioides Dodge, the only species known so far. Lidice innatae, disco primo a thallo tecto, dein aperto; parathecio nullo. Lirellae innate at first, covered by a thin layer of thallus, disc finally open and level with the thallus; parathecium absent. [Vor. 40 326 ANNALS OF THE MISSOURI BOTANICAL GARDEN The complete absence of the parathecium separates this section from all others in the genus. Perhaps it is closest to the dimidiate sect. Platygrammina Müll. Arg. and the entire sect. Platygraphina Müll. Arg., in both of which the pale para- thecium is less well developed than in the other sections. Platygrammodes bears the same relation to Platygrammina that the Lecanoraceae bear to the Lecideaceae. The complete absence of parathecium suggests Arthothelium of the Arthoniaceae, but all the structures of the thecium are clearly those of Graphina. GnaPHINA arthothelioides Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on twigs of Bauhinia tomentosa, Е. C. Deighton M4307F. Thallus epiphloeodes, pruinosus, 130 p crassitudine, pallide griseus; ecorticatus; stratum algarum 40 y crassitudine, filamentis Trenepobliae 3—4 р diametro, cellulis cylindricis vel subellipsoideis; medulla gelifacta, hyphis 2 u diametro laxe inter- texta. Lirellae orbiculares vel oblongae, rectae vel curvatae, usque ad 2 X 0.5 mm., ex thallo erumpentes, disco pruinoso pallide alutaceo; parathecium verum non evolutum; hypothecium 13 y crassitudine, parce evolutum; thecium 120 p altitudine; paraphyses tenues, apicibus non incrassatis; asci clavati dein ellipsoidei, 170 X 28 p; ascosporae juventute quaternae sed maturae binae, hyalinae, ellip- soideae, 20-loculares, 10-locellatae, 80 X 22 p. Thallus epiphloeodal, margin indefinite, pruinose, about 130 p thick, light mineral gray; ecorticate; algal layer 40 p thick, with occasional cells deep in the medulla, cells varying from cylindric to subellipsoid, the filaments of Trentepoblia curved and partly disorganized, 3—4 и in diameter; medulla 90 р thick, highly gelified, with loosely woven hyphae about 2 p in diameter. Lirellae round to oblong, straight or curved, up to 2 X 0.5 mm., erumpent from the thallus, cov- ered by a thin layer of thallus which finally peels off, leaving an open, flat, pruinose, pale buff disc level with the surrounding thallus; true lips and parathecium not differentiated; hypothecium about 13 p thick, scarcely differentiated; thecium 120 р tall; paraphyses slender, tips not thickened; asci clavate at first, becoming ellipsoid at maturity, 107 X 28 р, 4-spored at first but usually two ascospores abort, leaving only two mature ascospores; ascospores hyaline, ellipsoid, muriform, 20-locular, 10-locellate, 80 X 22 p. PHAEOGRAPHINA PHAEOGRAPHINA Müll. Arg., Flora 65:398. 1882. Thecaria Fée, Essai Crypt. Ecorces Officin. 97. 1824. Ectographa Trev., Spighe e Paglie, 11. 1853. T belograpbis Nyl., Mém. Soc. Sci. Nat. Cherbourg 5:130. 1857, nom. nud. Megalograpba Mass., Atti I. В. Ist. Veneto III, 5:317. 1860. Thecographa Mass., Atti I. В. Ist. Veneto III, 5:316. 1860. Pliarona Mass., Atti I. R. Ist. Veneto HL 5:318. 1960. Leucogramma Mass., Atti I. R. Ist. Veneto Ш, 5:320. 1860, non Meyer, 1825. 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 327 Creograpba Mass., Verhandl. K. K. Zool.-bot. Ges. Wien 10:686. 1860. Leiorreuma Mass., Atti I. R. Inst. Veneto III, 5:319. 1860, non Eschw., 1824. Type: not designated. Thecaria Fée was based on Т. quassiaecola Fée (now in Phaeographina). Ectograpba Trev. was based on Graphis scalpturata Ach. (now in Pbaeograpbina) and Opegrapha Poitiaei Fée (now in Graphina). Thelographis Nyl. was based on Graphis polymorpha Fée. Megalographa Mass. was based on M. bysterina Mass. Thecographa Mass. was based on T. ceramia Mass. (now in Phaeographina). Pliarona Mass. was based on Graphis Montagnei v. d. Bosch (now in Phaeographina). Leucogramma Mass., non Meyer, was based on Graphis chrysenteron Mont. (now in Phaeographina). Creographa Mass. was based on C. brasiliensis Mass. (now in Phaeographina). Leiorreuma Mass., non Eschw., was based on Opegrapha sordida Fée, O. depressa Mont. & v. d. Bosch, O. streblocarpa Bél., and Graphis scalpturatum Ach., the last two transferred to Phaeographina by Müller Argau, the rest probably incorrectly transferred to Graphina by Zahlbruck- ner, as Massalongo states the muriform ascospores finally become brown. From the above synonymy, it is evident that we have a case parallel to those of Phaeographis and Graphina, with many names antedating Phacographina Mill. Arg., although the latter has been in general use since its publication. In the same way, а type species should be chosen and Phaeographina conserved without preju- dice against older names, if г genus is divided. Thallus crustose, epi- or endophloeodal, ecorticate or with a cortex of peri- clinal hyphae; algae ыы, Lirellae immersed to sessile, elongate or branched, lips connivent to open, entire or sulcate; parathecium entire or dimidiate, carbo- naceous to hyaline; hypothecium thin, hyaline; paraphyses slender, unbranched; asci cylindric to somewhat rounded, 1—8-spored; ascospores muriform, brown, protoplasts rounded. l. Disc pruinose 5 1. "ine Es огы pruinose 7 True parathecium absent, a pseudo-parathecium of bark cells and bio forming dark brown 5 paler and пенй below; asci monosporous; ascospores Lx 19 p. 20- п. 10-locellate; Sierra Leone. iie Dodge 2. — parathecium pres 3 4 5 w . present monosporous; parathecium: dark or black above, paler below. : Asi treni locul ` ell Usambara 4. As —90 (—100 15-20 p, 10—18-locular, 3 e ocellate; Usam маан. PU 5 aesiopruinosa E striolata Zahlbr. Ascospores 72-95 X 21-24 р, 20-locular, 5-6-locellate; Lo g ye htoni Dodge . me ria wholly oisi or brownish, not darker above...... 6 Parathecium dark fus idiate; ascospores 42—48 n long, rus @ ан long, 0.7 mm. in diameter, cupulate, m есейе, нА subsessile, rounded to oblong gains e x ch uL d. 6. е ecium pale; ascospores 28-33 X 12-13 - & Са гоман е; veut a ier ет св e 12-13 уе генин Sierra 6. Parathecium pale brown; ascospores 29-32 X m ns itt ы; w мл м . i: res 5075 X 18 p; "ile Pararhecium во not е е рае; ‘ascospores . ыы opbora Gir Dodge - Disc dark blood- 3 lips open; AEE 7035 х 71-6 и; Ё us 70 pt lir ellae sessile, constri icted at che base; ied N ынын P deducts ev) — 190 vx 50—60 д; thallus о е эй De N FE BE TE if E E li i! ЛЕ ruf lirellae immersed; Socotra е ourii | Mill. А. Di eddish 7. isc not r "ca068000009009900000000009000000000 [Vor. 40, 328 ANNALS OF THE MISSOURI BOTANICAL GARDEN 8. Lirellae immersed 9 8. Lirellae sessile 10 9. Ascospores 25—35 X 10 и; thallus ashy olive, tuberculose, sorediose; Кепуа.................... Graphina sorediella Müll. Arg. 9. Ascospores 110 X 26 p; thallus dark fuscous, rimulose-areolate; Angola P. fuscescens (Vainio) Zahlbr. 10. Parathecium entire, hyaline, subobsolete; ascospores 25-30 Ж 11-13 и, 4—6-locular, 2-locellate; Kenya P. paucilocularis Müll. Arg. 10. Parathecium dimidiate 11 1. Parathecium thin, not winged, dark brown; ascospores 45—50 X 15-18 и, 10-locular, 5-locellate; Sierra Leone P. leptotremoides Dodge 11. Parathecium thicker, winged at the base, carbonaceous; ascospores 170 X 30 p, about 20-locular and 8-locellate; Sierra Leone P. alata Dodge m" PHAEOGRAPHINA innata Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on bark of Anisophyllea laurina, Е. C. Deighton M4625. Thallus obscure olivaceo-alutaceus, hypophloeodes, ad 40 p crassitudine, cellulis suberosis emortuis, hyphis periclinalibus et filamentis Trentepobliae 4—5 p diametro. Lirellae immersae, usque ad 4 Х 0.7 mm., curvatae, rare furcatae, apertae, disco albido-pruinoso; pseudoparathecium cellulis suberosis hyphisque, labiis nigro- brunneis inferne brunneum; hypothecium 15 p crassitudine, hyphis tenuibus dense contextum; thecium 110 д altitudine, 375 p latitudine; paraphyses tenues, semel vel bis dichotome ramosae, apicibus non incrassatis, crystallis minutis brunneis tectis; asci monospori, ellipsoidei juventute pachydermei, 80 X 24 и; ascosporae brunneae, murales, 72 X 19 р, saltem 20-loculares, 10-locellatae. Thallus deep olive buff, mostly hypophloeodal, but forming a layer up to 40 p thick, containing some disorganized bark cells, gelified, very slender, periclinal hyphae, and more or less disorganized filaments of Trentepohlia 4—5 p in diameter. Lirellae immersed, up to 4 X 0.7 mm., curved, sometimes once-dichotomous, open, disc white-pruinose; no true parathecium but a pseudo-parathecium of bark cells separated by strands of hyphae, forming dark brown lips, 16-20 џи thick and 50 p long, covering the margin of the thecium and sometimes represented by a brown line up to 8 thick under only a part of the hypothecium, otherwise indistinguish- able from the rest of the bark cells; hypothecium 15 p thick, of very slender, closely woven hyphae; thecium 110 № tall, 375 и broad; paraphyses slender, once or twice dichotomous, tips not thickened but partly covered by minute, brownis crystals in the brownish epithecial gel, 8—10 м thick; asci monosporous, ellipsoid, 80 X 24 p, thick-walled when young; ascospores brown, muriform, about 72 X 19 р, at least 20-locular, 10-locellate. The almost complete lack of parathecium and the dichotomous branching of the tips of the paraphyses are very unusual in Phaeographina, but the affinities of this species are clearly with this genus rather than with the Arthoniaceae (lack of parathecium) or with the. Opegraphaceae (branched and anastomosing рага- physes forming a thick epithecium). 1953] DODGE—LICHENS OF TROPICAL AFRICA 329 PHAEOGRAPHINA Deightoni Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on branch of Peltophorum africanum v. speciosum, F. C. Deighton M4338. Thallus epiphloeodes, 40—55 р crassitudine, olivaceo-alutaceus; cortex 25—40 p crassitudine, hyphis periclinalibus gelifactis, exteris brunneis; stratum algarum ca. 15 р crassitudine, filamentis verticalibus Trenfepobliae 6 p diametro. Lirellae curvatae flexuosaeve, furcatae aut radiatim ramosae, 3—6 Ж 0.2-0.5 mm., labiis thallinis conniventibus, elevatis, mox delapsis, discum pruinosum exponentibus, thallo aequantem; parathecium 15 р crassitudine, superne nigrum, inferne fulvum, integrum; hypothecium 10 р crassitudine, hyphis tenuibus dense contextum; thecium 95 р altitudine, 150 p latitudine; paraphyses tenues; asci monospori, 70 Х 18 p, juventute pachydermei; ascosporae brunneae, murales, ellipsoideae, 20-locu- lares, 5—6-locellatae, 72-95 X 21-24 p. Thallus epiphloeodal, 40—55 p thick, olive buff or darker depending on the color of the underlying bark; cortex 25—40 y thick, of slender, gelified, periclinal hyphae, the outer 10 p brownish; algal layer 15 p thick, of vertical filaments of Trentepoblia about 6 p in diameter; medulla not differentiated, the algal layer resting on the bark cells. Lirellae curved, flexuous, forked or radiately branched, 3—6 X 0.2-0.5 mm., covered by elevated, connivent thalline lips at first, which crack off exposing the pruinose disc level with the thallus; parathecium 15 p thick, black above, fading to tawny in the lower half, entire; hypothecium 10 p thick, of deeply staining, slender, densely woven hyphae; thecium 95 p high, 150 p wide; paraphyses slender, upper 10 м slightly brownish; asci monosporous, 70 Х 18 р, thick-walled at first; ascospores brown, muriform, ellipsoid, about 20-locular and 5—6-locellate, 72-95 X 21-24 y while still brown, shrinking to 55—69 X 15 p, and subfusiform when very dark brown. PHAEOGRAPHINA leucophora Dodge, comb. nov. Lecanactis leucopbora Nyl., Flora 69:176. 1886. PHAEOGRAPHINA scriptitata Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on branches of Citrus aurantifolia, Е. C. Deighton M4628. Thallus epiphloeodes, 90-100 ш crassitudine, cartilagineus, alutaceo-olivaceus; cortex 40—50 p crassitudine, gelifactus, hyphis periclinalibus septatis 3 р diametro, strato algarum discontinuo usque ad 30 p crassitudine, cellulis Trenfepobliae? 10-11 p diametro tectus; stratum algarum verum 40 p crassitudine, filamentis verticalibus Trentepobliae 6—8 p diametro. Lirellae elongatae flexuosaeque, 5 X .5 mm. aut ramosae, subdendroideae, immersae aut subimmersae, disco subcon- Cavo, caesio-pruinoso, madefacto livido, siccitate thecio ab labiis circumscisso; parathecium 55 р crassitudine, integrum, pallide brunneum; hypothecium vix evolutum; thecium 120 y altitudine; paraphyses tenues, apicibus non incrassatis, granulis brunneis tectis; asci ellipsoidei aut fusiformes, stipite 15 р longitudine, [Vor. 40, 330 ANNALS OF THE MISSOURI BOTANICAL GARDEN 115 X 22 p; ascosporae octonae, imbricatim distichae, brunneae, murales, ellip- soideae, 8-loculares, 3-locellatae, protoplastis rotundatis, 29—32 X 12-13 y. Thallus epiphloeodal, 90—100 y thick, cartilaginous, buffy olive; cortex 40—50 p thick, gelified, of periclinal, septate hyphae about 3 и in diameter, covered by a discontinuous algal layer of variable thickness, up to 30 и thick, of disorganized Trentepohlia filaments 10-11 p in diameter; true algal layer 40 p thick, of vertical filaments of Trentepohlia 6-8 p in diameter, resting on the bark cells, with oc- casional filaments penetrating much deeper into the bark. Lirellae immersed or very slightly elevated, long-flexuous, 5 Х 0.5 mm., or variously branched, sub- dendroid, closely aggregated, disc soon exposed, slightly concave and densely chalky-pruinose, livid when moistened, thecium tending to crack away from the lips when dry; parathecium 55 y thick, entire, pale brownish; hypothecium scarcely differentiated; thecium 120 y tall; paraphyses slender, tips not thickened, covered with brownish granules in the upper 10 р; asci 8-spored, ellipsoid to fusiform, stipe about 15 р long, 115 X 22 р; ascospores imbricately monostichous, brown, muriform, ellipsoid, mostly 8-locular, 3-locellate, protoplasts rounded, sometimes slightly constricted at the septa when mature, 29—32 X 12-13 p. The interpretation of this species is difficult. If the outer algal layer rep- resents purely epiphytic growths, it is strange that the algae should be so uniform. The cells are larger and the walls thicker than those of the Trentepoblia filaments in the true algal layer, but they have the usual color of Trentepoblia. "There are also somewhat irregular blackened masses in the parathecium and rarely also below the thecium, but they are apparently adventitious in contrast to the regularly darkened upper portion of the parathecium in P. caesiopruinosa. P. scriptitata seems closest to P. mozambica (Vainio) Zahlbr., from which it differs in its some- what darker thallus, shorter, more curved and branched lirellae, more brownish parathecium which does not extend above the thecium, much broader disc, and its more septate spores. PHAEOGRAPHINA DEDUCTA Zahlbr., Cat. Lich. Univ. 2:437. 1923. Lecanactis Montagnei subsp. deducta Nyl., Flora 69:176. 1886. Lecanactis Montagnei subsp. deducens Nyl, Lich. Insul Guineens. 32. 1889 (lapsus calami, repeating the description verbatim). Grapbis deducta Vainio, Cat. Welwitsch Afric. Pl. 2:436. 1901. Type: Lecanactis Montagnei subsp. deducta Nyl. was based on São Thomé, Rodia, 550 m., Monte Cafe, Saudade, 700 m., A. Moller, comm. J. Henriques, and subsp. deducens Nyl. on Sáo Thomé, 550-700 m., corticole, Moller. Thallus epiphloeodal, 70 р thick, olivaceus black; cortex 30 y thick, of highly gelified periclinal hyphae; algal layer 40 и thick, of rather disorganized, vertical filaments of Trentepohlia 5—8 № in diameter, tending to die off below, leaving lacunae between the hyphae but no medulla differentiated. Lirellae circular, about 1 mm. in diameter, to elongate, flexuous or lobulate, very rarely branched, 5—6 X 1 mm., elevated and somewhat constricted at the base; amphithecium prominent, 1953] DODGE—LICHENS OF TROPICAL AFRICA 331 slightly inflexed, lips spreading, disc open, not pruinose, claret brown; parathecium entire, carbonaceous, about 100 д thick above to 125-150 y thick at the angled base, vertical or slightly spreading, 125 р thick below the hypothecium which is scarcely differentiated; thecium 135 y tall; paraphyses slender with abundant oil droplets, with flexuous, deeply staining hyphae about 3 џи in diameter (resembling latex vessels of the Autobasidiomycetes) which apparently produce the amorphous red covering of the thecium; asci clavate with stipes 15 р long, thick-walled and 4- spored when young, only maturing a single spore; ascospores brown, muriform, 24-locular, 5-locellate, 80—100 X 22 pu. Probably Nylander had only a portion of a thallus with the circular lirellae, when superficially it would look like a large Haematomma, while its microscopic structures would place it in his Lecanactis, since Nylander seldom used spore septation as a generic character. After the ascospores are shed, the thecium quickly begins to disintegrate and is covered by thalline cortex and eventually some thalline algae. Later a new lirella forms on the base of the old parathecium. In contrast to most Graphidaceae where this occurs, only a part of the old parathecium is occupied, so that we may have 3—4 new lirellae along an old one, sometimes with the long axis parallel to that of the old one, sometimes nearly transverse. Fernando Po Island: Izaguirre estate near Botonos, 550 m., on Theobroma, C. A. Thorold 167. PHAEOGRAPHINA leptotremoides Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on bark of Bauhinia tomentosa, F. C. Deigh- ton M4307. Thallus epiphloeodes, superficie laevi, opaca, albidus, 40 p crassitudine; cortex 10 p crassitudine, hyphis periclinalibus gelifactis; stratum algarum 30 p crassi- tudine, filamentis periclinalibus Trentepohliae 5—6 р diametro; medulla non bene evoluta, vicinitate parathecii excepta ubi hyphae tenues crystallos magnos in- cludunt, ut in Leptotremate. Lirellae emersae, usque ad 2.5 X 0.3 mm., basi sub- constrictae, labiis conniventibus, ter profunde sulcatae, nigrae, rectae vel curvatae, rare furcatae; parathecium dimidiatum, ad labias 15 р crassitudine, in lateribus ad 20 в dilatatum, obscure brunneum, hyphis periclinalibus; hypothecium 8 и crassi- tudine; thecium cordiforme, 60 p altitudine latitudineque; paraphyses tenues, apicibus non incrassatis; asci ellipsoidei; ascosporae brunneae, muriformes, ca. 10- loculares, 5-locellatae, probabiliter 45-50 X 15—18 p, nunc collapsae, 30 X 11 p. Thallus epiphloeodal, surface smooth, dull whitish, 40 » thick; cortex 10 p thick, of gelified, periclinal hyphae 5—6 p in diameter; algal layer 30 p thick, of periclinal filaments of Trentepohlia 5-6 p in diameter, medulla not differentiated except in the vicinity of the parathecium where the hyphae enclose very large crystals as in Leptotrema. Lirellae emersed, up to 2.5 X 0.3 mm., somewhat con- stricted at the base, lips connivent, up to thrice deeply sulcate, black, straight or curved, sometimes once dichotomous; parathecium dimidiate, 15 p thick at the lips, expanding to 20 p on the sides, very dark brown, of periclinal hyphae; hypo- thecium about 8 p thick, resting on the bark cells; thecium cordate, 60 р tall and [Vor. 40, 332 ANNALS OF THE MISSOURI BOTANICAL GARDEN broad; paraphyses slender, tips not thickened; asci monosporous?, ellipsoid; asco- spores muriform, brown, at least 10-locular, 5-locellate, probably 45-50 Х 15—18 p, shrinking to 30 X 11 р. The youngest lirella sectioned has the sides covered with the thallus; the parathecium has lips up to 30 y thick, expanding to 55 u on the sides, with the uppermost bark cells darkened; the thecium is only 45 p tall and 80 р broad; the paraphyses are attached at both top and bottom; no asci seen. After the first thecium has disintegrated, a new thecium forms from the old hypothecium and new parathecial walls grow out of the lower inner margins of the old parathecium, spreading the old parathecial lips, leaving a curving space about 20 р wide extend- ing about halfway down the old parathecium, filled with the gelified remains of the old thecium. This process may be repeated, leaving at least three incurved lips. Sometimes the outermost lips break off and the thallus covers the broken stumps. The Cameroons lirella sectioned is still in the one-lipped stage, but the thecium has nearly disintegrated. А single brown shrunken ascospore was found in these sections, and one in the old thecium of an early 2-lipped stage from the Sierra Leone material. Probably before the spores shrank, they measured about 45-50 X 15-18 р, judging from my experience with other species where both mature and shrunken ascospores have been found in the same thecium. | Sierra Leone: Njala (Kori), on bark of Bauhinia tomentosa, Е. C. Deighton M4307. Cameroons: Tombel, on Theobroma, C. A. Thorold 125. PHAEOGRAPHINA alata Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on Peltophorum africanum v. speciosum, Е. С. Deighton M4339. Thallus epiphloeodes, 40 y crassitudine, alutaceo-brunneus; ecorticatus; cellulis Trentepobliae 7 р diametro, subellipsoideis; medulla hyphis tenuibus, verticalibus dense contexta. Lirellae emersae, simplices, rare curvatae vel furcatae, 300 p altitudine, 1-2 X 0.5 mm., dimidiatae, labiis conniventibus; parathecium superne 75 p crassitudine ad 180 p in lateribus basique alata usque ad 330 p dilatatum, carbonaceum, partim strato thallino tenui tectum; hypothecium 6-8 р crassitudine, hyphis tenuibus dense contextum in strato cellularum suberosarum brunnearum 25 p crassitudine impositum; thecium rotundatum, 300 y altitudine, 360 р lati- tudine; paraphyses tenues, apicibus non incrassatis, guttulis oleosis superne in- spersae; asci clavati dein ellipsoidei, 180 X 32 и pachydermei, monospori; ascosporae brunneae, muriformes, ad 20-loculares, dein quoque loculo diviso et cellulis irregulariter dispositis, 175 X 30 p. Thallus epiphloeodal, 40 м thick, between buffy brown and citrine drab; ecorticate; cells of Trentepoblia about 7 р in diameter, somewhat ellipsoid, irreg- ularly arranged in the upper part of the thallus; medulla of densely woven, pre- dominantly vertical hyphae. Lirellae emersed, 300 p tall, 1-2 X 0.5 mm. 1953] DODGE—LICHENS OF TROPICAL AFRICA 333 dimidiate, lips connivent, simple, rarely somewhat curved or forked; parathecium 75 м thick above, expanding to 180 u on the sides and extending 330 р beyond the thecium at the base, carbonaceous, covered on the lower two thirds by a thin decomposed layer of thallus; hypothecium 6—8 p thick, of densely woven, deeply staining, slender hyphae resting on a layer of bark cells about 25 p thick, deeper brown than those below; thecium rounded, 300 p tall, 360 u broad; paraphyses slender, unbranched, tips not thickened, with oil droplets in the upper portion of the thecium; asci monosporous, clavate at first, walls 5 » thick, becoming ellipsoid and thin-walled, 180 X 32 р; ascospores brown, muriform, about 20-locular at first, then each locule is further divided by septa in all three planes and the proto- plasts become quite irregularly arranged, 175 X 30 д. In thick sections, the dark brown bark cells may be mistaken for the base of the parathecium which would then appear entire instead of dimidiate. CHIODECTONACEAE Thallus crustose, homoeomerous or heteromerous, hypothallus often slightly developed, ecorticate or with a slight, almost amorphous cortex; algae Trentepohlia or Phycopeltis. Lirellae in stromata usually immersed, rounded or elongate, simple or branched; parathecium black or hyaline; paraphyses simple or branched and anastomosing; ascospores septate or muriform, protoplasts cylindric or rounded; spermatia exobasidial. The family as now constituted included all the stromatic lichens related to the Opegraphaceae and Graphidaceae. Here we have the same problems we have in the relationships of the Pyrenulaceae and Trypetheliaceae, although there are not so many intermediate species. Ptychographa Nyl. and Diplogramma Mill. Arg. perhaps should belong with this family as they have 2—4 parallel thecia immersed in a sort of stroma. The stroma with its immersed lirella is often called a sarco- thecium. 1. Corticole, rarely saxicole, "t Кенар algae. 2 l. Foliicole, with P. PN peltis 10 2. ры simple, free 3 2. Paraphyses branched and anastomosing 7 3. es with rounded to lenticular protop 4 ind ospor oplas - - ‹ quo with нтте бодон fe and thin dd muriform.............. Enterodictyon Mill. Arg. Asc созрогез septate & Ascospores muriform ? Glyphis Ach. : Ascospores hyaline Sireographa Е Эч с „А 6. AG ЫРДЫ “+ —— n & bee ыа ос оо E Sarcographina Mall An 4 rcd diode septate их . ospores muriform Р n Chiodec 8. Ascospores hyaline aa 2 EH ee Tuc Ach 8. Ascospores brown ee Minksia Mill. Аг i Ascospores hyali depu cain pen 9. Sen Müll. Arg. усто} ем M: An сонное нанео ноет e ree ЕЕЕ № oP Id „2 п own 10. Paraphyses unbranched and free... 10. Pic ise branched and anastomosing -....-.------- [Vor. 40, 334 ANNALS OF THE MISSOURI BOTANICAL GARDEN GLYPHIS GrvPHis Ach., Syn. Lich. 107. 1814. Grapbis subg. Scolaecospora sect. Glypbis Vainio, Etude Lich. Brésil 2:127. 1890. Type: not designated. С. cicatricosa Ach. may be chosen to conserve the name in its present use, as С. labyrinthica and С. tricosa have been transferred to Sarco- grapba. Thallus crustose, epi- or endophloeodal, ecorticate or with an almost amorphous cortex of periclinal hyphae; algae Trentepohlia. Lirellae immersed to subsessile on a black stroma, elongate, branched or somewhat rounded; parathecium black with entire lips, disc flat; hypothecium hyaline; paraphyses unbranched, free; asci clavate with thickened tips, 4—8-spored; ascospores ellipsoid to fusiform, septate, with lenticular protoplasts, hyaline. Ascospores 27—30 X 8 и, 8-locular; sarcothecia 3 mm. in diameter; Guinea..............G. cicatrosa Ach, Ascospores 40-52 X 10 u, 8—10-locular; sarcothecia up to 8 X 5.5 mm.; Mozambiqu G. latissima (Vainio) Zahlbr. GLypuis CICATRICOSA Ach., Syn. Lich. 107. 1814. Graphis cicatricosa Vainio, Etude Lich. Brésil 2:127. 1890. Type: Guinea, on Cordaria acutifolia Afz. (Dialia guineensis Willd.) Afzelius. Thallus mostly hypophloeodal, buffy brown, black-margined, at least next the thalli of other lichens; cortex gelified, about 25 р thick, of periclinal hyphae; fila- ments of Trentepohlia, 5-6 ш in diameter, mostly between the bark cells. Sarco- thecia circular, subcrenate, up to 3 mm. in diameter in our material, margin ashy, narrow, very slightly elevated, lirellae radiately dendroid-branched and lobulate, disc flat to slightly concave, brown, lips thin, spreading, black; stroma carbona- ceous, about 115 u thick; in the younger portions cortex of vertical brownish hyphae covered with minute granules which disappear in the older portions; para- thecium about 15 и thick, usually concrescent with the stroma in the older por- tions; hypothecium about 15 y thick, of densely woven, deeply staining hyphae; thecium 80 y tall; paraphyses slender, tips pyriform and brownish, about 8 X 3 p; asci clavate, 8-spored, thick-walled when young, about 67 X 19 pu; ascospores imbricately monostichous, somewhat twisted, hyaline, fusiform, thick-walled, 8- locular, protoplasts rounded, 27-30 X 8 p. Sierra Leone: Njala (Kori), on Baubinia tomentosa, F. C. Deighton M4307D; M4307G. Var. pulvinata Dodge, var. nov. Type: Sierra Leone, Njala (Kori), on Peltophorum africanum var. speciosum, Е. С. Deighton M4343, Thallus epiphloeodes, 65 н crassitudine, olivaceo-brunneus; cortex 25 p crassi- tudine, hyphis tenuibus periclinalibus gelifactis; stratum algarum 40 p crassitudine, filamentis verticalibus Trentepobliae; sarcothecia pulvinata, basi subconstricta, 1953] DODGE—LICHENS OF TROPICAL AFRICA 335 orbicularia, 1-1.5 mm. diametro, 0.33 mm. altitudine; stroma 220—250 p crassi- tudine sub theciis; parathecium 8 p crassitudine; ascosporae 8-loculares, uno apice obtuso, altero acuto, 32 X 8 д. Thallus 65 u thick, epiphloeodal, olive brown; cortex 25 p thick, of gelified, periclinal, slender hyphae; algal layer 40 p thick, of subvertical filaments of Trentepoblia above the bark cells; sarcothecia pulvinate, slightly constricted at the base, circular, 1-1.5 mm. in diameter, 0.33 mm. tall; stroma 220—250 p thick below the thecia; parathecium 8 p thick; ascospores 8-locular, one end obtuse, the other acute, 32 X 8 p. SARCOGRAPHA SARCOGRAPHA Fée, Essai Crypt. Ecorces Officin. xxxv, 38. 1824. Asterisca Meyer, Nebenstudien, 331. 1825. Actinoglypbis Mont., Syll. Gen. Sp. Crypt. 355. 1856. Type: not designated; S. Cascarillae, S. Cinchonarum and S. tigrina were de- scribed. Asterisca Meyer was based on Glyphis labyrintbica Ach. and С. tricosa Ach. Actinoglyphis Mont. was based on A. Гермеитй Mont. Thallus crustose, epi- or endophloeodal, ecorticate or with an almost amorphous cortex of periclinal hyphae; algae Trentepohlia. Sarcothecia carbonaceous in sect. Eusarcographa or hyaline in sect. Phacoglyphis; parathecium well developed and carbonaceous to rudimentary; lirellae usually branched; paraphyses unbranched, free; asci clavate with thickened tips; ascospores brown, septate with lenticular protoplasts. $ ; i ; 14-20 X 6-8 и, 4—6 (-8) -locular; Dor um нут А: аас e E labyrintbica (Ach.) Müll. Arg. Rar us brownish; parathecium pale; ascospores 18—19 X 5—6 My joco Peras cay Dis ее оО SARCOGRAPHA LABYRINTHICA Müll Arg, Mém.: Soc. Phys. Hist. Nat. Genéve 29:8:62. 1887. Glyphis labyrintbica Ach., Syn. Lich. 107. 1814. Asterisca labyrinthica Meyer, Nebenstudien, 161. 1825. Graphis labyrintbica Vainio, Cat. Welwitsch Afric. Pl. 2:439. 1901. Type: Guinea, corticole, Afzelius. Thallus epiphloeodal, about 80 thick, deep olive buff to citrine drab; cortex 40 р thick, gelified, of densely woven, mostly periclinal hyphae; algal layer 40 ГР thick, of somewhat disorganized, vertical filaments of Trentepoblia 5—6 p in diam- eter. Sarcothecia rounded, up to 4 mm. in diameter, then coalesced into much lirellae labyrinthiform, very narrow, open, d cracking away from one side of the parathecium when dry, giving the appearance of minute, partly connivent lips, [Vor. 40, 336 ANNALS OF THE MISSOURI BOTANICAL GARDEN 8 р thick above to 15 p below, immersed in the stroma of lighter brown pseudo- parenchyma; hypothecium not well differentiated, of closely packed, vertical sep- tate hyphae, somewhat more deeply staining in the lower 25 р of the thecium; thecium 100 м tall; paraphyses slender, tips thickened and brown in the upper 8 p; asci cylindric-clavate, thick-walled when young, 60 X 12-13 y, 8-spored; ascospores imbricately monostichous to subdistichous, oblong-ellipsoid, brownish, 4—6 (-8) -locular, protoplasts rounded, 14—20 X 6-8 p. Sierra Leone: Njala (Kori), on Peltophorum africanum v. speciosum, Е. C. Deighton M4336, on Anisophyllea laurina, F. C. Deighton M4403, and on Citrus aurantifolia, F. C. Deighton M4424, M4626. Nigeria: Ehor near Benin City, on Theobroma, C. A. Thorold 123. Cameroons: Tombel, on Theobroma, C. A. Thorold 122, 136. SARCOGRAPHA (PHAEOGLYPHIS) Thoroldi Dodge, sp. nov. Type: Fernando Po, Izaguirre estate, 550 m., near Botonos, on Theobroma, C. A. Thorold 166. Thallus epiphloeodes, 70—80 ш crassitudine, obscure olivaceo-griseus; cortex gelifactus, 15 м crassitudine, hyphis periclinalibus; stratum algarum 55—65 p crassitudine, filamentis Trentepohliae subverticalibus laxe dispositis, 4—5 p diametro. Sarcothecia rotundata, usque ad 5 mm. diametro, in gregibus irregularibus con- fluentia, 2.5 Ж 1 cm.; stroma pallide brunnea, 130—160 y crassitudine; parathecium superne 20 y crassitudine, inferne ad 40 и dilatatum, pseudoparenchymate lepto- dermeo, sub hypothecio 20 y crassitudine, hyphis periclinalibus pallide brunneis; hypothecium 25 y crassitudine, hyphis subverticalibus dense contextum; thecium 95 p altitudine; paraphyses tenues, apicibus incrassatis brunneis, epithecium 8 р crassitudine formantes; asci juventute pachydermei, cylindrico-clavati, 65 X 8 #3 ascosporae octonae, submonostichae, brunneae, oblongae, 4-loculares, protoplastis rotundatis, 18-19 X 5-6 y. Thallus epiphloeodal, 70-80 в thick, dark grayish olive; cortex 15 p thick, gelified, of periclinal hyphae; algal layer 55-65 p thick, of loosely arranged, sub- vertical filaments of Trentepohlia 4—5 р in diameter, some filaments penetrating deeper between the bark cells. Sarcothecia rounded, up to 5 mm. in diameter, confluent into irregular areas up to 2.5 Х 1 cm., stroma pale brownish, about 130—160 p thick; parathecium 20 p thick above to 40 y below, of thin-walled pseudoparenchyma, 20 p thick below the hypothecium, of periclinal light brown hyphae; hypothecium 25 y thick, of densely woven, deeply staining, subvertical hyphae; thecium 95 p tall; paraphyses slender, tips slightly enlarged and brownish, forming an epithecium 8 и thick; asci thick-walled when young, 8-spored, cylin- dric-clavate, 65 X 8 p; ascospores submonostichous, brownish, oblong, 4-locular, protoplasts rounded, 18-19 X 5-6 y. 1953] DODGE—LICHENS OF TROPICAL AFRICA 337 CHIODECTON Снюрестом Ach., Syn. Lich. 108. 1814. Hypochnus Ehrenb. in Nees, Horae Phys. Berol. 84. 1820, non Fr. 1818. Syncesia Tayl. in Mackay, Fl. Hibern. 2:103. Melanodecton Mass., Atti I. R. Ist. Veneto III, 5:324. 1860. Type: not designated. С. sphaerale and C. seriale described, both in subg. Euchiodecton. Hypochnus Ehrenb. was based on Н. rubrocinctus Ehrenb., a synonym of C. sanguineum (Sw.) Vainio in subg. Byssophorum. Syncesia was based on S. albida Taylor, usually included in subg. Euchiodecton. Melanodecton Mass. was based on Chiodecton sphaerale Ach. and M. indicum Mass. Since the subgenus Enterographa (Fée) Müll. Arg. with hyaline stromata and parathecia is often recognized as a separate genus, I have not included its synonymy here, as none of the names would displace those listed above if the genus were to be divided. Thallus epiphloeodal (sometimes epilithic in subg. Enterographa); ecorticate; algae Trentepohlia. Sarcothecia with immersed or sessile lirellae which may rounded or elongate; parathecium carbonaceous, well developed, greatly thickened below to thin and rudimentary above, hyaline below (in subg. Enterographa) ; paraphyses branched and anastomosing; ascospores fusiform to acicular, hyaline, septate with cylindric protoplasts. 1, True stromata ее A a the bases of parathecia very thick, usually confluent below; | subg. Еос 2 1, Pseudostromata еа и e. containing several ect often sterile; thallus of very loosely oven hyphae and flame nts of Trentepohlia, margin byssoid, wide, usually of a dif- ferent its the rest of the thallus; poe da BYssoPHORUM d 2. es 4-locular 2. т ош, 22—35 u long; disc 0.8-1.3 mm., disces aen poorly d bique rotundatum Vainio 8 2. Ascospores 6—8 Seca! ar 9 2. Ascospores 10—12-locular i 3. Ascospores more than 5.5 м broad; thallus white 3. Ascospores not over road ....... * Ascospores 23—26 xu 5—6.5 ш; Socot С sorts a Mill. xis ospores 30—35 X 6-7 и; j ponencia A ак C. amyloplacoides Yeh мл . Asc зада SENG SO $4 с Sao у м E ae ее res 39—44 Х 3—4 ш; sarcothecia 1-2.5 X 0.8-1.5 mm., disc blackening; Ss » gettin Vainio -.-.............................................. dd pruinose; Moza Thallus margin inconspicuous or ре Фаг еее Sarcothecia aig cular енн ng, . broad; disc 50-100 и in — ascospo: 26—30 и; Ken и inutulum п Mill. Arg. MS Laban apie oblong to со, .8—2 X 0.8-1.5 mm.; ascospores 30 x. тз) 25 Mozam bi м = ozambicum Vainio - eter; sarcothecia 0. T mm.; Socot 8. Run 21-24 X 5 p; disc 20—50 и in diameter; е tê D d i жении ===. вырост bU o mtm енко 8. Азсозрогез 28—44 X 3—4 ш; disc 80—100 и; загсо веса 0.4—2 mm.; Кепуа................ о. E ER PAE E a ' I 2 ME Юю з p: eed x eu ш; Socotra бА и ма rg. 9. Ascospores 34—40 x 2 2 ; disc 300—600 и; Angola. ا‎ С. amyloplacum M 12 -Thallus white; ster ы us 10. Thallus Хунайн ог yellowish; "Ed. LL OE ананетонаа [Vor. 40, 338 ANNALS OF THE MISSOURI BOTANICAL GARDEN 11. Thallus easily separating from the substrate, white above, yellow below, margin some- what tawny; Usambara pocbryseum Müll. Arg. 11. Thallus not easily separating, covered with white isidioid soredia, white below, margin rra Leone C. album Dodge 13 white; Sie 12. Ascospores 4-locular, 45—50 X 4.5-5 п 2. Ascospores 8-locular, 70-80 X 3.5—4 u; t isidiose; many minute rounded lirellae per sarcothecium; hallus glaucescent, black below, surface : Usambar C. Brunnthaleri Zahlbr. 13. Thallus thin, pale green to ashy; lirellae forked or branched, many per sarcothecium; sambara C. intercedens Müll. Arg. 13. Thallus thick, pale yellow; lirellae rounded, few per sarcothecium; Usambara....C. molle Müll. Arg. CHIODECTON (ByssopHoRUM) album Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on trunk of Phyllanthus discoideus, Е. C. Deighton M4396. Thallus albus, byssinus, margine radiato-fibrilloso, minutis sorediis isidioideis tectus, 25—50 y crassitudine; ecorticatus; filamentis Trentepohliae 11 u diametro, cellulis 15 u longitudine et hyphis tenuibus ramosis laxe intertextis cum crystallis usque ad 3 и diametro. Sterilis. Thallus white, byssine, margin radiately fibrillose, covered with minute white isidioid soredia, 25—50 и thick; ecorticate; filaments of Trentepohlia mostly be- tween the bark cells with an occasional branch pushing up through the thallus and forming the soredia, about 11 и in diameter, cells 15 р long; the rest of the thallus of slender, branched and intricately but loosely tangled hyphae 1.5 p in diameter, enclosing abundant hyaline crystals up to 3 p in diameter. Sterile. The habit is similar to Chiodecton (Byssopborum) sanguineum (Sw.) Vainio, but rather thinner, more sorediose and containing hyaline instead of red crystals. As it is sterile, the systematic position of this species is uncertain, since sometimes Cryptotheciaceae have somewhat similar thalli but usually are more closely woven and clearer differentiation of algal layer and medulla. LECANACTIDACEAE Thallus crustose, attached to the substrate by hyphae of the medulla; ecorti- cate; algae Trentepohlia. Apothecia round or nearly so, innate or sessile; amphi- thecium present or absent; parathecium well developed or rudimentary; paraphyses branched and anastomosing; asci 8-spored; ascospores unilocular, septate or dwarf- muriform. Although placed in the Cyclocarpineae by Zahlbruckner, this family shows much closer relationships with the Arthoniaceae and Opegraphaceae, differing from the former by the presence of at least a rudimentary parathecium and from the latter by more-rounded apothecia. l. Parathecium well developed, entire; ascospores hyaline. ss 1. Parathecium rudimentary ...... i ERRE E MEE 2. Ascospores unilocular ......... Pseudolecanactis Zahlbr. 2. Ascospores bilocular, ellipsoidal ................. ...Catinaria Vainio 2. Ascospores 4—16-locular, fusiform to acicular Lecanactis Eschw. 3. Amphithecium present; ascospores septate .... Schismatomma Mass, 3. Amphithecium absent; ascospores septate or dwarf таня .Melampidium Stirton 1953] DODGE— LICHENS OF TROPICAL AFRICA 339 LECANACTIS Lecanactis Eschw., Syst. Lich. 14. 1824; Fries, Syst. Orb. Veg. 274. 1825. Type: based on Opegrapha astroidea Smith & Sow., non Ach., and Lichen lynceus Smith & Sow. Eschweiler also figures L. lobata Eschw. None of these species are now included in Lecanactis, as their apothecia are elongate. Fries added Opegrapha illecebrosa Duf., which now is considered a synonym of Г. amylacea (Ehrh.) Arn. Hence Fries’ usage, based on Opegrapha illecebrosa Duf., should be conserved. Thallus crustose, mostly homoeomerous, attached to the substrate by medullary hyphae; algae Trentepohlia. Apothecia innate to sessile, round, with black, thin parathecium; paraphyses branched and anastomosing, forming a thick epithecium; asci 8-spored; ascospores fusiform to acicular, septate, protoplasts cylindric. Spermogonia spherical, upper half of wall dark; spermatia exobasidial, ellipsoid to cylindri 1. Disc black; ascospores 17-21(—24) X 3.5 м, (4—)6—8-locular; apothecia 1 mm. in diameter; Sierra Leone L. Deigbtoni Dodge l. Disc yellow-pruinose 2 2. Ascospores 20—26 X 4—6 и, 7-10-locular; apothecia 0.6-1 mm. in diameter; Mozam- ique L. flavescens Vainio f 2. Ascospores 15—16 X 4—5 и, 4-locular; apothecia 0.4—0.6 mm. in diameter; Angola У І. flava Vainio Lecanactis Deightoni Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on Cathormion Dinklagei, F. C. Deighton M3734, Thallus hypophloeodes, cortex 18—20 p crassitudine, hyphis tenuibus con- glutinatis; stratum algarum 10-15 y crassitudine super cellulas suberosas, filamentis Trentepobliae 5-6 и diametro, corticem arboris penetrantibus usque ad 140 р. Apothecia sessilia, basi constricta, rotundata, 1 mm. diametro aut greges ad 5 mm. diametro formantia, margine sicco prominente, nigro; parathecium integrum, 15 р sub hypothecio superne ad 40 u, hyphis nigro-brunneis, pachydermeis, periclinal- ibus; hypothecium subbrunneum, 8 м crassitudine, hyphis periclinalibus 2 р diametro; thecium 80 p altitudine; paraphyses sparsae, tenuissimae, superne dense ramosae anastomosantesque, epithecium nigro-brunneum formantes; asci clavati, 56 X 10 р; ascosporae octonae, ellipsoideae, rectae vel subcurvatae, (4—)6-8- loculares, protoplastis cylindricis, 17-21(-24) X 3.5 p. Thallus mostly hypophloeodal, cortex 18—20 р thick, of slender, conglutinate hyphae; algal layer 10—15 м thick above the bark cells, filaments of Trentepoblia 5—6 и in diameter, rather disorganized but penetrating the bark 140 p deep, sep- arating layers of bark cells and forming cavities filled with contorted filaments. Apothecia sessile, constricted at the base, round, 1 mm. in diameter or forming groups up to 3 mm. in diameter, then distorted by mutual pressure but not coalescent, margin slightly prominent when dry, black both wet and dry; para- ecium entire, 15 д thick below the hypothecium expanding to 40 р thick around the thecium, of very dark brown, periclinal, thick-walled hyphae; hypothecium [Vor. 40, 340 ANNALS OF THE MISSOURI BOTANICAL GARDEN about 8 д thick, of periclinal, pale brown hyphae about 2 u in diameter; thecium 80 p tall; paraphyses sparse, very slender, densely branched and anastomosing above to form the dark brown epithecium 20 р thick, thecial gel I blue; asci clavate, 8-spored, 56 Ж 10 р; ascospores hyaline, long-ellipsoid, straight or slightly curved, (4—) 6—8-locular, walls and septa very thin, protoplasts cylindric, 17—21 (-24) X 3.5 p. THELOTREMACEAE Thallus crustose, ecorticate ог with a more or less amorphous cortex; with Trentepoblia ог Pbyllactidium algae, usually with a loosely woven medulla, en- closing large hyaline crystals. Apothecia usually immersed in thalline warts or partly emersed, or several united into a pseudostroma (Tremotylium) or prolif- erating from the margin to form rows of apothecia (Polystroma) ; amphithecium and parathecium well developed, usually partly covering the thecium and opening by a wide ostiole; thecium entire or pierced by a central columella; paraphyses usually simple and free, rarely branched and anastomosing (Gyrostomum) ; asci 1—8-spored; ascospores septate or muriform, hyaline or brown. Apothecia кенеш from the margin of the older apothecia, forming meon. or less erect, forked c of apothecia Polystroma Clement 1. Apothecia seke into — paraphyses branched and anastomosing; ascospor muriform, hyaline or bro anges i 1. Apothecia мэ ог ратан aggregated, but not united into stromata © 2. Corticolous, very rarely saxicolous; algae Trentepohlia уз 2. Foliicolous; algae Phyl ас — ascospores hyaline OSEE HEN Sr: 7 3. Paraphyses br Er and anastomosing Gyrostomum 3. Paraphyses unbranched se ee ЕИ EE AscosporMé septate Еи n LI анаа Ascospores muriform 5. Ascospores hyaline Oce Talaria Nae 5. Ascospores brown Phaeotrema Müll. A 6. Ascospores hyaline ..T helotrema Mh 6. Ascospores brown Leptotrema Mont. & v. d. Bosch 7. Ascospores septate Phyllophthalmaria Zahlbr. 7. Ascospores muriform Chroodiscus “Май. Агу. TREMOTYLIUM TREMOTYLIUM Nyl., Bull. Soc. Linn. Normandie Н; 2:513. 1868. Type: T. angolense Nyl. Thallus crustose, ecorticate or with an amorphous cortex; algae Trentepoblia. Apothecia crowded, forming pseudostromata, each apothecium with its own well- developed parathecium; paraphyses branched and anastomosing; asci 1-8-spored; ascospores hyaline (brown in T. occultum Stirton), muriform, protoplasts rounded. Thallus white; asci 2-8-spored; ascospores 95—170 X 23—46 ш; Апро!а................—- T. angola Nyl. Thallus pale ochre: asci 1—3-spored; ascospores 120—160 X 24—30 д; Cameroons....T. afric m Ras. TREMOTYLIUM AFRICANUM Ris., Arch. Soc. Zool Bot. Fenn. Vanamo 3:185. 1 Type: Cameroons, Bipinde, corticole, С. Zenker (anno 1889). 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 341 Thallus epiphloeodal, smooth, continuous, pale ochraceous, К reddening. Stromata pulvinate, rounded to somewhat irregular, 1-2 mm. in diameter, about 0.8 mm. tall, containing (1-)2—4(—5) apothecia, discs very concave, 0.2—0.7 mm. in diameter, nude or slightly white-pruinose; parathecium black, entire, 125 p thick on the sides, thinning to 75 p thick below the hypothecium; hypothecium thick, I—; paraphyses discrete, branched, slender; asci cylindric, 1—3-spored, I—; ascospores hyaline, muriform, oblong, 120—160 X 24—30 p, protoplasts spherical. Our material is very old, having lost its thecia except for a single ascus ad- hering to the parathecium, showing three ascospores falling within the size range given by Казапеп. Nigeria: Moor plantation near Ibadan, on Theobroma, C. A. Thorold 100, 101. THELOTREMA THELOTREMA Ach., Meth. Lich. 130. 1803. Antrocarpum Meyer, Nebenstudien 326. 1825. Volvaria Mass., Richerche Autonom. Lich. Crost. 141. 1852, probably not DC. 1805. Coniochila Mass., Atti I. R. Ist. Veneto III, 5:258. 1860. Schistostoma Stirton, Proc. Phil. Soc. Glasgow 11:312. 1879. Type: not designated, but Lichen lepadinus would conserve the genus in the sense that it has been used for the last 150 years. Antrocarpum Meyer and Volvaria Mass. were also based on this species. Coniochila Mass. was based on Thelotrema varioloides Hampe nom. nud., from Ceylon, not T. varioloides (Pers.) Ach. from Europe. From Massalongo’s generic description, Т. varioloides Hampe seems close to (if not identical with) Schistostoma dehiscens Stirton from South India, and closely related to T. schistostomoides Zahlbr. (T. schistostoma Müll. Arg. non Tuck.). Schistostoma Stirton was based on S. debiscens Stirton. Thallus epi- or endophloeodal, crustose, ecorticate or with an amorphous cor- tex; algae Trentepohlia; medulla usually loosely woven. Apothecia immersed in the substrate or in thalline warts; parathecium well developed, covering the thecium when young, then rupturing and forming an ostiole; paraphyses simple, unbranched; asci with 8 or fewer ascospores; ascospores hyaline, muriform with rounded protoplasts. Spermatia exobasidial, short-cylindric. . = - -locular; columella subconic; thallus w | 1 se фаг багу, 110—114 X 22-35 р, 34-locular; c cages red Vainio red; Mozambique A дшш шины нс ш шый ыш 1. -locular; columella absent; thallus dark olive / Arne ot СЕ с pi m 1. Ascospores 8 per ascus, under 25 м long; colume Па not mentioned... SN ae 2. Thallus olivaceous; ascospores 18-25 X 5-10 А, omen е d gola TE OR аня [Vor. 40, 342 ANNALS OF THE MISSOURI BOTANICAL GARDEN THELOTREMA (PsEUDo-AsciiUM) cameroonensis Dodge, sp. nov. pe: Cameroons, Banga, on Theobroma, C. A. Thorold 141. Thallus epiphloeodes, 110—120 y crassitudine, obscure olivaceo-alutaceus; cortex 55 p crassitudine, hyphis pachydermeis periclinalibus, 5—6 м diametro, exteris brunneis; stratum algarum 55—65 y crassitudine, filamentis Trentepohliae minutis cum crystallis (rarius magnis) inter hyphas; medulla non evoluta. Apothecia in verrucis thallinis, 1—1.2 mm. diametro, 1 mm. altitudine, ostiolo non papillato; parathecium integrum, 210 p crassitudine superne, ad 40 д sub hypothecio tenu- escens, non alatum, ostiolo 250 и diametro, areola nigra, 200 и latitudine circum- datum; hypothecium vix evolutum; thecium 430 y altitudine; paraphyses tenues, flexuosae, ramosae, apicibus liberis; asci non bene visi; ascosporae binae vel quaternae, hyalinae, muriformes, ca. 42-loculares et 6-locellatae, 185 X 22 p. Thallus epiphloeodal, 110—125 и thick, dark olive buff; cortex 55 p thick, of thick-walled, periclinal hyphae 5—6 p in diameter, the outer ones brownish; algal layer 55—65 м thick, of filaments of Trentepohlia 6-7 ш in diameter, with many minute crystals and occasional larger ones; medulla not differentiated. Apothecia іп thalline warts, 1-1.2 mm. in diameter, 1 mm. tall, ostiole not papillate, about 250 p in diameter; parathecium exposed as a black area 200 p wide around the ostiole, in turn surrounded by a paler area of about the same extent; parathecium carbonaceous, entire, 210 џи thick above, thinning to 40 u below the hypothecium, not winged at the base, resting on the cork cells of the bark; hypothecium scarcely differentiated; thecium 430 м tall; paraphyses slender, flexuous, branched, tips free in the thecial gel; asci not clearly seen, ascospores in pairs or fours in the thecial gel; ascospores hyaline, muriform, about 42-locular, 6-locellate, 185 X 22 p. Occasionally there are two confluent thalline warts with two ostioles but the apothecia are not crowded as in sect. Tremotylopsis Zahlbr. OCELLULARIA OcELLULARIA Meyer, Nebenstudien 327. 1825. Thelotrema subg. Ocellularia Vainio, Etude Lich. Brésil 2:262. 1890. Ascidium Fée, Essai Crypt. Ecorces Officin. xlii, 96. 1824 Ocellularia sect. Ascidium Müll. Arg., Flora 64:525. 1881. Myriotrema Fée, Essai Crypt. Ecorces Officin. xlix, 103. 1824. Ocellularia sect. rem Zahlbr. in Engler & Prantl, Die Nat. Pflanzenfam. 1. 1:176, 1905 Stegobolus Mont. in Hooker's London Jour. Bot. 4:4. 1845. Ectolecbia Mass., Alcuni Gen. Lich. 10. 1853, лоп Trev. 1853. Brassia Mass., Atti I. R. Ist. Veneto III, 5:259. 1860. Coscinedia И Atti I. В. Ist. Veneto Ш, 5:256. 1860. Chapsa Mass., Atti I. В. Ist. Veneto III, 5:257. 1860. Ocellis Clements, Gen. Fung. 80. 1909. Type: not designated. Two species were transferred from Thelotrema (T. obturatum and T. urceolare, both now in Ocellularia) and six from Pyrenula, of which belong in Pyremula as now used). Ascidium Fée was based on 1953] DODGE—LICHENS OF TROPICAL AFRICA 343 Cinchonarum Fée. Myriotrema Fée was based on M. olivaceum and M. album Fée. Stegobolus Mont. was based on S. Berkeleyanus Mont. Ectolechia Mass. was based on Ascidium rhodostoma Mont. Brassia Mass. was based on Thelotrema porinoides Mont. & v. d. Bosch, now in Ocellularia. Coscinedia Mass. was based on Thelotrema microporum Mont., now in Ocellularia. Chapsa Mass. was based on C. indicum Mass. Ocellis Clements was a segregate for species with bilocular ascospores, and was probably based on Ocellularia myriopora (Tuck.) Müll. Arg., but the combination was not formally made. Since Ascidium Fée and Myriotrema Fée are both older than Ocellularia, the last should be conserved, based on Thelotrema obturata or T. urceolare, in order to avoid a very large number of new combinations. However, it should be with- out prejudice of Ascidium and Myriotrema, if а future monographer decides to split the genus. Both names are already in use for very distinct sections of Ocellularia. hallus crustose, uniform, heteromerous; ecorticate or with an amorphous cortex; algae Trentepoblia. Apothecia more or less immersed in substrate or in thalline warts; parathecium and amphithecium covering the thecium when young, then rupturing stellately or forming a circular ostiole, or wholly breaking away above the usually white pruinose thecium; central columella present or absent; paraphyses unbranched; asci 1—8-spored; ascospores septate with lenticular proto- plasts, hyaline. l. Ascospores 4—6-locular Ascospores 6—10-locular; thallus ashy, yellowish or fuscous 2. Thallus hice ak ега present іп the 2. Thallus glau ucous; columella absent; ascospores ST E 45-53 X 11-14 д; ostio only 40 д in diameter; Guinea gea n (ac Dodge 2. Th 11 -gray; 4(—6)-locular, 18-25 X 8-10 з гие, olive E nmm ape O. VEM eS vem бә) Zahlbr. 3. Asc -1 15—17 6 u; apothecia 0.5-1 mm. in diameter; Sierra spore ee x giu O. disc муун (Ach.) ? Mall. Arg. 3. А - —20 6-7 и; ostiole 0.1-0.2 mm. in тая eter; Mozam- Mae ii a 26 а О. mozambica (Vainio) Zahlbr. 4. "Thallus n K red, margin indeterminate; ascospores 27.24 x 8-10 в, 9 1 Кеп Е 9 a— d: a а Hi рч ә i» е А А Hue Ocular; Kenya ©... чиш ере ншнен brio tret ioni rite grise kien . Poncinsiana 4. Thallus feror ien columella present; ascospores 20-30 X 7-8 p, 6—8-1осшаг; Sierra Leo: . h; columell t mentioned; ascospores ие pe ser Sierra Leon E eti SENS ote (Ach) Mall. Ace. 1 4. ть T olive; ascos ata; apotheci: Shae ы. size, mo pore essed with a smaller de a Se ot mentioned; а Ие ОЯ ea oes = и med Zahlbr. a th 135 м tall; ‘она 0.4 5. Pee rl 22-30- dicli; jue ie 3 (-5.5) ш; thecium и thallus dark olive buff; Nigeria ......———————————memmmm О. scolecospors Dodge OcELLULARIA SUBTEREBRATA Zahlbr., Cat. Lich. Univ. 2:601. 1924. Thelotrema subterebrata Nyl., Flora 69:174. 1886. Type: São Thomé, Pico, 1500-2100 m., А. Moller, comm. J. Henriques. Thallus epiphloeodal, light olive gray, 100 p thick; cortex 6-8 p thick, de- мл . Ascospores Майы, 36—60 Х 6-9 и; thecium 90 p tall; apothecia 0. 5-1 e [Vor. 40 344 ANNALS OF THE MISSOURI BOTANICAL GARDEN composed; algae periclinal filaments of Trentepohlia, 5—6 p, filling the rest of the thallus, closely packed and partly disorganized, and penetrating the bark at least 65 p; medulla absent. Apothecia rather crowded but distinct, 0.3—0.4 mm. in diameter, lip elevated about 0.1 mm. above the surface of the thallus, innate, ostiole 150 м in diameter; parathecium dimidiate, 55—60 № thick, pale brownish, darker at the lips, of slender, interwoven, periclinal hyphae, nubilated with minute granules; hypothecium 12 y thick, of conglutinate, interwoven periclinal hyphae; thecium 115 р tall, 140 p in diameter; paraphyses slender, not dense, once or twice dichotomous above the asci, tips clavate; asci 8-spored, cylindric, 95 Х 15 y; ascospores imbricately monostichous, hyaline, 6-locular, protoplasts slightly rounded, 16 Х 5-6 и. Our material, described above, has slightly smaller ascospores but as Nylander stated, has the appearance of О. microspora. Nylander also stated "subsimilis T. terebratae," but it is not clear if this implies the presence of a columella which is lacking in our material. Sierra Leone: Кепета (Nongowa), on Copaifera copallifera, F. C. Deighton M5012. OcELLULARIA trypanea Dodge, comb. nov. Pyrenula trypanea Ach., Syn. Lich. 119. 1814. Verrucaria trypanea Sprgl., Syst. Veg. 4:1:244. 1824. ype: Guinea, corticole, Afzelius. Thallus epiphloeodal, 135 м thick, deep sea-foam green; cortex scarcely dif- ferentiated, 5—6 д thick, of two layers of conglutinate, periclinal hyphae; algal layer about 130 y thick, usually a narrow layer 20 p thick, nearly separated by a perithecioid, opening by an ostiole about 40 и in diameter, surrounded by a narrow blackened area, immersed in thalline warts, about 1 mm. in diameter and 0.3 mm. tall; parathecium hyaline (pale yellowish in thick sections), 13—14 y thick, of conglutinate, periclinal hyphae, surrounded by the algal layer of the thallus; hypo- thecium 15 д thick, of slender, densely woven hyphae; thecial gel filling the cavity, paraphyses somewhat dichotomously branched, tips free, not compact; asci clavate, thin-walled when young, 8-spored; ascospores hyaline, fusiform, 6- locular, protoplasts rounded, 45-53 X 11-14 p. е thallus and very thin, hyaline parathecium clearly relate this species to the Thelotremaceae rather than to the Pyrenulaceae, although the habit and cross- section suggest the Pyrenulaceae, especially with the low magnifications available to Acharius. In old apothecia, the thecial gel turns brown, agreeing with Acharius’ observation. On the other hand, I have not found the ostiole clearly papillate at any stage as reported by Acharius. Nigeria: Ondo Province, Erinmo, on Theobroma, C. A. Thorold 134. 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 345 OcELLULARIA CAVATA Müll. Arg., Flora 65:499. 1882. Thelotrema cavata Ach., К. Vetensk. Akad. Nya Handl. 33:92. 1812. Type: Sierra Leone, corticole, Afzelius. Thallus cartilaginous, verrucose, wrinkled, ashy greenish-fuscous. Apothecia gibberulous, margin of broad ostiole entire, tumid, somewhat verrucose, disc black, covered with a glaucous pruina; parathecium black; ascospores 25—43 X 6-8 p, (6—) 8-10-locular, fide Müller Argau. In our collections, a single thallus may belong here. The thallus agrees in color with Acharius’ description (deep olive buff) and is 125-185 y thick, cortex about 25 р thick, of thin-walled, conglutinate, mostly periclinal hyphae; algal layer 15 р thick, continuous, of short filaments of Trentepohlia, 5-6 и in diameter, partly disorganized; medulla 85—145 p thick, of thick-walled, closely interwoven hyphae, penetrating deeper between blackened cork cells. Apothecia 0.5-0.7 mm. in diam- eter, 0.3 mm. tall, flattened above and coarsely white-granular, easily mistaken for a member of the Pertusaria velata group when the thecium is replaced by soredia. Parathecium about 25 и thick, hyaline, of large-celled pseudoparenchyma surrounded by a yellow layer 20 и thick, of disintegrated bark cells and very minute crystals, which is in turn surrounded by a thin layer of thallus. The thecium has disintegrated and been replaced by huge crystals 65 X 25 p or more. Near the parathecium in a bit of disintegrated thecium, a single ascospore 37 X 8 р was found with 8 rounded protoplasts, well within the range of size reported by Miller Argau. Nigeria: Abdo Ekiti near Ondo, on Theobroma, C. A. Thorold 139. OcELLULaRIA scolecospora Dodge, sp. nov. Type: Nigeria, Moor plantation near Ibadan, on Theobroma, C. A. Thorold 60 Thallus hypophloeodes, obscure olivaceo-alutaceus, ex isidiis granularibus 15—25 p diametro; stratum algarum 25 p crassitudine, 25 p sub superficie corticis T beobromatis, filamentis Trentepobliae 5-6 p diametro, periclinalibus inter cellulas suberosas. Apothecia orbicularia, 150 р altitudine, 400 y latitudine, sine columella, non aut parce basi constricta, erumpentia, disco albo-pruinoso; pseudo-amphi- thecium 25 y crassitudine, cellulis suberosis, hyphis paucis cellulisque Trentepobliae; parathecium 15—25 y crassitudine, integrum, hyphis pachydermeis, periclinalibus, fuligineis, lateribus verticalibus intus cum cellulis hyalinis magnis laxis; hypo- thecium vix evolutum; thecium 135 p altitudine; paraphyses tenues, subdichotome ramosae, rarissime anastomosantes, apicibus non incrassatis, pruina crystallorum tectae; asci fusiformes, 130 X 15 pm; ascosporae octonae, hyalinae, 22 (-30)- loculares, protoplastis rotundatis, curvatae vel flexuosae, 50-66 X 3 )-5.5( p. allus phloeodal, dark olive buff, from minute granular isidia, 15—25 р in diameter; algal layer about 25 p thick, situated about 25 y below the surface of the bark, filaments of Trentepohlia, 5—6 p in diameter, periclinal between the cork cells, somewhat disorganized, with occasional filaments above and below the main [Vor. 40, 346 ANNALS OF THE MISSOURI BOTANICAL GARDEN layer, the lower cork cells blackened, forming a layer 15 и thick. Apothecia round, 150 д tall, about 400 u broad, without a columella, not or slightly constricted at the base, disc densely white-pruinose, carrying a layer of cork cells 25 и thick, with hyphae and a few algal cells up to the top of the parathecium, thus forming a pseudo-amphithecium; parathecium 15—25 р thick, entire, of dark fuliginous, periclinal, thick-walled hyphae, appearing carbonaceous in thick sections, inside which is a nearly hyaline layer of very large, loosely packed cells; hypothecium scarcely differentiated; thecium 135 yp tall; paraphyses slender, somewhat dicho- tomously branched and anastomosing in the thecial gel, tips not thickened, covered with a thick pruina of crystals (which mostly wash away in sectioning and mounting); asci fusiform, about 8-spored, 130 X 15 y; ascospores hyaline, fusi- form, 22-30-locular, protoplasts rounded, 50-66 X 3(—5.5) и, when free from the ascus, breaking apart into short sections of 6-8 cells, flexuous or curved, twisted about each other in the ascus. Perhaps this species should be taken as the type of a new genus, homologous with Gyrostomum Fr. in the series with brown, muriform ascospores. The spore measurements are rather unsatisfactory, as I have been unable to free whole spores from the ascus without breaking them. In the ascus they are so twisted about each other that I have been unable to see both ends at once. In Thorold 161 and 117, the ascospores are less twisted in the ascus and measure approximately 50—66 X 3 рапа are at least 30-septate. This species differs from О. albescens (Vainio) Zahlbr. in lacking a columella, a darker thallus, a taller thecium, and narrower, more septate ascospores. Nigeria: Moor plantation near Ibadan, C. A. Thorold 160, type; Ina near Ibadan, C. A. Thorold 162; Ondo Province, Owena near Akure, C. A. Thorold 117, 161, 163; Abdo Ekiti, C. A. Thorold 118; Akure, C. A. Thorold 162; all on Theobroma. GYALECTACEAE Thallus crustose, homoeomerous or heteromerous, usually ecorticate, with Trentepoblia or Phyllactidium algae. Apothecia immersed to sessile, solitary; amphithecium often present; parathecium hyaline (dark in Sagiolechia); asci 8- many-spored; ascospores hyaline, from unicellular to muriform, with thin septa and cylindric protoplasts when septate and cubical protoplasts when muriform. 1. Corticole or saxicole, with Trentepohlia Wee ee 2 1. Foliicole with Phyllactidium п 6 . Asci SPOR use ыа еы аи 3 2. Asi T2-many-epored |... л Вене. 3. Parathecium dark, entire Auc qp А Sagiolecbia Mass. 3. Parathecium hyaline or light-colored, У uod E Ои. 4. Ascospores оке a OT lonaspis Th. Fr. 4. Ascospores bi Е DEI Dimerella Trev. (Microphiale Zahlbr.) 4. Ascospores 4 or more locular .......... Secoliga Norm. 4. Ascospora MIHO |... n Gyalecta Ach. 5. Ascospores bilocular, fusiform -a Ramonia Stzbgr. 5. Ascospores 6—pluri-locular, fusiform to асісијаг. Pachyphiale Lónnr. 6. Дороге bilotuler mT Lecaniopsis Zahlb ГА 6. Ascospores 4-locular, acicular ........................ Semigyalecta Vainio 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 347 IONASPIS Ionaspis Th. Fr., Lichenogr. Scand. 1:273. 1871. Type: not designated, seven species listed, all still included in recent mono- graphs. Thallus crustose, uniform or effigurate, epi- or endolithic (epiphloeodal in I. ascidioides) , homoeomerous, algae Trentepohlia. Apothecia immersed to adnate; parathecium pale or dark; paraphyses simple or sparingly branched above; asci clavate, 8-spored; ascospores unicellular, hyaline, ellipsoidal with a thin wall. Spermatia short, cylindric, straight. IoNaspis ascidioides Dodge, sp. nov. Type: Nyasaland, Kasungu Hill, 1100 m., L. J. Brass 17458a’. Thallus epiphloeodes, minute verrucosus, rimulosus, 150—175 p crassitudine, albidus, anguste nigro-marginatus; ecorticatus; filamentis Trentepobliae 8 p diametro, dense aggregatis, verticalibus, lacunis sphaericis exceptis, 40-80 p diametro, crystallis magnis hyalinis impletis; medulla non evoluta. Apothecia 0.5-0.6 mm. diametro, sessilia, basi constricta, margine crasso involuto dein tenuiori et thecio aequante, integro aut verticaliter striolato; discus nigro-brunneus, dense albo-pruinosus; amphithecium 90 p crassitudine, stratum algarum 40 p crassi- tudine, filamentis periclinalibus, stratum interius lacunis, magnis cum crystallis; parathecium integrum, hyalinum, pseudoparenchymaticum, 18-24 p crassitudine; hypothecium 10 р crassitudine, hyphis tenuibus; thecium 65 р altitudine; para- physes conglutinatae, tenues, dichotome ramosae, apicibus non incrassatis, brunneis; asci clavati dein cylindrici, pachydermei, 55 Х 14 p; ascosporae octonae (an 12nae?), hyalinae, uniloculares (juventute false biloculares), leptodermeae, 8—9 x 6-7 Ё Thallus epiphloeodal, minutely verrucose, rimulose, 150-175 p thick, whitish with a very narrow black margin; ecorticate; algae Trentepoblia, filaments vertical, closely packed except for more or less spherical lacunae, 40—80 p in diameter, filled with large hyaline crystals. Apothecia 0.5—0.6 mm. in diameter, constricted at the base, margin thick and inrolled when young, becoming thinner and level with the very dark brown, densely white, pruinose disc, entire, becoming slightly vertically striolate; amphithecium 90 y thick, of an algal layer 40 p thick of periclinal filaments, the rest of the lacunae filled with large crystals; parathecium entire, hyaline, of small-celled pseudoparenchyma, 18—24 p thick; hypothecium 10 р thick, of slender, deeply staining hyphae; thecium 65 р tall; paraphyses con- glutinate, slender, dichotomous, tips not thickened in the brown epithecial gel; k-walled, tip of protoplast acute to mamillate, 8 (-12?) - asci clavate-cylindric, thic falsely bilocular when young, spored, 55 X 14 p; ascospores hyaline, unicellular, wall thin, 8—9 x 6-7 p. ; The thallus and amphithecium with large lacunae filled with very large crystals and the shape of the apothecium suggest 2 relationship to Ocellularia subg. Ascidium of the Thelotremaceae, which has quite different ascospores. The algae are clearly filamentous (Trentepohlia), hence exclude this species from Lecanora. [Vor. 40, 348 ANNALS OF THE MISSOURI BOTANICAL GARDEN I have hesitated to refer this species to Ionaspis, which is otherwise saxicolous with immersed apothecia so that the older species were first described in Гесапота subg. Aspicilia, but the homoeomerous thallus with Trentepohlia algae, as well as the spores, seems definitely to belong here, unless one enlarges the concept of the Thelotremaceae to include a new genus with unicellular ascospores. The mature spore is certainly unilocular, although young ascospores have a narrow zone of deeply staining granules across the middle, making them appear falsely bilocular, as one often sees in the Antarctic species of Lecidea. SECOLIGA ЅЕСОІЛСА Norm., Nyt Mag. Naturvidensk. 7:230. 1853. Cryptolechia Mass., Alcuni Gen. Lich. 13. 1853. Phialopsis Koerber, Syst. Lich. Germ. 169. 1854. Bryophagus Nitschke in Arn., Flora 45:38. 1862. Gyalecta sect. Secoliga Tuck., Syn. N. Amer. Lich. 1:218. 1882. Type: not designated. Of the six species included, Lichen ruber Hoffm. may be chosen, as it best fits the generic description. Phialopsis Koerb. was also based on this species. Cryptolechia Mass. was based on Lecanora carneolutea Ach. Bryopbagus Nitschke was based on B. Gloeocapsa Nitschke. allus crustose, uniform, epilithic, growing over mosses, or corticole; ecorti- cate; algae Trentepohlia; medulla loosely woven. Apothecia long immersed in the thallus then becoming sessile; parathecium waxy or horny, light-colored, nude or more usually covered by an amphithecium, disc concave; paraphyses unbranched; asci 8-spored; ascospores fusiform to ellipsoidal, septate with cylindric protoplasts. Spermatia exobasidial, cylindric, straight. 1. Ascospores 6-locular, 16—18 X 3 u; thallus s isl ues apothecia 0.3—0.7 mm diameter, disc chap ochre to clay color; Sier $ Dupin Dolg 1. Ascospores Art ах 18-30 X 4-5 p; thallus е в 2. Apothecia 0.3-0.5 т mm., сє reddish bd cde; Usambara ersicolor Müll. 2. арзы нка 8 mm., disc whitish; Апроја....................... S. vite (Vainio) Dodge SEcoricA plurilocularis Dodge, comb. nov. Gyalecta plurilocularis Vainio, Cat. Welwitsch Afric. Pl. 2:427. 1901. SEcouicA Deightoni Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on pebbles, F. C. Deighton M4840. Thallus epilithicus, laevis, partim subrimulosus, 80 p. crassitudine, olivaceo- alutaceus, homoeomerus; filamentis Trentepobliae periclinalibus, 7-8 p diametro. Apothecia sessilia, basi constricta, 0.3—0.7 mm. diametro, disco ochraceo-argillaceo, margine pallidiori; parathecium 65 p crassitudine, superne ad 40 р tenuescens, pseudoparenchymate cellulis radiantibus, pachydermeis, 4-6 и diametro; hypo- thecium 55 y crassitudine, brunneum, hyphis tenuibus, 3 p diametro, periclinalibus, leptodermeis; thecium 65—80 y altitudine; paraphyses conglutinatae, apicibus non incrassatis, brunneis, subdichotome ramosae super ascos; asci pachydermei, clavati, vel cylindrici, 60 X 16 р; ascosporae octonae, hyalinae, leptodermeae, fusiformes, 6-loculares, 16-18 X 3 p. 1953] DODGE—LICHENS OF TROPICAL AFRICA 349 Thallus epilithic, smooth, partly subrimulose, 80 u thick, deep olive buff, homoeomerous; algae Trentepohlia, filaments mostly periclinal, 7-8 р in diameter. Apothecia sessile, constricted at the base, 0.3-0.7 mm. in diameter, disc yellow ochre to clay color, margin paler; parathecium 65 y thick below, thinning to 40 р above, hyaline, of radiating pseudoparenchyma, thick-walled cells 4—6 и in diam- eter; hypothecium 55 y thick, brownish, of thin-walled, mostly periclinal hyphae 3 ріп diameter; thecium 65—80 y tall; paraphyses conglutinate, tips not thickened in the brownish epithecial gel, somewhat dichotomous above the asci; asci 8-spored, clavate-cylindric, thick-walled, 60 X 16 p; ascospores hyaline, thin-walled, fusi- form, 6-locular, 16-18 X 3 y. COENOGONIACEAE Thallus spongy-byssoid, adnate or forming dimidiate, shelving masses; homoeo- merous with Trentepoblia or Cladophora, whose filaments are only partially sur- rounded by hyphae. Apothecia with pseudoparenchymatous parathecium; азс! 8- spored; ascospores hyaline, uni- or bilocular; spermatia exobasidial. ....Coenogonium Ehrenb. Thallus with Trentepoh Racodium Pers. Бора Thallus with Cladophora, apothecia unknown .............. ern COENOGONIUM COENOGONIUM Ehrenb. in Nees, Horae Phys. Berol. 120. 1820. Type: C. Linkii Ehrenb. pU А Thallus loosely spongy-byssoid, either adnate or forming dimidiate shelving masses (suggesting a thin species of Polystictus in shape but not in texture), omoeomerous with Trentepohlia filaments partially covered by a network of hyphae. Apothecia scattered on the upper surface, scutiform, usually with a very short stipe; parathecium of thin-walled pseudoparenchyma; paraphyses орысы; often with swollen tips; asci clavate, 8-spored; ascospores hyaline, fusiform or long-ellipsoid, 1—2-celled. CoENoconium Deightoni Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on trunk of Phyllanthus discoideus, F. C. Deighton M4396a. Thallus tenuissimus, filamentis repentibus Trentepobliae 26 p diametro, cellulis 75 p longis, cylindricis, hyphis tenuibus, 5 p diametro, partim corticatis. Apo- thecia 0.5—0.6 mm. diametro, in filamentis sessilia, juventute urceolata, dein plana, disco cinnamomeo-brunneo, margine subpallidiori, maturitate non elevato; para- thecium inferne 55 p crassitudine, non superne tenuescens, dimidia parte extera pseudoparenchymate radiante, pachydermeo, cellulis са. $ j diametro, extus granulis flavis insperso, dimidia parte interna hyphis tenuibus periclinalibus dense contextum; hypothecium vix evolutum; thecium 40 Р altitudine; paraphyses tenues, conglutinatae, apicibus moniliformibus sed non incrassatis; asci clavati, 25 X 5 р; ascosporae octonae, fusiformes, hyalinae, uniloculares, 6-8 X 2.5-3 p. [Vor. 40, 350 ANNALS OF THE MISSOURI BOTANICAL GARDEN Thallus very thin, of repent filaments of Trentepoblia 26 u in diameter, corti- cate, with hyphae 3 » in diameter. Apothecia 0.5-0.6 mm. in diameter, sessile on the algal filaments, urceolate at first, becoming plane, disc cinnamon brown, margin slightly lighter, not elevated at maturity; parathecium 55 p thick below the hypo- thecium, not thinning above; the outer half composed of radiating thick-walled pseudoparenchyma, cells up to 5 р in diameter, the outer 15 p. inspersed with yellow granules, the inner half of slender, periclinal, densely interwoven hyphae; hypo- thecium scarcely differentiated from the inner layer of the parathecium; thecium 40 р tall; paraphyses slender, unbranched, conglutinate, tips moniliform but not thickened; asci clavate, 8-spored, about 25 X 5 и; ascospores hyaline, fusiform, unilocular, 6-8 X 2.5-3 y. CoENOGONIUM congensis Dodge, sp. nov. Type: Belgian Congo, Yangola, 20 km. west of Yangambi, on twigs of Ouratea brunneo- purpurea Gilg, J. Louis 12070. Thallus dimidiatus, radius 1 cm., filamentis Trentepohliae 5—6 y diametro, cellulis ad 8 р longitudine. Apothecia convexa, sessilia in filamentis, ad 1 mm. diametro, disco ochraceo-aurantiaco, margine concolore, subdentato; parathecium 135 p crassitudine sub hypothecio, superne ad 105 № tenuescens, dimidia parte extera hyphis rectis septatis, pachydermeis, hyalinis extus subbrunneis, pseudo- parenchyma formante, dimidia parte interna hyphis pachydermeis periclinalibus; hypothecium vix evolutum, 10 № crassitudine, subbrunneum; thecium 55 y alti- tudine; paraphyses tenues, conglutinatae, apicibus pallide brunneis non incrassatis; asci cylindrici, 50 X 5 y; ascosporae octonae, monostichae, fusiformes, biloculares, hyalinae, leptodermeae, 6 X 2.5 y. Thallus dimidiate, radius 1 ст., of filaments of Trentepoblia 5—6 p. in diameter, cells about 8 » long. Apothecia convex, sessile on the algal filaments, up to 1 mm. in diameter, disc ochraceous orange, margin concolorous, slightly dentate; para- thecium 135 р thick below, thinning to 105 p thick at the margin, outer half of vertical, thick-walled, septate hyphae 4 шіп diameter, forming а pseudoparenchyma, hyaline, the outer 8 p slightly brownish, the inner half of thick-walled, densely woven periclinal hyphae; hypothecium scarcely differentiated, represented by a slightly brownish layer about 10 м thick; thecium 55 р tall; paraphyses slender, unbranched, conglutinate, tips not thickened in the pale brown epithecial gel; asci 8-spored, cylindric, about 50 Х 3 p; ascospores monostichous, fusiform, bilocular, hyaline, very thin-walled, 6 X 2.5 y. COLLEMACEAE Thallus gelified, crustose to foliose or dwarf-fruticose, with or without rhizinae; ecorticate or corticate with one or two layers of isodiametric cells. Apo- thecia immersed to sessile, amphithecium often present; parathecium often rudi- unicellular to muriform. Оо оао 21232... 8 LESE а ыл. + со 1953] DODGE—LICHENS OF TROPICAL AFRICA 351 Besides the species of Collema included below, there are a few scraps of at least two species of Leptogium, one completely unidentifiable and the other in the L. vesiculosum group, probably not that species and too meagre for description. The other tropical genera are small and seldom collected. | COLLEMA CoLLEMA Wiggers, Primit. Fl. Holsat. 89. 1780. Latbagrium S. Е. Gray, Nat. Arr. Brit. Pl. 1:399. 1821. Letbagrium Mass., Mem. Lichenogr. 90. 1853. Synecboblastus 'Trev., Caratt. Tre Nuov. Gen. Collemac. 3. 1853. Blenothallia Trev., Caratt. Tre Nuov. Gen. Collemac. 3. 1853. Type: C. Lactuca (Web.) Wiggers, a synonym of Lichen crispus L. now in sect. Blenothallia. Lathagrium S. Е. Gray was based on С. flaccidum Ach., C. nigrescens (Huds.) DC., both in sect. Collemodiopsis Vainio, and C. scotinum Ach., later transferred to Leptogium. С. nigrescens may be chosen as type if the section Collemodiopsis is raised to generic rank. Lethagrium Mass. is probably only a variant spelling of Lathagrium. It included C. fasciculare Wigg., С. rupestre (Sw.) Rabh., C. ascaridiosporum Mass., C. conglomeratum Mass. and C. turgidum Ach., the last now placed in Leptogium. Synechoblastus Trev. was based on C. aggregatum (Ach.) Röhl., C. nigrescens (Huds.) DC., and C. rupestre (Sw.) Ra Since the latter two species belong in the section Collemodiopsis, C. aggregatum may be chosen as the type when Synechoblastus is used as a separate genus. Blenotballia Trev. was based on C. cheilea Ach. and C. baemaleia Smrft., the latter now placed in Pyrenopsis. Thallus squamulose to foliose, gelified, without rhizinae, homoeomerous; ecorti- Cate (except the amphithecium in sect. Collemodiopsis); algae Nostoc, hyphae loosely arranged in the algal gel. Apothecia immersed at first then erumpent, sessile to scutellate, constricted at the base; amphithecium well developed, para- thecium present but more or less rudimentary; paraphyses simple, septate; asci 8-spored; ascospores acicular, fusiform or ellipsoidal (2-locular in sect. Dicollema, 3-pluri-locular in sect. Collemodiopsis and Synechoblastus), broadly ellipsoid and dwarf-muriform in sect. Blenothallia. ' COLLEMA NIGRESCENS (Huds.) DC. var. ммотом Hue, Mém. Soc. Bot. France 8:130. 1910 Type: French Tropical Africa, Chari, Baguirmi, corticole, A. Chevalier. Thallus dark olive, foliose, membranous, 32 и thick, monophyllous, 2—4 cm. in diameter, lobes 0.4—12 mm. broad, radiating, contiguous or somewhat imbricate, tips rounded, surface smooth with radiating wrinkles, paler beneath, costate, partly clinging to the underlying bark by hyphae, but not developed into rhizinae. Hyphae 2—3.75 и, mostly perpendicular to the surface of the thallus, thin-walled, sparsely septate, in the center of the thallus periclinal and forming a loose medulla about 20 р thick, without algae. At the surface a few periclinal hyphae forming а [Vor. 40, 352 ANNALS OF THE MISSOURI BOTANICAL GARDEN highly gelified and almost amorphous cortex 8—10 р thick, in contrast to the clearly cellular cortex of Leptogium; algae Nostoc, cells 2.5—3.75 p in diameter, in curved, moniliform chains. Apothecia 0.4—0.6 mm. in diameter, at first immersed in the thallus, then short-pedicellate, disc rufous, plane to convex; cortex in amphithecium and pedicel 15—20 и thick, of cells 8—10 м in diameter, with thalline hyphae radiating between the parathecium and the amphithecial cortex between the algal colonies; parathecium hyaline, 50—80 м thick below the thecium, thinning to 20 » on the sides and 10 y at the top, cells 5-15(—18) д in diameter below the thecium, smaller and less conspicuous above; hypothecium 40 y thick, of densely woven, periclinal hyphae; thecium 100 y tall; paraphyses 4—5 p in diameter, thick- walled, lumen 1.5-1.7 р, up to 3 м in the terminal cell; asci 8-spored, clavate, tips thickened, 75 X 12 и; ascospores hyaline, fusiform, 4-locular, 27-32 X 4—5 р In this small variety, the ascospores are smaller, as those of the species are about 3442» Yu Nigeria: Agbaola and Айа near Ibadan, on Theobroma, С. A. Thorold 114, 115, respectively. Cameroons: Tombel, on Theobroma, С. A. Thorold. LECIDEACEAE Thallus crustose, uniform or with effigurate margins, continuous, areolate or squamulose (foliose in Psora and dwarf-fruticose in Sphaerophoropsis), attached to the substrate by hyphae of the medulla or hypothallus, without rhizinae; ecorti- cate or with a cortex of fasciculate, thick-walled hyphae, never pseudoparenchy- matous; algae Protococcus or Trebouxia; medulla loosely woven or with a basal layer suggesting the structure of the upper cortex. Apothecia round, sessile, oc- casionally immersed or with a very short stalk; amphithecium absent; parathecium entire or dimidiate, hyaline or carbonaceous; hypothecium hyaline; paraphyses simple or sparingly forked above the asci, free or conglutinate; asci usually 8- spored (fewer than 8 in Mycoblastus and Megalospora, or 16—32 in a few species of Lecidea and Bacidia); ascospores hyaline (brown in Rhizocarpon), unilocular, septate or muriform, with or without a halo. Spermogonia immersed, spermatio- phores unicellular; spermatia elongate-ellipsoid to cylindrical, often very long. A a <. Азсорфогы MERE Lacu шс RR eeu EL л я 1. Ascospores muriform .......... . Ascospores small and thin-walled, uninucleate 2. ае ed thick-walled and large, multinucleate — ыы H 3. Ascospores bilocular ............ 3. Ascopore 4—shuri-localer ео 5 4. Ascospores under 30 u long, slender, thin-walled Catillaria Th. Fr. 4 pores over 40 м long, stout, thick-walled. _Megalospora Mey. & Fw 5. Thallus ecorticate, smooth or mo n granular. DNtrs 5. Thallus corticate, verrucose to subsquamulose. Toninia Mass. 6. Paraphyses unbranched; ascospores hyaline. Lopadium Koerb. 6. Paraphyses branched; ascospores brown with halo Rhizocarpon Ram. 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 353 LECIDEA LEcmEA Ach., Meth. Lich. 32. 1803. Type: not designated. It is probable that when we have a modern monographic treatment of this genus, it will be divided into several genera on the morphology of the apothecium. Until then, it seems best not to select a type species from the many included in the genus when it was segregated from Licben L. Most of those included belong in Lecidea as now recognized. Thallus crustose, continuous, areolate, verrucose, attached to the substrate by hyphae of the medulla or hypothallus, without rhizinae; ecorticate or with a thin Cortex, sometimes sorediose, very rarely with true soralia; algae protococcoid. Apothecia round or angular from mutual pressure, very rarely slightly elongate, immersed, sessile or very short-stalked; parathecium hyaline, colored or black, en- tire or dimidiate; hypothecium hyaline or somewhat brownish; epithecium bright- colored or black; paraphyses unbranched or sparingly so above the asci; asci usually 8-spored, rarely 16-spored; ascospores hyaline, unicellular (sometimes falsely bi- locular), spherical, ellipsoid or allantoid. Spermogonia immersed, with a dark mouth, spherical; spermatiophores unicellular; spermatia short-cylindric to filiform, straight or curved. All of our material belongs in the subgenus Biatora with hyaline or brightly colored, more or less waxy parathecium and colored disc, sometimes darkening in age, usually corticolous or lignicolous. For completeness, I have included a key to the subgenus Eulecidea with a very dark, usually carbonaceous parathecium and black disc, usually saxicolous. Parathecium light-colored, waxy, never carbonaceous, disc reddish, yellowish fuscous to — very dark brown (almost black); corticolous (very rarely saxicolous) ; subg. BIATORA............ 2 l. Parathecium very dark brown, usually carbonaceous disc black or greenish black; saxicolous or terricolous; su E Х 2-3 hi Daly granular-furfuraceous; apothecia u Я 2. Ascospores Pe er ascus, 5—8 0 p И TS : L. furfurosula "E diues uscous, entire; $20 Th Ascospo: us 3. Disc eat аса hair-brown (benzo brown to fuscous when pruina is rubbed off); parathecium hyaline; hypothecium fuliginous above; thallus of hemi- spheric warts with a к margin; medulla yellowish; Guinea........ L. granifera (Ach.) Vainio 3. Disc not whi in 4 4. Disc testaceous, eh cde. reddish, rarely yellowish when young 5 4. Disc fuscous 11 4. Disc almost bla 3 5. Ascospores 4.5—6 Х 2-3 и; thallus olive green; disc flesh-red; hypothecium yellowish- hyaline; Usambar L. pannosa Müll. Arg. 5. Ascospores 8—11 X 5. кон larger and eade: thallus ashy eret 7 E ENS to gd white; disc коё pale yellow flesh-colors a” Mall Arg. n— eee ия 6. Thallus dark ashy to clay woo — subvitelline, vex; ascospores 8—11 X 2-2.5 и gola... crescens recente tenente tete [Vor. 40, 354 ANNALS OF THE MISSOURI BOTANICAL GARDEN 7. Apothecia 0.3—0.7 mm., disc rufous-fleshcolor, plane; ascospores 13—14 X 6-7 enya L. carneoru a ма. Arg. 7: Apotheci 0.5—0.6 mm., org buckthorn brown; hypothecium pale nitas: ascospores 4—5.5 и; Nigeri nigeriensis Рон 7. Apotheci 0.5—1.2 mm., cat testaceous or yellowish athecium асы black below, disc testaceous; ascospores 15 ш long; medulla yellow ; Usambar І. bypomeloides Müll. Arg. 8. Sarecbaeium узе ен outer half hyaline, inner half fuliginous, disc isabelline; asco- ores 11-12 X sp 1- 5—6 и; я А-н ite; Guinea L. rubina Ach. B. bah bem dimidiate, at least p к disc EE Angola 9 9. Hypothallus black; ascospores 9—12 = 6-8 L. caliginosa у. rhacocarpa Vainio 9. Hypothallus white or inc i 10 10. Ascospores — X 4—5 и; parat thecium pale L. gyalectoides Vainio 10. Ascospores 8—1 —7 д; parathecium fuscescent L. capo, nA p: Vainio it: di ame ecium fuscous black, apothecia 0.5-0.8 mm.; ascospores 12-16 X 67 Ery E ra Vainio 11. Hypocheci ecium 12 12. Thecium ba apothecia 0.5—0.75 mm.; ascospores 9—11 X = ш; Kenya........ L. endochrysea Müll. Arg. 12. Thecium hyaline; apothecia 0.3 mm.; ascospores 9—11 X 6-7 в; Ilha Principe........ L. citima Nyl. 12. Thecium hyaline; apothecia 1 mm.; ascospores 12 X 6 д; Usambara phaeopbtbalma Zahlbr. 13. Apothecia 0.4—0.5 mm.; thecium 45 д tall; ascospores 7—15 X 3-5 ш; Angola................ L кн Vainio $3. Sota М 5 mm.; ascospores 11—13 X 8-9 ци; saxicole; Kenya р belon Б асуга Mall. "N Ix ема 14. Thallus нот thecium 75-84 u tall; ascospores 12 X 6 в; Usambar L bhecophtbéine Zahlbr. 14. Thallus smooth; thecium 60—65 ш tall; ascospores 15 X 4.5 и; Usambara І. amaniensis Zahlbr. verruculose to granulose; Angola 8 Ay 6. o o eme ersed LE —————— советники 19 i ate; Lem 0.5—0.9 mm., disc chalky pruinose; parath rk, gen m reditus e L. litbagoga Qi) Vainio 17. Тыа ра1е С mendis mm.; disc black; parathecium and doen r fus enya Log eid Steiner 17. Thallus yellow : ashy; disc black; parathecium hyaline........................ ccce 18 18, oo гей: apothecia 0.15—0.25 mm.; ascospores mi X 45 L. angolensis Mall. Arg. 18. Thallus а not black-margined; apothecia 0.3 (0.4) mm.; mc 9-13 X 5-65 и; angolensis v. orientalis Steiner 19. Thallus « squamu the e margin, gray or nA Central A 3d. а 20 19. Thallus prse gram, yellowish; و‎ Webs 0.13—0.3 mm., disc — Lorie um entire, fuscous es 8—9 X 6-7 и; Kenya binula Müll. Arg. 19. Thallus fes reed, priced pale glaucous; apot chine ia 0.5 mm., disc та ое entire, ini but top of thecium lectus ascospores 11-13 X 8-9 и; Ke L. tra сэнса Mill. Arg ntt — ÀÀ 19. н ا‎ reee disappearing 8 × 3-4 u EM Jganda Js Cagnii s ss т ان‎ 5-15 X 5,5 ; Ru І. argillicola Lindau 21. Apothecia white-pruinose e Os ae -— black; ascospore size not er: Abyssinia L. bogosensis Zahlbr. И .L. leptobola Ny. LECIDEA GRANIFERA Vainio, Cat. Welwitsch Afric. Pl. 2:424. 1901. Lecanora granifera Ach., Syn. Lich. 164. 1814. Type: Guinea, corticole, Afzelius. Thallus crustose, verrucose, 100—125 p. thick, vetiver green; cortex 10-12 p 21. Apothecia black, 0.5-0.7 mm.; ascospores 8-11 Ж 2.5-3 д; Angola. 1953] DODGE—LICHENS OF TROPICAL AFRICA 355 thick, gelified, amorphous; algae T'rebouxia, cells subspherical, 5—6 м in diameter, closely packed in spherical colonies about 20 p in diameter; medulla of thick-walled hyphae 7-8 p in diameter, very loosely woven in the larger warts; cortex finally disappearing and the colonies of algae escaping as soredia, not in definite soralia. Apothecia biatorine, round at first, becoming very irregular, sublobate from un- equal expansion of the parathecium, margin white, disc benzo brown to fuscous, slightly pruinose when young, partly immersed to sessile оп thalline warts, the thallus extending nearly to the top of the parathecium when young, forming a thin pseudo-amphithecium, but covering only the base of the parathecium when mature; parathecium hyaline, 370 y thick in the center, thinning to the slightly elevated margin, slightly inflexed when dry and disc slightly concave to plane, when moist the parathecium is almost level with the slightly convex disc, com- posed of radiating pseudoparenchyma, highly gelified, the upper 40 u under the hypothecium fuliginous, the rest hyaline; thecium 95 р tall; hypothecium not clearly differentiated; paraphyses conglutinate, slender, once or twice dichotomous above the asci, tips not thickened in the slightly brownish epithecial gel; asci 8- spored, clavate, tip thickened when young, 55 X 8 p; ascospores unicellular, monostichous, subspherical, 7—8 р in diameter. Due to the deep fuliginous color which extends from the upper part of the parathecium through the hypothecium shading off gradually through the lower 25 u of the thecium, it is very difficult to see how much is hypothecium and its structure. The systematic position of this species is somewhat uncertain. In the young stages, the sides of the parathecium are nearly covered by thallus and very rarely one can find an algal cell which has penetrated the outer side of the para- thecium, so that one might be justified in following Acharius and leaving the species in Lecanora. On the other hand, the very highly developed, biatorine parathecium and lack of algal layer or medulla of an amphithecium in the mature apothecium suggest a much closer relation to Lecidea sect. Biatora where it was placed by Vainio. This is a very different situation from the ambiguous species between Rinodina and Buellia where a true amphithecium is formed, then the algae die and the amphithecium blackens but leaves traces of medulla and lacunae where the algae have disappeared. Vainio recognized two varieties: lecanoroides, the typical form, and leucotropoides, with a testaceous disc, which he later raised to specific rank. I have seen no material referable to the latter. Nigeria: Ina near Ibadan, on Theobroma, C. A. Thorold 151a pro parte, 156; Ondo Province, Owena and Aponmu near Akure on Theobroma, C. A. Thorold 157, 158, respectively. LECIDEA TENUIS Müll. Arg., Linnaea 63:34. 1880. Type: Angola, Quillu River, growing over sterile Chiodecton, Pechuel-Loesche. Thallus smooth to minutely verrucose, deep olive buff, not black-margined, 80 р thick; cortex 6-8 и thick, decomposed, of periclinal hyphae; algae Trebouxia?, in discrete cylindric colonies 25—40 и in diameter, cells 4—8 м in diameter; medulla [Vor. 40, 356 ANNALS ОЕ THE MISSOURI BOTANICAL GARDEN scarcely differentiated. Apothecia 0.4—0.5 mm. in diameter, margin white, disc waxy, plane at first, becoming very convex, warm buff; parathecium 180 y thick below the hypothecium, thinning to 40 y or less at the top of the thecium, hyaline, the inner 100 р of gelified, thick-walled, periclinal hyphae, the outer 80 p of radiating, septate hyphae 6 и in diameter, forming а pseudoparenchyma; hypo- thecium 30 м thick, of slender, more deeply staining, vertical hyphae, not sharply differentiated from the thecium; thecium 40 р tall; paraphyses slender, once or twice dichotomous above the asci, tips not thickened in the very pale brownish epithecial gel; asci cylindric-clavate, 8-spored, 30 X 6 p; ascospores hyaline, unicellular, long-ellipsoid, thin-walled, 6—7 X 2-3 p. Our material differs in some respects from Müller Argau’s description. The apothecia are more strongly convex at maturity, the parathecium (hypothecium of Müller Argau) is not slightly yellowish below, and the ascospores are slightly shorter and broader (Müller Argau states: 8—11 X 2-2.5 д). Г. micros permoides Müll. Arg., from Usambara, has a chalky thallus, larger apothecia, а darker disc and ascospores 7-9 X 2-3 p. Nigeria: Ondo, on Theobroma, C. A. Thorold 110. LECIDEA (Biatora) nigeriensis Dodge, sp. nov. Type: Nigeria, Ekialodor near Benin City, on Theobroma, C. A. Thorold 135. Thallus epiphloeodes, laevis vel subrugulosus, nigro-marginatus, obscure olivaceo-alutaceus, 80 p crassitudine; cortex vix evolutus; algae protococcoideae, cellulis 6-8 р diametro; medulla nulla. Apothecia 0.5—0.6 mm. diametro, margine thallo concolore, disco plano, subpruinoso, brunneo, linea angusta nigra circumdato; parathecium hyalinum, 110—120 м crassitudine sub hypothecio, superne ad 40 p tenuescens, pseudoparenchymaticum, cellulis 5-8 и diametro, radialiter elongatis; hypothecium 20 м crassitudine, hyphis subbrunneis, dense contextum; thecium 95 и altitudine; paraphyses tenues, semel bisve dichotome ramosae super ascos, apicibus non incrassatis; asci clavati, 80 Ж 8 p; ascosporae octonae, inbricatim monostichae vel subdistichae, hyalinae, ellipsoideae, uniloculares (false biloculares juventute), 8-11 X 4-5.5 y. Thallus smooth to slightly wrinkled, black-margined, deep olive buff, 80 № thick; cortex not differentiated; algae protococcoid, cells 6—8 u in diameter, filling the whole thallus but more densely packed above. Apothecia 0.5-0.6 mm. m diameter, margin concolorous with the thallus, disc plane, slightly pruinose, buck- thorn brown, surrounded by a very narrow black line due to blackening of the top of the parathecium next the thecium; parathecium hyaline, 110—120 p thick below the hypothecium, thinning to 40 p at the top of the thecium, pseudo- parenchymatous, cells 5-8 и in diameter, elongated radially; hypothecium 20 р thick, of densely woven, slightly brownish hyphae; thecium 95 p tall; paraphyses slender, once or twice dichotomous above the asci, tips not enlarged in the very pale brownish epithecial gel; asci clavate, 8-spored, 80 8 и; ascospores im- bricately monostichous to subdistichous, hyaline, ellipsoidal, unilocular (falsely bilocular when young), 8-11 Х 4-5.5 p. 1953] DODGE—LICHENS OF TROPICAL AFRICA 337 LECIDEA RUBINA Ach., Syn. Lich. 40. 1814. Biatora rubina Bél., Voy. Ind. Orient. 2:Crypt.:128. 1846. Type: Guinea, on bark of Uvaria etc., Afzelius. Thallus smooth, somewhat rimose, citrine drab, 55 м thick; cortex 8 p thick, decomposed, of periclinal hyphae; algal layer 20—25 р thick, of rounded colonies of Trebouxia, cells about 5 р in diameter, closely packed; medulla of densely woven, subvertical hyphae, penetrating between the cork cells and disorganizing them to a depth of 40 и. Apothecia 0.7-1 mm., round, sessile, margin concolorous with the thallus, disc plane then convex, isabelline, darkening; parathecium 105 p below the hypothecium, thinning to 55 y at the level of the thecium, the inner half deep fuliginous, the outer half hyaline, of thick-walled pseudoparenchyma, cells 5—6 p in diameter, elongated radially; hypothecium about 15 р thick, not sharply differentiated; thecium 95 p tall; paraphyses coherent, gelified, once or twice dichotomous above the asci, ending in the slightly brownish epithecial gel; asci clavate, 8-spored, 65 X 12 p, wall and tip thickened when young; ascospores imbricately monostichous to subdistichous, hyaline, unilocular, ellipsoidal, 11—12 X 5-6 p. After the ascospores are shed, new apothecia proliferate from the margins of the disc and become partly confluent, resulting in lobed and distorted apothecia. Thorold 131 has the fuliginous portion of the parathecium 135 p thick in the center and the hyaline portion 55 р» otherwise it is similar to the other specimens. Nigeria: Ife near Ibadan, on Theobroma, C. A. Thorold 152; Ondo Province, Aponmu near Akure, on Theobroma, C. A. Thorold 153; Igede near Ado Ekiti, on Theobroma, C. A. Thorold 154; Ondo, on Theobroma, C. A. Thorold 109; Ado Ekiti near Ondo, on Theobroma, C. A. Thorold 131. BACIDIA Bacipia DNtrs., Giorn. Bot. Ital. 1:189. 1846. Type: Not designated, based on Lecidea rosella Ach. and L. carnea Ach. Since L. carnea Ach. has been transferred elsewhere, we may accept L. rosella Ach. as the type, as this would conserve the genus in its sense commonly used. The synonymy of this very large and variable genus is very confused, hence I have not cited the various synonyms. In our collections there are only two species, one in sect. Weitenwebera Zahlbr. (Bilimbia DNtrs. non Reichb., Weitenwebera Opiz поп Schrank) with fusiform ascospores, and one in sect. Eubacidia Zahlbr. with acicular ascospores. Аз these two sections are well represented in tropical Africa, I have prepared keys. Thallus crustose, homoeomerous or heteromerous; ecorticate or cortex poorly developed; algae protococcoid. Apothecia sessile, rarely somewhat immersed or almost stipitate, with a flat or very convex disc; parathecium light-colored; hypo- thecium hyaline or darkening; asci usually 8-spored; ascospores hyaline, long-fusi- form to acicular, 3—many-septate, walls and septa thin, protoplasts cylindric, straight, curved, or helical. Spermatia exobasidial, acicular, straight or curved. [Vor. 40, 358 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. Ascospores vg freue to fusiform, not over 25 u long; sect. WEITENWEBERA (BILIMBIA tr et ct) 2 т. г кзз оуег 25 са long, usually not over 3 mw in diameter; sect. EUBACIDIA............ 10 2. Asc i sepes e ш long, 4— Le ular 3 Asc ^ [pe ng, 4-loc 7 s less than 14 бао Е ea testaceous, ascospores Е, 14-17 X 4—5 p; Sao Thomé...... Catillaria Ferd NDS ез 3. Apothecia finally black or nearly so 4. Ascospores 4-locular, 14-17 X 4—5 и 4. Ascospores 6—8-locular, 17—22 u long 5. Thallus white, disc black; $80 TA ns (Nyl.) Za ктен 5 NNN № . 13. 13. lg ashy white, disc olive black; Usambara B йа го, Arg.) Zahlbr. ypothecium hyaline; ascospores MN X 5-6 р, disc fuscous; Abyssinia................ B ب‎ “Май rg.) Zahlbr. 6. Hypothecium fuscous; песе 18 X 3 д; disc black; Uganda....B. Scottii T a Thallus white, somewhat farina Thallus уе lowish Thallus green; apothecia 0.15—0.25 mm., disc Пер ne EN IS eer (Vainio) Zahl 8. Apothecia black, acter tel 10-12 X 3.5 м; São Th vagula (Nyl.) Za bbe 8. Apothecia rose fleshcolor; ascospores 8-12 X 2. 3-35 5 м де. B. farinulenta s Arg) Zahlbr. Thallus yellowish ashy; apothecia 0.3—0.5 mm., disc and hypothecium fuscous; Ango sublecanorina (Mall. reu Zahlbr. Thallus pale porem apothecia 0.5—0.8 mm., disc arana orange; Ариан hyaline; Nigeri: nigeriensis Dodge 10. Disc vineis -— VA hypothecium fuscous; ES. 27- 34 X 5—5.5 м, 6—8-locular; Central Africa B. griseoalba Lindau 10. Disc deh саг to КЫ at least ena Vul НЯ Ба 20: Disc кий ЫЕ sy a) es ee ee ааа ааа 14 . Hy SES rufous becoming fuscous; ascospores 62-70 X 2-4 ш; Кепуа..............------ araneosa (Hue) Zahlbr. . Hypothecium deep yellow; ascospores 40-65 X 2.5-3 и, 8—10-locular; Kenya.............— submillegrana x . Hypothecium hyaline (sometimes slightly yellowish in B. migrocincta).............————— 12. — 45—55 X 3.5-4.5 ш; apothecia 0.5-1 mm., disc flesh color; hypothallus ; Usambara nigrocincta (Müll. Arg.) a 12. тнт lei than 42 р lohg LS a a e arier r Ascospores 25-30 X 1.5 u; did 0.4—0.6 mm.; mm тер olive; Usambara........ richosperm fie Are) Zahlbr. Ascospores 28—32 X 2.5 u; apothecia 0.25—0.4 mm.; ын ыкы Ascens a (Mill. Arg) Zahlbr. Ascospores 26—42 X 2.5-3 ш; apothecia 0.5 mm.; thallus ва арча € Е сана s (Vainio) Zahlbr. о) . нир 38—41 X 4 u; thallus ochraceous; Abyssinia............ B. Ko ahi “(Krmphbr.) ME Hypothecium tawny fuscous to rufous; ferui EUR or glaucescent..........——-— F Hypothecium — нае ow; thecium 75—80 4; аѕсоѕрогеѕ 72—80 x3 locular; Usam B. endoleucella вч Butte тол 14. Hypothecium —€€ а Ascospores 34-38 X 2-2.5 и, 4—6-locular; apothecia 0.7-1 mm.; Guinea (Togoland) B infus сє (Müll. Arg.) Zahlbr. Ascospores 42—58 X 3.5-4 и, 8—9-locular; apothecia 0.8-1 mm.; An B. [лы Т (Vainio) Zahlbr. Ascospores 62 X 3-3.5 и, 10—14-locular; apothecia 0.5-1 mm.; SOCOtra.----------------------- B. socotrana (Müll. А) Zahlbr. g.) Zahlbr. 16. Ascospores 46-54 X 2-3 и; Angola ый Vainio 16. Ascospores 65—75 X 3—4 u; São Thom B Aber Riy1.) Zahlbr. 16. Азсозрогез 36—56 X 2-3.5 m, rege Angola i ..B. beteroloma v. elongata (Vainio) Zahlbr. 16. Азсозрогез 17—28 X 3.5 и; thecium 55—60 hid Аара OA се eee B. heteroloma v. bacteriospora (Vainio) Zahlbr. 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 359 Bacipia (WEITENWEBERA) nigeriensis Dodge, sp. nov. Type: Nigeria, Ondo Province, Owena, on Theobroma, С. A. Thorold 111. Thallus laevis vel minute verruculosus, pallide flavo-olivaceus, 40-80 y crassi- tudine; cortex vix evolutus; stratum algarum 25 y crassitudine, cellulis proto- coccoideis 5—6 ш diametro; medulla 15-55 д crassitudine, hyphis 3 и diametro, laxe intertextis. Apothecia sessilia, 0.5—0.8 mm. diametro, margine albo, disco ceraceo, ochraceo-aurantiaco, madefacto albo; parathecium hyalinum, inferne 55 џи crassitudine, superne ad 25 y tenuescens, pseudoparenchymaticum, dimidia parte exteriore hyphis radiantibus 5—6 и diametro, cellulis terminalibus rotundatis major- ibusque, dimidia parte interiore hyphis periclinalibus 3—4 и diametro; hypothecium са. 15 p crassitudine; thecium 55 p altitudine; paraphyses tenues, liberae, 2.5 p diametro, pachydermeae, apicibus non incrassatis; asci clavati, 30 X 6 и; ascosporae octonae, ellipsoideae vel subfusiformes, hyalinae, leptodermeae, 4-loculares, 9—12 X 2.5-3 p Thallus smooth to minutely verrucose, 40-80 p thick, light yellowish olive; cortex not well differentiated; algal layer 25 д thick, protococcoid cells 5—6 p in diameter, in a compact layer; medulla 15-55 и thick, of loosely woven pls about 3 и in diameter. Apothecia sessile, 0.5-0.8 mm. in diameter, margin white, disc waxy, ochraceous orange becoming almost white when moistened; parathecium hyaline, 55 џи thick below the hypothecium, thinning to 25 р thick above, pseudo- parenchymatous, the outer half of radiating hyphae 5—6 д in diameter, outermost cells rounded and somewhat larger, inner half of mostly periclinal hyphae 3-4 p in diameter; hypothecium about 15 p thick, not sharply differentiated; thecium 55 р tall; paraphyses slender, free, about 2.5 д in diameter, thick-walled, tips not thickened; asci clavate, 8-spored, about 30 X 6 p, tips somewhat thickened when young; ascospores ellipsoid to subfusiform, hyaline, 4-locular, thin-walled, 9—12 X 2.5-3 y It is with some hesitation that I have referred Deighton М4307А to this species, as the parathecium is 95—105 и thick below, thinning to 55 u above, and the ascospores are smaller, 7-9 X 2.5-3 и; otherwise it agrees well with the Nigerian material. Nigeria: Ondo Province, Owena, on Theobroma, C. A. Thorold 111, type; Ikere near Ondo, on Theobroma, C. A. Thorold 112. Sierra Leone: Njala (Kori), on twigs of Bauhinia tomentosa, Е. C. Deighton M4307A Васила (EUBACIDIA) GOLUNGENSIS (Vainio) Zahlbr., Cat. Lich. Univ. 4:202. 1926 Lecidea golungensis Vainio, Cat. Welwitsch Afric. Pl. 2:419. 1901. Type: Angola, Golungo Alto, near Sange, 320-775 m., Welwitsch 135a, 136 as v. pauciseptata, 149 p.p. as v. pluriseptata; Serra de Alta Queta, Welwitsch 150c, 151 as v. pauciseptata, 154 as v. pluriseptata. Мо specimen cited. as type of the species (see discussion below). [Vor. 40, 360 ANNALS OF THE MISSOURI BOTANICAL GARDEN Thallus granular-verrucose, continuous, citrine drab, 40 u thick; cortex gelified, 5—6 p thick; algal colonies discrete, 15—20 и in diameter, of Trebouxia, closely packed cells 3—4 u in diameter; medulla 14—19 y thick, gelified, of densely woven hyphae. Apothecia sessile, margin very thin, soon immarginate, 0.5 mm. in diam- eter, sometimes concrescent in groups up to 1 mm. in diameter, disc avellaneous to wood brown; parathecium 80-150 м thick below the hypothecium, thinning to 25 р at the top of the thecium, hyaline, inner half of slender, periclinal gelified hyphae, the outer half of radiating, thick-walled, conglutinate hyphae 5—6 y in diameter, occasionally penetrated by a few algal cells; hypothecium 15 и thick, of densely woven, mostly periclinal, slender hyphae; thecium 55-70 y tall; para- physes slender, coherent, once or twice dichotomous above the asci, tips clavate in the slightly brownish epithecial gel; asci cylindric-clavate, 8-spored, about 45 X 8 p; ascospores acicular, fascicled, 3—6-septate, 26—42 X 2.5—3 и. Spermo- gonia spherical, half emersed from the thallus at the edge of the parathecium, about 80 p in diameter, wall of exposed portion blackened to a depth of 15 p, rest hyaline, filamentous, 15 и thick; spermatiophores flask-shaped, about 10 X 2 y; spermatia bacilliform, straight, about 6 X 1 y. Vainio separated this species into two varieties, pauciseptata, with the thecium 55—65 p tall, ascospores 4-locular, 26—36 p long, and var. pluriseptata, with the thecium 60-70 р tall, ascospores 5—7-locular, 38—42 p long. In my material, using gentle pressure on the cover glass, I have found free ascospores 32 and 40 p long up to 8-locular. Nigeria: Ina near Ibadan, on Theobroma, C. A. Thorold 151; Ondo Province, Aponmu near Akure, on Theobroma, C. A. Thorold 172b. ВАСИЛА HETEROLOMA Zahlbr., Cat. Lich. Univ. 4:204. 1926. Lecidea beteroloma Vainio, Cat. Welwitsch Afric. Pl. 2:418. 1901. Type: Angola, Golungo Alto, near Sange, 360—775 m., corticole, Welwitsch 125, 148, 149. Thallus slightly verrucose, uneven and rimose, margin pale, indefinite on bark, pale olive buff, 65-80 р thick; cortex 10—12 p thick, of slender thin-walled peri- clinal hyphae, the outer ones somewhat decomposed; algae Trebouxia, filling the rest of the thallus, cells about 6 и in diameter; medullary hyphae partly disinte- grating bark cells to a depth of 140 и. Apothecia sessile, constricted at the base, round or somewhat distorted by mutual pressure, 0.5—0.7 mm. in diameter or in groups 1 X 2 mm.; margin prominent, pale ashy or brownish becoming nearly black in age, disc concave to plane, fuscous black; parathecium sulphate green when first sectioned soon fading in lacto-phenol, 140 p thick below the hypo- thecium, thinning to 65 p at the top of the thecium, of radiating pseudoparen- chyma, cells 6-8 м in diameter, thick-walled; hypothecium probably hyaline (pale brown in my moribund specimens), about 25 р thick, of closely woven, slender 1953] DODGE—LICHENS OF TROPICAL AFRICA 361 hyphae; thecium 60 y tall; paraphyses slender, dichotomous above the asci, tips not thickened in the pale brownish epithecial gel; asci long-ellipsoid, 8-spored, 45 X 6 р; ascospores hyaline, acicular, plurilocular, 35 X 2.5 p. My material is moribund, and much of the thecium has begun to disintegrate. The one ascus near the parathecium clearly seen is probably somewhat immature, as I could not see the septa of the ascospores clearly enough to count them and I found no free ascospores. Vainio describes the ascospores as 36—56 X 2—3.5 p, 6-10-locular. He described two varieties: v. elongata which should be taken as his varietas typica since he repeats the dimensions of the ascospores for it, and у. bacteriospora with paler apothecial margins and smaller ascospores 17—28 Х 3—3.5 p. Sierra Leone: Njala (Kori), on Phyllanthus discoideus, Е. C. Deighton M4581. LOPADIUM LoPADIUM Koerb., Syst. Lich. Germ. 210. 1854. Heterotbecium Mont. in Gay, Hist. Fís. Polít. Chile, Bot. 8:175. 1852, mon Flotow, 1850. Brigantiaea Trev., Sihet e Paglie 7. 1853. Туре: Г. pezizoideum (Ach.) Koerb. Heterothecium Mont., non Fw., was based on H. Berteroanum Mont. Brigantiaea Trev. was based on Heterotbecium Berteroanum Mont., B. Mariae Trev., B. tricolor 'Trev., B. tristis Trev., and B. argentea Trev. When Lopadium sensu latiore is monographed, it seems probable that Brigantiaea Trev. will be recognized as a separate genus, and there will be no need to conserve Lopadium Koerb. Thallus crustose, ecorticate or nearly so; algae protococcoid. Apothecia sessile; parathecium fleshy, cartilaginous, and light colored, or carbonaceous; hypothecium hyaline to dark colored; asci 1—8- spored; ascospores hyaline, muriform, thin- walled, without a halo; spermatia ellipsoidal, short, straight. _ = Palticoloas аа анн ане Corticolous; asci monosporous, rarely 2—3-spored.............. 2.' Asci 8-spored; ascospores 18-19 X 8-9 м, PEE a слана disc black 620 Thomé ш... Ен г riques) Sant. 2. Asci 1- d; ascospores 65—85 X 16—23 p, disc "pim Pipes colo pins te cocco т у L ger rt Мек БЫШ Тыш ыс сы зше о миа Thallus glaucous; disc testaceo-fuscous, margin pale; ascospores 60-73 X 20-24 golà. ee EON inicr Lecidea olivacea v. ju ndi Vainio Thallus olive buff or darker ...............—— = ет : IU ыы 5 es; ascospores 4. Disc watery fuscous, margin с e à —— uis s Mi. Ar. а x 19-21 p; ee са sepia — . ww в > . асеит Dodge ro Leone elf pceksccdednasnescsasssoeeresososooseces: MEE g РА © B a 8 Е б. 4 я E $ g B 8 [2 2 E 8 ч b © х $ © A О 26 5 a F ° n, 5 о. 3 1 á p X © = à i E 8 > | = A = Vlerm T4008. о Шы a ——- Lopapium sepiaceum Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on Baubinia tomentosa, F. C. Deighton M4307B. [Vor. 40, 362 ANNALS OF THE MISSOURI BOTANICAL GARDEN Thallus verruculosus, margine pallido, 80 м crassitudine, albidus; cortex vix evolutus; algae protococcoideae, cellulis 5—6 м diametro. Apothecia sessilia, basi constricta, margine subelevata, thallo concolore, subpruinosa, disco sepiaceo; para- thecium inferne 55 р, superne ad 15 р tenuescens, hyphis pachydermeis radiantibus, luminibus 4 р diametro, hyalinum, extus brunnescens; hypothecium brunneum, centro 55 p ad marginem 15 y tenuescens, hyphis 4—5 и diametro; thecium 105- 110 м altitudine; paraphyses super ascos dichotome ramosae, apicibus non in- crassatis; asci pachydermei, cylindrici, 95 X 20-24 р; ascosporae binae vel ternae, muriformes, pallide brunneae, ellipsoideae, 32—43 (—60) X 19-21 y. Thallus whitish, minutely verrucose, margin pale, 80 м thick; cortex not well differentiated; algae protococcoid, filling the whole thallus, but not closely packed, cells 5—6 p іп diameter. Apothecia sessile, constricted at the base, margin slightly elevated, concolorous with the thallus, slightly pruinose, disc sepia; parathecium 55 u thick below, thinning to 15 y above, of thick-walled radiating hyphae, lumen about 4 д in diameter, hyaline, brownish on the outside; hypothecium deep brown, 55 p thick in the center, thinning to 15 p at the margin, of hyphae 4—5 y in diameter; thecium 105—110 џ tall; paraphyses dichotomous above the asci, tips not thickened in the brown epithecial gel; asci thick-walled, cylindric, 2—3-spored, 95 Х 20-24 p; ascospores monostichous, muriform, pale brownish, without a halo, ellipsoid, 32—43 (—60) X 19-21 p. Lopapium Deightoni Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on Bauhinia tomentosa, Е. C. Deighton M4307C. Thallus minute verruculosus, margine pallido, 65 ш crassitudine, obscure olivaceo-alutaceus; cortex vix evolutus, са. 8 и crassitudine, hyphis periclinalibus; stratum algarum 25 p crassitudine, continuum, cellulis protococcoideis, 5—6 p diametro, laxe dispositis; medulla 30-35 p crassitudine, yphis dense contexta. Apothecia 0.6—0.8 mm. diametro, basi constricta, margine subelevato, disco con- colore, pruinoso; discus isabellinus, planus, granuloso-pruinosus; parathecium inferne 65 p crassitudine, superne ad 30 p. tenuescens, hyalinum, pseudoparenchy- maticum, cellulis radiantibus; hypothecium centro 40 ш crassitudine, ad marginem 15 р tenuescens, hyphis periclinalibus dense contextum; thecium 130 y altitudine, superficie inaequali; paraphyses pachydermeae, super ascos dichotome ramosae, cellula apicali subsphaerica, 5—6 и diametro; asci cylindrico-clavati, pachydermei, 120 X 27 p; ascosporae solitariae vel rare binae, ellipsoideae, hyalinae vel sub- brunneae, sine halone, muriformes, 61-88 X 16-20 p (solitariae) vel 50—60 Х 16 и (binae). Thallus minutely verruculose, margin pale, 65 u thick, deep olive buff; cortex scarcely developed, about 8 р thick, of periclinal hyphae; algal layer continuous, protococcoid, cells 5—6 д in diameter, loosely packed; medulla 30—35 д thick, of densely woven hyphae. Apothecia 0.6-0.8 mm. in diameter, constricted at the base, margin somewhat elevated, pruinose, colored like the disc which is plane, isabelline, covered by a coarse granular pruina; parathecium 65 и thick below, 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 363 thinning to 30 д above, hyaline, pseudoparenchymatous, cells radiating; hypo- thecium 40 p thick in the center, of closely woven, periclinal hyphae; thecium 130 p tall, surface uneven; paraphyses thick-walled, dichotomously branc ed above the asci, the terminal cells subspherical, 5—6 џи in diameter; asci cylindric- clavate, thick-walled, 1—2-spored, 120 X 27 p; ascospores ellipsoid, hyaline or slightly brownish when old but without a halo, muriform, 61-88 X 16-20 «u when single, 50—60 16 u when two per ascus. Deighton 4344 is older than the type and the apothecia have begun to dis- integrate, but the structure is the same and the spores are of the same size. The thallus bears abundant campylidia, otiform, conchiform or cyphelliform, cinnamon buff, about 1 mm. tall and broad; stalk 125 p tall, 250 р in diameter with hairs 40 p long; context 120-135 м thick, the outer 15 р forming a fine tomentum of curved, brownish, thick-walled hairs 5—6 р in diameter, with protococcoid algae penetrating in an irregular layer 80—90 и thick, the rest of conglutinate, densely woven, thick-walled, hyaline hyphae 6 p in diameter; subhymenium about 15 p thick, of slender vertical hyphae covering the inner (proximal) portion, leaving the distal portion of the cup sterile; conidiophores somewhat flexuous, 30 X 3 p, with 1—2 short sterigmata; conidia 16 X 2 p, very thin-walled, slightly curved, unicellular. The campylidium has often been considered a parasitic fungus and described as a species of Cyphella or Chlorocyphella, but the layer of protococcoid algae is similar to those of the species of Lopadium on which it develops, and continuous with the algal layer of the thallus. This seems to indicate that it is a genuine conidial state of the fungus component of Lopadium, as originally described by Müller Argau. Frequently I have seen this conidial state on various species of Lopadium and on the foliicolous Sporopodium from Texas southward in tropical America. Sierra Leone: Njala (Kori), on Bauhinia tomentosa, Е. С. Deighton M4307C, type; on Lagerstroemia speciosa, F. C. Deighton M4344. Lopapium nigrobrunneum Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on twigs of Lagerstroemia speciosa, F. C. Deighton M4346. allus verruculosus subrimulosusque, in hypothallo nigro, 65 p crassitudine, olivaceo-alutaceus; cortex vix evolutus; stratum algarum 20-25 р crassitudine, protococcoideum, cellulis 6-7 м diametro; medulla 25-30 р crassitudine, hyphis 3-4 р diametro laxe intertextis; hypothallus 15 н crassitudine hyphis periclinalibus brunneis. Apothecia 0.5—0.7 mm. diametro, subconvexa, basi constricta, margine et disco nigricantibus; parathecium inferne 40 p crassitudine, superne ad В р tenuescens, pseudoparenchymaticum, hyalinum, hyphis radiantibus, cellulis 6-7 p diametro; hypothecium obscure brunneum, centro 40 p crassitudine ad marginem 20 р tenuescens; thecium 100—105 p altitudine, superficie inaequali; paraphyses conglutinatae, pachydermeae, super ascos dichotome ramosae, apicibus non in- [Vor. 40, 364 ANNALS OF THE MISSOURI BOTANICAL GARDEN crassatis; asci cylindrico-clavati, ca. 100 X 30 д; ascosporae solitariae vel binae, hyalinae vel subbrunneae, sine halone, muriformes, 43—56 (—66) X 16-21 (-23) p. Thallus minutely verrucose and subrimulose, on a black hypothallus, olive buff, 65 и thick; cortex scarcely differentiated; algal layer 20—25 р thick, proto- coccoid, cells 6-7 и in diameter; medulla 25—30 y thick, of somewhat loosely woven hyphae 3—4 и in diameter; hypothallus 15 и thick, of periclinal brown hyphae. Apothecia 0.5-0.7 mm. in diameter, slightly convex, constricted at the base, margin and disc clove brown, blackening; parathecium 40 и thick below, thinning to 15 р at the slightly inflexed top, НЕ of radiating hyphae, cells 6-7 м in diameter; hypothecium very dark brown, 40 y thick in the center, thinning to 20 p at the margin; thecium 100—105 м tall, surface very uneven; paraphyses conglutinate, thick-walled, dichotomous above the asci, tips not enlarged in the brownish, epithecial gel, filled with minute granules; asci 1 (—2)-spored, cylindric-clavate, about 100 > 30 р; ascospores hyaline to slightly brownish, without а halo, muriform, 43—56 (-66) X 16-21 (-23) p. PHYLLOPSORACEAE Thallus squamulose to subfoliose, often somewhat erect, attached to the sub- strate by rhizinae; upper cortex well developed; algae protococcoid. Apothecia rounded, sessile; amphithecium absent, parathecium light or dark; paraphyses un- branched; ascospores hyaline, unilocular or septate. Although there is no clear differentiation of primary thallus and podetia which one finds in the Cladoniaceae, this family seems much closer to it than to the Lecideaceae. Only two genera are known: Phyllopsora with unilocular ascospores, and Psorella with 4—16-locular ascospores. Our material belongs in the former genus. PHYLLOPSORA Рнутторзовл Müll. Arg., Bull. Herb. Boissier 2:append. 1:11, 45. 1894. ?Psoromidium Stirton, Proc. Phil. Soc. Glasgow 10:304. 1877. Туре: not designated but P. parvifolia (Pers.) Müll. Arg., one of the best- known species, may be chosen. Psoromidium Stirton was based on P. Wellingtonii Stirton. Characters of the family: ascospores unilocular, long-ellipsoid, hyaline. 1. Squamule not long white-ciliste: fertile ори ile 1. Squamules long, snowy-white ciliate, upper surface pale yellowish; sterile; re Pune и P. pannosa ‘a Müll. Arg. P A E P. Buettneri (Müll. Arg.) Zahlbr. 2. Ascospores relatively short ellipsoid, 5-7.5 X 3.5-5 p; Usambafa cic ا‎ ОЕ P. brachyspora Müll. Arg. PHYLLOPSORA BuETTNERI Zahlbr., Cat. Lich. Univ. 4:396. 1926. Psora Buettneri Müll. Arg., Bot. Jahrb. [Engler] 15:506. 1893. Lecidea Buettneri Stzbgr., Ber. Thatigk. St. Gall. Naturw. kis 1893/4:246. 1895. 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 365 Type: Togoland, Bismarcksburg, corticole, R. Buettner. Hypothallus fuscous black, almost completely covered by light yellowish olive squamules, imbricate, less than 1 mm. broad, deeply lobed, the lobes becoming erect isidia, so that the whole thallus appears isidiose and minutely granular- sorediose. Apothecia convex, immarginate, constricted below into a substipitate base, 0.35 mm. broad, 0.25 mm. tall, disc cinnamon rufous; parathecium of hyaline, thick-walled, densely woven hyphae 3-4 и in diameter, containing а few small colonies of Trebouxia cells 6-8 м in diameter; hypothecium 70 p thick, of thick- walled, vertical hyphae, very dark brown next the thecium, shading to hyaline next the base; thecium 25 y tall; paraphyses conglutinate, tips not thickened, some- what dichotomous in the upper portion; asci cylindric, 8-spored, about 20 X 8 р; ascospores fascicled, unicellular, long-ellipsoid, 10-12 X 2.5-3.5 p. Some ascospores contain 3—5 large oil droplets, when the ascospores appear 3—5-septate and might lead one to look for the species in Psoretta. Gold Coast: Tafo, West African Research Institute, on Theobroma, C. A. Thorold 155. Nigeria: Ondo Province, Aponmu near Akure, on Theobroma, C. A. Thorold 123. LECANORACEAE Thallus crustose, uniform or with effigurate margins, rarely dwarf-fruticose, branched, attached to the substrate by hyphae of the hypothallus or of the medulla, without rhizinae; heteromerous (except in Harpidium); ecorticate or corticate; algae Protococcus or Trebouxia. Apothecia immersed in the thallus, to sessile, round; amphithecium well developed; parathecium very poorly developed or lack- ing; hypothecium hyaline or brownish, usually with algae below it; paraphyses unbranched (except in Ochrolechia and Phlyctidia) ; asci 8—32-spored; ascospores hyaline (brownish in some species of Myxodictyon), unilocular, septate or muri- Thallus homoeomerous, pseudoparenchymatous; ascospores sickle-shaped; known only Lh Eran Fue LL боді Harpidium Koerb. 1. Thallus -heteromerous, . plectenchymatous | ittm 2 je Aicopara unilocullt i cene ennemi egt te 3 5 JAxemote Бак ие ertt ПУ 5 2. Авёоврогев muriform | 1mm iere irn etie e УНЕСИТЕ ЧНТУ 3. Paraphyses unbranched or sparingly dichotomous above the asci............ M ——M— 4 3. Paraphyses branched, sparse in the thecial gel; ascospores large but relatively re acad d [perhaps belongs in the Pertusariaceae]....—————777 olechia reca 4. Thallus not bright yellow to orange; spermatia acicular, straight or Deeper tat >= ; 4. Thallus bright yellow or orange; spermatia ellipsoidal, straight.........- Candelarie! A all. гв. 5. Ascospores ellipsoid to subcylindric; spermatiophores unbranched, spermatia terminal................. а 5. Ascospores fusi ; spermatiophores branc te, rmatia ме ——--;:- 6. Thallus gray or brown; spermatia straight or curved, acicu о n vei 6. allus bright yellow to orange; spermatia ipsoidal, straight....... „Сай " ie г) ü a rg 7. Thallus cranes, und odi. о ee о тте pos opbila 'Trev Y Thallus disc 2. 1 o OS о ет enopsora Mass 8. Thallus ecorticate; apothecia immersed а 8. Thallus corticate; apothecia sessile 7777070707077 77g Heiss wc 9. Paraphyses unbranched суусоо a n cea е сатр г. 9. Paraphyses branched and anastomosing.....——7700007 enne Phlyctidia mo ыа 10. Apothecia immersed in the thallus, А» Mass. [Vor. 40, 366 ANNALS OF THE MISSOURI BOTANICAL GARDEN LECANORA Lecanora Ach., Lichenogr. Univ. 77. 1810. Type: not designated. Thallus crustose, uniform, effigurate, rarely squamose or dwarf-fruticose, at- tached to the substrate by the hyphae of the hypothallus or of the medulla, without rhizinae, heteromerous; ecorticate or corticate; algae protococcoid. Apothecia immersed or sessile, circular; amphithecium well developed; parathecium usually absent or poorly developed; hypothecium hyaline or brownish; paraphyses un- branched, free; asci normally 8-spored, rarely 16—32-spored; ascospores hyaline, ellipsoidal to spherical, rarely reniform. Spermatia cylindric to acicular, straight or curved. The above generic description is based on the concept of Zahlbruckner. Of the six sections into which he divided the genus, I would recognize four as distinct genera, leaving Eulecanora and Placodium. Our material belongs in Eulecanora, of which the key to tropical African species follows. — Asci 16-spored; ascospores 8—11 X 6-8 pu, disc pale flesh color; apothecia about 0.7 mm.; thallus yellowish АЕ granular, wrinkled, margin subarachnoid; Kenya L. pleospora Müll. A 1. Asci 8-spored + De white o pale сетш. thallus. white... Leve seed icio asked, sS 3 с ye die to cinnamon or yellowish Е зару Бае 4 3 Di "ODE fah COE IAS ее 7 Dec fuscous. to reddi Fuse ac a es yS 10 * Dise black; at Net when deg о макы ыи И 19 3. Thallus pale gray, margin radiate-rimose with white id. мез apothecia MM angular from mutual pressure; ascospores 9—11.5 X 6-7.5 u; on Euphorbia voa ACE DAS L pototallin Lindau 3 н таи white or пар tartareous, obsoletely чамь, тагріп DEDE apotheci: 3 х 5 аѕсоѕрогеѕ 9—12 -7 м; saxicole; St. Helena.......... L. Sanctae on Mal. Arg. 3: Thallus verruculone, Миы; apothecia 0. он mm.; ascospores 11—13 X 5 и corticole; Angola L Monodore Vainio 4. = allus white to ashy, margin indistinct, verruculose; apothecia 0. e mm.; es 12-16 X 6-8 и; corticole; Kenya L романы Vainio 4. Thallus злее гы margin slightly radiate-fibrillose, smooth; apothecia 0.5—0 s: 9-9 X 6— ji corticole; Мужава и L. Brassii ma 4. Thallus eR. WO: БЕЛИН а ааа олень ИЗ КЕ! 5. Ascospores slender, 13 X 4 u; apothecia 0.7-1.2 mm.; thallus — en: Mozamb В Ао ог carneosul wi — yellow босат to livid, apothecial margin granulose x5 Thallus margin black; Pg ascospores 9—11 Х 6-7 ш; saxicole; Erit L ем Zahlbr. 6. Thallus margin fibrillose, pale; apothecia 15-2 mm.; margin gen p pale yellow; ascospores 12—14 -= m k; — хара: 5 bo ambar aed n ll. Arg. 6. Thallus margin indefinite, e, thallus granulose-verrucose to sq d бие Уди slightly ащ рое АА сіа 0. 71. mm., margin t а er 10. 5 E x и terricole; Rüshda ... 5 7 ани _ 7. Disc pruinóéa IT bs а козлы ed net SS 7. сие not tese Pymes white ie Me glaucous т e - . Disc grayish r apothec : =, terricole; inate A ee profer mm.; ascospores 9.5-11 X 5.8—6 ЛЕ ШЕЙН, Lp dus 8. Disc dr flesh color; apothecia 0.4—0.8 mm.: ; — 9-14 5 corticole; j О iii ef etis v. carneola (Vaini nio) Zahlbr. . Disc livid flesh color, margin granulose; ascospores 9-11 x 6-7 p; saxicole; Eritrea... = І tropoides Zahlbr. Disc flesh-rufous margin very thin; crenulate; asco: icole; Zahl! ^ ; spores 10—12 X 6-7 p; corticole; Angola — T. on once о L. leprosa v. rufocarnea (Vainio) Zahlbr. 1953] DODGE—LICHENS OF TROPICAL AFRICA 367 9. Disc dark fuscous, margin entire; corticole; Mozambique ecoronata Vainio 9. Disc flesh color, margin thin, entire to slightly crenulate; apothecia 0.2—0. i mm.; asco- spores 10-14 X 6-10 п; corticole; Kenya L чан Hue 9. Disc pale rufous, margin thin; apothecia 0.5—0.7 mm.; ascospores 11-13 X 6-7 corticole; French Equatorial Afri L eiae Hue 10. p аа obsolete; apothecia "pen 6 mm.; margin thin, entire; ascospores 10-13 X m ns saxicole Kenya L. subcongruens Müll. M 10. Thalls yellow 10. Thallus white e ashy 13 11. Thallus rimulose, thin; nibii 1-1.5 mm.; medulla yellowish; а ен 9-11 X 55.5 ш; saxicole; Socotra Г. socotrana Müll. Arg. 11. Thallus rimose-areolate, black-margined, sulfur-tartareous, thi а medulla yellowish; apothecia sibi: 0.5 mm.; ascospores 8—9 о бщ; A: имей ару га pedata pue 11. ee рле ог erimus spei margin tumid, entir . Apothec 8(—1.5) mm es 8 X 5.5-6 4; iballos ashy 2 рса drab; pared Belgis Соп beo octialis Stzbgr. a 0.4 mm.; ascospores 8—9 X 4—5 и; thallus ashy yellow to gage saxicole; africana Zahlbr. 12. Apschc 0.2—0.5 mm.; ascospores 10-12 X 4—5 и; thallus ochroleucous; terricole; ran anulescens Jatta 12. Arie 0.5—0.8 mm.; ascospores 10-13 X 6-9 ш; thallus sulfur pn corti- sabulosa Steiner 14 m > T TE cole; Ke 13. Thall n white-byssine argi 13. т тагып black 15 13. Thallus margin inconspicuous (or not described) 17 uous ( я Thallus glebu eem dine apothecial margin flexuous; мено» size corticole; Abys абме у. ти Fw. 14. Lag Pei wrin WAR pada wis apothecia 0.5-0.75 mm.; margin entire ospores 15 X 7 4; corticoles Usambara- -L subfusca v. ЖУЛ ЫП Arg. 15. Thallus aptam to glebulose, ashy t e drab; apothecia 0.8 cm T angular in age; ascospores 8 X 5.5 pu тысе ole; Congo. aequinoctialis Stzbgr. 15. Thallus rimulose, exe apothecia up to 0.5 mm.; ascospores 10—15 = ur p; saxi- cole; Angola (m ong in Aspicilia) L. bomaloplaca Nyl. 15. Thallus ди оѕе-аего liw: apothecia 0.4—0.5 mm.; ascospores 8—10 X 4-5 п; — Lene 15 Thallus ashy to ashy терин ascospores 0-9 x + 4—5 и; Eritrea.............. y africana Zahlbr. allus white; жыл уге 0 X 4.5-5 p; Ken I. encopac Vainio 17. “Thales smoot e or verrucose; чы t 0.6 mm.; наси 10-14 X 6.5-7. T ш; cort vubinité Steiner 17. Thallus gatis е ct apothecia 0.4—0.6 mm.; ascospores 16—20 Х 4 hi ide ole; tri iguttulata жа ritrea 17. Thallus granulose 18. Ascospores 8-12 X 4-6 ц; a ee 0.3-0.6 m xine І, subfusca v. ferax Müll. Ascospores 8—14 X 5-7 и; São Thomé and Ilha Dense a L. tropica Lii. 19. Thallus ee ^ri ashy, lls og 19. „Thallus yellow: Thallus mers white, fibrous; apothecia 0.5 mm.; ascospores 13-18 X 7-8 p corticole; A ngola : x m Arg. 20. E oie margin inconspicuous; apothecia 1.5 mm.; ascospores unknown; corticole; Р a L. subfusca v. melaleuca мап. Arg. re mbara 21. Hyporhecium orange rufous; apothecia 1-1.5 mm.; ascospores 17—20 5—6 ш; corti- ee 84 5 эй L f igrans Müll. Arg. 21. Hyp mA tS ck; ascospores 8—10 X 7-8 и; saxicole; $осокга.................. L. notba Müll. Arg 21. Hypochecium hyaline; apothecia 0.5-1 mm.; not corticole. 22 — saxicole; Usam ncm m dark green; thallus yellow ochre; ascospores orci pear brace Mill in у 11-12 X 65-7 д 22, араса та aoe black; thallus stramineous; ascospores i LA € jt UA Lecanora Brassii, Dodge, sp. nov. Type: Nyasaland, Kasungu Hill, corticole, L. J. Brass 174582. Thallus indeterminatus, 44 р crassitudine, albus, pruinosus, margine incon- spicue radiato-fibrilloso; шаш! algae Trebouxia, cellulis 6 р diametro; [Vor. 40, 368 ANNALS OF THE MISSOURI BOTANICAL GARDEN medulla deest aut ad 15 y crassitudine hyphis subverticalibus magnis cum crystallis hyalinis. Apothecia 0.5—0.6 mm. diametro, margine crasso, elevato, disco pallide flavo-carneo, dense pruinoso; amphithecium 105—120 и crassitudine, cellulis algarum dense dispositis; parathecium deest; hypothecium 40 џи crassitudine, hyalinum, hyphis 3 и diametro dense contextum; thecium 55—60 м altitudine; paraphyses tenues, pachydermeae, apicibus clavatis; asci clavati, juventute apicibus incrassatis, protoplastis submamillatis, 42 Ж 12 р; ascosporae octonae, imbricatim monostichae, dein subdistichae, hyalinae, uniloculares, late ellipsoideae, 8—9 X 6-7 p. Thallus crustose, indeterminate, margin inconspicuously radiate-fibrillose, white, pruinose, not changing color when moistened; cortex not differentiated; algae T'rebouxia, cells 6 p in diameter, in places completely filling the thallus, in others leaving a medulla, 15 u thick, of coarse, closely woven, subvertical hyphae, penetrating between the cork cells, often with large hyaline crystals. Apothecia 0.5—0.6 mm. in diameter, margin thick, elevated, disc pale yellowish flesh colored, densely pruinose; amphithecium 105—120 p thick, of densely packed algae; para- thecium absent; hypothecium 40 u thick, hyaline, of densely woven hyphae 3 p in diameter; thecium 55—60 м tall; paraphyses slender, relatively thick-walled, tips slightly clavate, epithecial gel hyaline with minute crystals (brownish in thick sections); asci clavate, tips thickened when young, protoplasts somewhat mamil- late, 8-spored, about 42 X 12 p; ascospores imbricately monostichous, becoming subdistichous, hyaline, unilocular, broadly ellipsoidal, 8-9 X 6-7 p. LECANORA AEQUINOCTIALIS Stzbgr. Ber. Thitigk. St. Gall Naturw. Ges. 1888/9:218. 1890. Type: Belgian Congo, Banana, at mouth of Congo River, corticole, P. Hesse. Thallus epiphloeodal, verrucose to glebulose, thickness variable, up to 160 р, citrine drab with a narrow black margin; cortex about 8 y thick, gelified, of peri- clinal hyphae, often disappearing on the tops of the glebulae and revealing the sulfur-colored interior, but not truly sorediose; algae protococcoid?, somewhat yellowish green, cells subspherical, 3-5 и in diameter, mostly arranged in vertical rows and filling the smaller warts, while forming a layer about 40 p thick aroun the larger, hollow warts which contain a few short, loosely tangled, subvertical filaments of Stigonema; medulla not differentiated. Apothecia innate in the tops of the glebulae, becoming sessile at maturity, round with an entire margin at first, then becoming lobulate from the confluence of several apothecia with a somewhat crenulate margin, 0.8(—1.5) mm. in diameter, disc plane, warm sepia becoming convex when moist and appearing immarginate; amphithecium 55 р thick, of the same structure as the thallus; parathecium 55 y thick in the center below the hypothecium, thinning to 20 p at the margin, very dark brown (black in thick sections) ; hypothecium not clearly differentiated from the parathecium; thecium about 55 y tall; paraphyses conglutinate, slender, tips not thickened in the brownish epithecial gel; asci clavate, thick-walled when young, 8-spored, about 40 X 8 #5 ascospores hyaline, unilocular, relatively thick-walled, subspherical, 8 X 5.5-6 ^ 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 369 Stizenberger's description is brief, but our material agrees well with the char- acters recorded. igeria: Ondo Province, Owena, on Theobroma, C. A. Thorold 132; Ipetu, on Theobroma, С. A. Thorold 130; Ina near Ibadan, on Theobroma, C. A. Thorold 129 OCHROLECHIA OcHROLECHIA Mass., Ricerch. Autonom. Lich. Crost. 30. 1852. Lecanora sect. Ocbrolecbia Flagey, Mém. Soc. Emul. Doubs, 272. 1886. Type: not designated, based on Verrucaria parella Wigg. and Lichen pallescens L. Thallus crustose, uniform, attached to the substrate by hyphae of the hypo- thallus or of the medulla, without rhizinae, ecorticate or with a cortex of erect or irregularly woven, septate, gelified hyphae; algae protococcoid; medulla of thin- walled hyphae; soredia frequent. Apothecia immersed at first, then sessile, con- stricted at the base; amphithecium well developed; hypothecium hyaline; paraphyses branched and anastomosing in the thecial gel (similar to those of the Pertusari- aceae); asci 2—8-spored; ascospores hyaline, ellipsoidal, unilocular, large. Spermo- gonia in verrucae, cavity often labyrinthiform; spermatiophores unicellular; spermatia long-cylindric, straight. OCHROLECHIA palmicola Dodge, sp. nov. Type: Sierra Leone, Sefadu (Gbense), on trunk of Elaeis guineensis, P. Adames, comm. F. C. Deighton M4750. Thallus epiphloeodes, verrucosus aut rugosus, subrimoso-areolatus, albidus, 300 м crassitudine, laevior continuiorque ad marginem lobatum minute plumoso- fibrillosum; cortex 25 p crassitudine, hyphis conglutinatis periclinalibus 2-2.5 р diametro, inferne tenuioribus intertextisque; stratum algarum 25 p crassitudine, protococcoideum, cellulis 5—6 и diametro; medulla 130 p crassitudine, hyphis 2-5 p diametro laxe intertextis, granulis minutis aut crystallis inspersis, dein stratum alterum inter strata cellularum suberosarum 80-105 p crassitudine, structura simili; soralia elevata, 1 mm. diametro, soredia granularia, pallide sulfurea, 50—60 p diametro. Apothecia sessilia, urceolata, margine integro, 1 mm. diametro, disco vinaceo-alutaceo, albo-pruinoso; amphithecium inferne 125 р crassitudine, superne ad 50 p tenuescens; cortex et stratum algarum ut іп thallo, medulla hyphis hyalinis conglutinatis 4 и diametro; hypothecium 25 y crassitudine, hyphis tenuibus, peri- clinalibus super stratum algarum; thecium 200-210 p altitudine; paraphyses tenues, dichotome ramosae et sparse anastomosantes; asci cylindrico-clavati, 175 X 65 p, pachydermei, usque ad 8 y crassitudine; ascosporae octonae, hyalinae, leptodermeae, protoplastis granularibus, uniloculares, 55 X 24 p. - Thallus epiphloeodal, verrucose or wrinkled, somewhat rimose-aerolate, whitish, smoother and continuous toward the lobed, white, minutely plumose-fibrillose margin, 300 p thick; cortex 25 р thick, of conglutinate, periclinal hyphae 2-2.5 p in diameter, more slender and interwoven below; algal layer 25 р thick, proto- [Vor. 40, 370 ANNALS OF THE MISSOURI BOTANICAL GARDEN coccoid, cells 5-6 и in diameter; medulla about 130 м thick above the uppermost layer of bark cells, of loosely woven hyphae 2-3 и in diameter, inspersed with minute granules or crystals, below which is another layer 80—105 u thick between the bark cells, of similar structure; soralia elevated, 1 mm. in diameter, soredia coarsely granular, pale sulfur-colored, about 50—60 и in diameter. Apothecia sessile, urceolate, margin entire, 1 mm. in diameter, disc vinaceous buff, white- pruinose; amphithecium 125 u thick below the hypothecium, thinning to 50 p the somewhat inflexed top; cortex and algal layer as in the thallus, meiste of vertical, conglutinate, hyaline hyphae about 4 и in diameter, resting on the algal layer; thecium 200-210 р tall; paraphyses slender, dichotomously branched and occasionally anastomosing in the thecial gel; asci cylindric-clavate, 8-spored, 175 X 65 p, wall 8 u thick when the ascospores are mature; ascospores thin-walled, hyaline, unilocular, protoplast granular, 55 X 24 p Sierra Leone: Sefadu (Gbense), on trunk of Elaeis guineensis, P. Adames, comm. Е. С. Deighton M4750, fertile, type, 4754a, sterile. HAEMATOMMA НАЕМАТОММА Mass., Ricerche Autonom. Lich. Crost. 32. 1852. Loxospora Mass., Ricerche Autonom. Lich. Crost. 137. 1852. Ophioparma Norm., Nyt Mag. Naturvidensk. 7:230. 1853. Type: H. vulgare Mass. (Lecanora haematomma Ach.). Lichen ventosus L. was also transferred here. Loxospora was based on Lecanora elatina Ach., probably on immature material as Massolongo failed to see the septa and reported the asco- spores as unilocular. Ophioparma Norm. was based on Lichen ventosus L., L. haematomma Ehrh. and L. puniceus Smith. Thallus crustose (to squamulose in Н. dactylopbolis (Nyl.) Zahlbr.), corti- cate, of subvertical, conglutinate, thin-walled hyphae; algae protococcoid; medulla of loosely woven hyphae. Apothecia usually sessile, rarely subimmersed; amphi- thecium present; parathecium present or absent; hypothecium hyaline; asci 8- spored; ascospores hyaline, fusiform to acicular, septate, 4—pluri-locular, proto- plasts cylindric. Spermogonia in thalline warts, dark about the ostiole; spermatia cylindric, straight or curved. — . Thallus UM (resembling that of the Е ee mm.; ascospores 4-locular, 22—32 X 2 u; Sio Thomé.............. H. de еси qoi user 1. Thallus c oie tes E E 2. Adis immersed or nearly so, more or less кр; disc reddish to testaceous; Ka ss subarthonioideum (Zahlbr.) Hille. Apothecia sessile 3. Disc fering white-pruinose, mar, gin subcrenulate; ПН fo aie H. “Brassil pos 3. Disc not pruinose, ERR margin entire; apothecia often aggregated and pe from mutual pressure; Abyss . similis Bagl. HAEMATOMMA Brassii Dodge, sp. nov. Type: Nyasaland, Kasungu Hill, 1100 m., corticole, L. J. Brass 174582. Thallus epiphloeodes, continuus, laevis vel minutissime verruculosus, sub- rimulosus, subpruinosus, albidus, 250 p. Crassitudine, margine indistincto; cortex 1953] DODGE—LICHENS OF TROPICAL AFRICA 371 15 p crassitudine, hyphis subverticalibus tenuibus; stratum algarum 75 y crassi- tudine, cellulis Trebouxiae, 6-8 м diametro, sphaericis vel subangulosis pressione mutua; medulla 160 р crassitudine, hyphis subverticalibus laxe intertextis, 3 p diametro. Apothecia rotundata, basi constricta, juventute urceolata, margine elevato, crasso, integro, dein disco plano, dense albo-pruinoso, margine subcrenulato; amphithecium inferne 105 y crassitudine, superne ad 30 д tenuescens, cortex et stratum algarum ut in thallo, medulla deest; hypothecium conicum, substipitatum dein subpulvinare, centro 300 y crassitudine, ad marginem thecii tenuescens, hyphis verticalibus conglutinatis, rare cum coloniis algarum ad 100 X 40 ш; thecium circa 80 р altitudine; paraphyses tenues apicibus non incrassatis; asci stipitati, clavati, apicibus incrassatis, 72 X 16 y; ascosporae octonae, hyalinae, aciculares, multiloculares, 40 X 4 p. allus epiphloeodal, continuous, smooth or very minutely verrucose, slightly rimulose and pruinose, 250 м thick, white, margin indistinct; cortex 15 р thick, of slender, subvertical hyphae; algal layer 75 д thick, with occasional colonies in the medulla, Trebouxia, cells 6-8 м in diameter, spherical or somewhat angular from mutual pressure; medulla 160 џи thick, of loosely woven subvertical hyphae 3 p in diameter. Apothecia circular, constricted at the base, urceolate at first, margin elevated, thick, entire, disc concave, at maturity margin becoming sub- crenulate and disc plane, densely white-pruinose; amphithecium 105 р thick below, thinning to 30 д above, of@the~same’ structure as the thallus but lacking the medulla on the sides of the thecium; hypothecium conic, substipitate, becoming somewhat pulvinate, 300 p tall in the center, thinning to the margin of the thecium, of vertical conglutinate hyphae, occasionally with large, ellipsoid colonies of algae up to 100 Х 40 » which may push up into the base of the thecium, re- sembling internal cephalodia, but the algae clearly Trebouxia; thecium not sharply differentiated from the hypothecium, about 80 ш tall; paraphyses slender, septate, tips not thickened in the epithecial gel; asci stipitate, clavate, 8-spored, about 72 X 16 p, tips thickened, protoplasts hemispheric above; ascospores acicular, multilocular, hyaline, 40 X 4 HAEMATOMMA DACTYLOPHOLIS Zahlbr., Cat. Lich. Univ. 5:765. 1928. Lecanora dactylopholis Nyl., Flora 69:172. 1886. Туре: São Thomé, Bom Successo, 1050—1250 m., A. Moller. l Thallus epiphloeodal, imbricate-squamulose, squamules whitish, 0.5 mm. in diameter, white-puberulent below, often with dactyloid fibrils above, up to 1-2 mm. long, crowded. Apothecia 0.5-1 mm. in diameter, constricted at the base, disc pale testaceous or pale yellowish; amphithecium crenate; asci 8-spored; asco- spores acicular, 3-septate, 22-32 Ж 2 p; paraphyses not very distinct, epithecium and hypothecium hyaline.—N ylander. | Only a single small thallus, with 8 apothecia оп a small twig, was available for study. The thallus was probably somewhat immature, small discrete squamules on a black hypothallus, up to 0.5 mm. in diameter, margins dactyloid, ascending [Vor. 40, 372 ANNALS OF THE MISSOURI BOTANICAL GARDEN but not reaching the dimensions recorded by Nylander, about 130 р thick; cortex 16 p thick, of erect hyphae 2—3 џи in diameter; algal layer 35 р thick, Trebouxia, cells 6-8 м in diameter; medulla 55 p thick, of loosely woven vertical hyphae; hypothallus 25 и thick, of periclinal hyphae 6-8 и in diameter, septate, somewhat brownish, with occasional short rhizinal branches. Apothecia 0.5-1 mm., con- stricted at the base; amphithecium crenate, 130 м thick below the thecium, 80 p thick on the sides where the algal cells are more crowded and the medulla is absent; no parathecium; hypothecium 55 p thick, of slender, densely woven hyphae; thecium 60 p tall; paraphyses slender, conglutinate, tips not thickened in the pale epithecial gel; asci subcylindric, 8-spored, about 30 X 8 p; ascospores 4-locular, acicular, hyaline, about 22 X 2 y. Sierra Leone: Sugar Loaf Mountain, 645-775 m., оп twig, Е. C. Deighton M4441b. PARMELIACEAE Thallus foliose, resupinate to erect and almost fruticose, but always dorsiventral, usually attached to the substrate by rhizinae; usually corticate on both surfaces (ecorticate below іп Anzia and Pannoparmelia) ; algae protococcoid, lower surface nearly nude or more usually covered with rhizinae, which rarely anastomose to form a spongy hypothallus (Anzia and Pannoparmelia). Apothecia circular, sessile to short-stipitate; amphithecium well developed; parathecium absent or inconspicuous; paraphyses branched or unbranched; asci 8-spored (16—32-spored in Anzia and Candelaria, 2—4-spored in Menegazzia); ascospores unilocular, hyaline (septate in Megalopsora and Physcidia). Of the dozen genera, all but Parmelia are characteristic of subarctic and tem- perate zones, or are endemic on other continents. PARMELIA ParMELIA Ach., Meth. Lich. 153. 1803. Imbricaria Ach., К. Vetensk. Akad. Nya Handl. 15, 250. 1794. Physcia S. Е. Gray, Nat. Arr. Brit. Pl. 1:455. 1821, non Schreber, 1791. Type: Lichen saxatilis L. Thallus foliose, appressed or ascending, laciniate with rounded or linear lobes, often imbricate; upper surface often sorediose or isidiose, lower surface either darker or lighter than the upper, usually covered with rhizinae except at the mar- gins (absent in subg. Hypogymnia); upper cortex of vertical hyphae forming 2 pseudoparenchyma, lower cortex usually similar (but of periclinal hyphae in the Antarctic sect. Physcioideae) ; medulla of loosely woven, mostly periclinal hyphae; algae protococcoid. Apothecia on the upper surface, never marginal, sessile ог short-stipitate, disc concave to flat, often chestnut brown; amphithecium promi- nent; parathecium poorly developed or absent; hypothecium with algae below; paraphyses imbedded in a gel, usually somewhat branched above the asci, septate tips clavate or pointed; asci clavate, usually 8-spored; ascospores hyaline, uni- locular. Spermogonia immersed in the surface of the thallus or amphithecium, 1953], ' DODGE—LICHENS ОЕ TROPICAL AFRICA 373 spherical or pyriform, opening by blackened ostioles; wall black or dark brown above, light brown to hyaline below; spermatiophores simple or branched; spermatia cylindric or fusiform, often slightly constricted in the middle. Only the sections Hypothrachyna and Amphigymnia are represented in our collections, although the section Xanthoparmelia has been reported from our area. HYPOTRACHYNA 1. Thallus both isidiose and sorediose, rimulose, pale glaucous; wo SOCOLEA......1 tiliacea v. с Mill. Е E Vistas isidiose but d disi a 1. Thallus sorediose but n ee ose 6 1. Thallus neither 1519105 redios 11 2. Thallus vum broad, Е dissected 4 2. Thallus lobes 3. Thallus glaucous, poen £u "isidiose; 1l mbar P. cetrata v. subisidiosa Müll. ^d 3; Noted Pale yellow, center not ч Кен о 6-10 mm.; asc Pe 12-1 ; Soe otra P bweinfurthii Müll. Arg. 4. rie dos scrobiculate and isidiose, margins microphylline; Uu. P. revoluta v. ambigua Stein ce not scrobiculate nor margins microphylline 5 6. е а = xum medulla K yellow then red, C pale yellow; saxicole; French Guinea and Ivor P. hee к des Abb. 5. Lobes concrescent, n д ucous; medulla K—, C—; corticole; Angola........ scens Vainio 6. En m of lobes reticulate-rimose with nta along the MP ups center а иді confluent pang: Usambara irima f. sorediata Müll. igi Thallus not as abov 7. а, granular, беду large areas; thallus pale glaucous, under-side yellowish, center pale fuscous; apothecia 5—10 mm.; ascospores 16-18 X 10-12 д; Kenya P. Hanningtoniana Müll. Arg. 8 rd Soredia i in discrete or confluent soralia jns с— 8. Маг а еэ 9 8. 10 9. Soralia кү їп in ceneral Portion є of thallus; medulla K—; sterile; Eritrea............ Р. asmarana Vainio 9. Soralia on lobes nfluent, sorediose, git late ‘ad Е суре] ик yellow, ا‎ леа, in spots; apothecia 4—6 mm -15 ara d fi number 24 С Zahlbr. 10. р few, laciniae longer and narrower than in ee seis o г mosses; cinior v. Байна ophila p 10. Soralia abundant in the center with a few on the си thallus 10 (15) c eee pale ashy, lobes 3—5 i J mm. wide; medulla K—, C pink; ا‎ bo both cole rei на es Ivory C .P. Mangenoti des me 11. Thallus b black 11. Thallus pale fuscous, Bila cac oris Е ed! nts above; ultimate lobes 3 wide and long; as es 7-13 X 5 Би ‚Р. leptasca Sana & emt ll. Thallus pale or hie "dod is Tak the А d fai 1 11 dulla K yellow then orange 12. Lobes with cilia, K т slowly an р DE 7 — medulla К уе е red, С, n roots of orchi orchidophila у 13. Assad small, 2.5-3. r in damer 13. —; 14-18 X 9-12 р; Angola................-.- Apothecia 2.5-6 mm.; modella 1 К—; ascospores oes dpa onse ч уаш 13. Б 24—34 a 12-17 p; Fer- зонт 7—12 mm.; medulla К yellow then red; ascospores LE ert ЛЕЙ 14. "Medulla К. yellow; Uganda aska a о — Vainio - 7-11 X f; jy: Emi 14. Medulla K yellow then rufous or blood-red; ascospores и Е - Zahlbr d Mene and poorly described; Tanganyika .......-..------- - рай erbe dd 228 боран v. rapi cng Sambo 16. rbi d ыы P. tiliac v. bypoleuca M Arg. [Vor. 40, 374 ANNALS OF THE MISSOURI BOTANICAL GARDEN iz. менки 6—15 mm. in diameter; lobes ач mm., densely rhizinose below e in P icornis Tuck.); ascospores 8—9 X 3—4 д; Gaid a P. daik Jatta 17. крыса small, sessile; lobes 1 a mm. — medulla C— 18. Medulla E yellow; growing over mosses; Uganda P. leucorbiza Vainio 18. Med ulla K yellow then red; На ee ia x mm.; ascospores 9-12 X 6-7 и; corti- ; Angola Р. Руростаеа Vainio PanMELIA orchidophila Dodge, sp. nov. Type: Uganda, Western Province, Toro District, Nyinabitaba, in ridge forest, 2500 m., on roots of Tridactyla bicaudata (Lindl.) Schltr., Н. A. Omastin 1184. Thallus 3—4 cm. diametro, 95-100 p crassitudine, olivaceo-alutaceus, crenatus, sinibus excisis, lobis rotundatis, lobulis ultimis ad 3 mm. latitudine, marginibus ciliatis, ciliis ca. 0.5 mm. longitudine, laevis, subimpressus, albo-reticulatus, sub- rimulosus, inferne niger, marginibus castaneis, rhizinis nigris, semel bisve dichotome ramosis, centro thalli 1 mm. longitudine, ad margines brevioribus, omnino tectus; cortex superior 12 р crassitudine, pseudoparenchymaticus, hyphis verticalibus adglutinatis, cellulis leptodermaticis, 5 и diametro, granulis minutis nubilatis; stratum algarum 16-20 р crassitudine, cellulis protococcoideis 5 д diametro; medulla 55 р crassitudine, hyphis periclinalibus, pachydermeis, 3 и diametro, minutis cum granulis albis vel pallide alutaceis (aurantiacis in partibus moribundis) nubilatis, dense contexta; cortex inferior niger, 12 p crassitudine, pseudoparenchy- maticus, cellulis 4 д diametro; rhizinae 20 и diametro. Steri Thallus 3—4 cm. in diameter, 95—100 p thick, deep olive buff, K very slowly light yellow, lobes rounded, crenate, sinuses excised, ultimate lobules up to 3 mm. wide, margins with cilia about 0.5 mm. long, surface smooth, slightly impressed, white-reticulate, slightly rimulose, underside black with chestnut margins; rhizinae covering the whole under-surface, black, once or twice dichotomously branched, about 1 mm. long in the center of the thallus, shorter toward the margin; upper cortex about 12 р thick, pseudoparenchymatous, hyphae vertical, cells thin-walled, 5 p in diameter, nubilated with minute granules; algal layer 16—20 p thick, of compactly interwoven, mostly periclinal hyphae, heavily incrusted with minute granules white to pale buff, becoming orange in moribund thalli, K yellow then orange red, С, KC—; lower cortex black, about 12 и thick, of very dark brown pseudoparenchyma, cells about 4 и in diameter; rhizinae 20 p in diameter. Sterile. AMPHIGYMNIA l. Thallus isidiose (for о lacerate and microphylline margins, see No. 11 below ).........--- 2 1. Thallus хе soredios (ON И 6 1. Thallus without isidia Dr amedid ОРООНО о ааа EM 9 2. lus white to ashy or pex olive buff; lobes rounded, 15 mm. or more Ьгоай.........----------- 3 2. Thallus yellow- green .. d rim Ie. س h‏ Isidia confined to nx center of the thallus, not marginal; es и P. nitens f. indios “Mall. Arg. . Isidia on upper surface, "iia бадаг 5 К yellow then ferruginous; apothecia 2—4 mm.; v ascospores 9—12 X 6-8 кы que. ‚Р. isidiza Ме 3. EE abundant on margins ire eese xs bres pecia Isidia up to 2 m tdi some contio, ieri K—, su KC—; thall and hid» yllo owing; Sierra Leone................... » L bulascens v. pA eis Dodge slowly 4. рми ети short, 0.5 mm. tall, almos Sce medulla peel C р pink to blood thallus K егыла А yellow; Ivory Coas P. pseudotinctorum des Abb. 1953] DODGE— LICHENS ОЕ TROPICAL AFRICA 375 5. Surface fane with isidia in the depressions; Kenya P. creto v. isidiophora aos 5. Surface not wrinkled; center densely isidiose; Tang: ganyika тах ta Müll. A 5. Surface pale alles, isidiose and microphylline margins; apothecia 6-10 mm pale fuscous; ascospores 22-25 X 12—15 mw; Socotra Р ana ман P Mill. А, Hus eins with short cilia allus without cilia ^ 75 ا‎ age? ple olive buf, rugulose; medulla K—, C pink, KC—; subcil x OUS qe » ntm rugulose, glaucous; medulla K—, C—; Abyssinia........ Жеб у. sored эха Arg 8; Thales pale glaucous X аита К yellow, ец gun PARI ds жа bs Archipelag Hilden brandti cee hbr. 8. Thallus dio ous, жук medulla C iuge Ай оа ЫЕ. Р; Жы v. sorediosa Vainio 8. Thallus pale ashy, medulla K—, C—; sterile; French Guinea eid Ivory Coast........-- тарауда des 286. 9. Thallus ciliate 9. Bu. m cil D s bilanci to lacerate; thallus K yellow. H 6 t lac nor lobulate 11. выть K ‘yellow, С aor thallus lobes 1.5 (-2) cm. wide; Cameroons........ P. lobulascens Sine 11. Medulla K—, 12. Ascospores 1930 X 12-17 ns — pale lurid, unchanged when moist; apu al lobes m. wide; 1 P. amaniens & Zahlbr. 12. Ascospores 1 pue x Н 2 th wien glaucous, surface rugulose; Abyssinia... i ду e Nyl. Re — ores 8—14 X 6.5-8 u; thallus ashy lurid, yellow green а moist; peripheral 3—5 mm. wide; enn neirobiensis Stnr. & en i сн 00р та argin white or very pale below TX PE margin deep brown to black below M 14. Ascospores rhet xX3101-12- i; Usambara....e = . Hildenbrandtii v. Sasss Müll. Arg. 14. Ascospores 14-16 X 8—10 д; Usambara...........--.-------- z dei f. gla & Zahlbr. 14. Ascospores 20-24 X 10-13 и; Abyssinia i Sc Me deve м Il. Arg. 15. Medulla K—; ascospores 13-17 X 5.5-7 м; Usambara .P. procera Stnr. & Zahlbr. 15. Medulla K yellow 6 16. Ascospores 19-29 X 11.5-18 ш; ОзапаБага...................-———- .P. eurycarpa Stnr. & Zahlbr. 16. Ascospores 13-19 (-20) X 7-10 м; Kenya APs pe ciate Steiner 16. Азсозрогез 1 —11 р; Kenya а Hue 17. Ascospores 15-23 X 7-11 и; Ѕодап...........------------------ т rican gery Arg. 17. Ascospores 19 X 11 п; along Zambesi River Menyhartii Steiner 17. Ascospores 8—15 X 7-10 р cenesnsense--neseesneensrnsenennseacenssnnsunennsnannstasenannnannennennsssse# 19 17. Ascóspotes 15-18 Х 7-08 LL ee E a e 18 од x: 18. Apothecia 2-11 mm.; ascospores 15—17 X 7-8.5 и; along мина eon И 18. Apothecia 4 mm.; ascospores 14-16 X 6-7 и; Angola........ P. usar f esorediata Vainio 18. Apothecia 7-20 mm.; ascospores 15-18 X 8-10 и; ARGOS byporysalea мег 19. Apothecia perforate .............. eet IX Арлен Dua fa р ————— ^ 20. Ascospores 11-12 X 6-7 и; apothecia up to 3.5 cm. іп diameter; M Reset sore E паи e ee a ases Л ЕЕЕ in diam =2 шей +. ету Stnr. Гг Zahlbr. 21. Apothecia 2—4 mm.; ascospores 11—15 X 7-9 p; Kenya... Р. а Ное 21. Apothecia =: mm.; ascospores 9—11 X ‘6-7 8, "d ERU EAMUS P. ronds Mäll. га 21. Apothecia mm.; ascospores 10—12 p long; Usambara............———7 ens Müll. A PARMELIA LOBULASCENS Steiner var. isidiosissima Dodge, var. nov. Type: Sierra Leone, Sefadu (Gbense), on trunk of Elaeis guineensis, P. Adames, comm. F. C. Deighton M4754. Thallus foliosus, 10 cm. diametro, 135—150 р crassitudine, pallide olivaceo- alutaceus, irregulariter lobatus, lobis aliis rotundatis, usque ad 15 mm. latitudine, marginibus laevibus, crispatis, sinibus irregulariter excisis, eciliatis, aliis 3—5 mm. latitudine, erectis, marginibus isidiosis aut lobulatis; isidiis granularibus aut [Vor. 40, 376 ANNALS OF THE MISSOURI BOTANICAL GARDEN stipitatis, ramosis, 2 mm. altitudine, 0.5 mm. diametro; lobulis non isidiatis raris, 1 mm. longitudine, 0.5 mm. latitudine, basi subconstrictis; superficies inferna nigra, opaca, minute rugulosa, rhizinis centralibus, lobis latioribus marginibus castaneis aut pallide brunneis, laevibus, nitentibus; lobis angustioribus aut rugulosis et nigris ad marginem aut alba cum zona marginali, 1 mm. latitudine; cortex superior ca. 25 р crassitudine, pseudoparenchymaticus, cellulis leptodermeis 6 p diametro, ad superficiem minoribus; stratum algarum 15—25(—40) и crassitudine, protococcoideum, cellulis 6—8 и diametro, minutis cum granulis; medulla ca. 65 и crassitudine, hyphis pachydermeis, 3 и diametro, laxe intertextis, granulis inspersis; cortex inferior niger, 15—25 р crassitudine, pseudoparenchymaticus. Sterilis. Thallus about 10 cm. in diameter, 135—150 и thick, pale olive buff, К faint yellow, irregularly lobed, some lobes rounded, up to 15 mm. broad, margin smooth, crisped, sinuses irregularly excised, eciliate, other lobes only 3—5 mm. broad, more erect, margins isidiose or lobulate, bearing granular to stalked and branched isidia 2 mm. tall, 0.5 mm. in diameter; non-isidiose lobules very rare, about 1 mm. long, 0.5 mm. broad, somewhat constricted at the base; under-side black, opaque, minutely rugulose, rhizinae about 1 mm. long, black, confined to the central por- tion, attaching the thallus to the bark; broader lobes shading through chestnut to light brown at the margin, nearly smooth and shining; the narrower lobes either rugulose and black to the margin or abruptly white in a narrow zone about 1 mm. broad; upper cortex an irregular palisade about 25 p thick, of thin-walled pseudo- parenchyma, cells about 6 д in diameter, somewhat smaller next the surface; algal layer 15—25 (—40) и thick, protococcoid, cells 6—8 и in diameter with abundant minute granules; medulla about 65 p thick, of loosely woven, thick-walled hyphae about 3 p in diameter, more periclinal and compact below, interstices nearly filled with granules, K—, C—, KC—; lower cortex black, 15-25 и thick, pseudo- parenchymatous. Sterile. The relationship of this variety to P. lobulascens is somewhat doubtful, as I have found no short, marginal cilia; the upper surface is wholly smooth and the hyphae of the medulla are more loosely woven and slightly smaller than in Steiner’s description. PARMELIA PSEUDOTINCTORUM des Abb., Bull. Inst. Franc. Afrique Noire 13:973. Type: Ivory Coast, Mt. Tonkoni (cercle de Man), 1150 m., saxicole, H. des Abbayes; Mankono (cercle de Séguela), on granite, H. des Abbayes; Séguela, on granite, H. des Abbayes. Thallus at least 16—20 cm. broad, 135 и thick, pale olive buff to deep olive buff where covered by isidia, K yellow, lobes rounded, 2.5 cm. broad, sinuses not excised, surface smooth to slightly undulate in the outer 1.5 cm., the rest densely isidiose, isidia 0.2 mm. tall, simple or dichotomous; under-side black, minutely verrucose and rugulose, margins shading to isabelline, very rarely almost white; rhizinae scarce, simple, tip densely branched, less than 1 mm. long; upper cortex about 1953] DODGE—LICHENS OF TROPICAL AFRICA 377 16 р thick, a palisade of thin-walled hyphae, 5—6 р in diameter, forming a pseudo- parenchyma, somewhat nubilated with grayish granules; algal layer 15-18 p thick, cells protococcoid, 6-8 м in diameter; medulla 75-80 p thick, of densely woven, periclinal hyphae, thick-walled, 3 » in diameter, nubilated with grayish granules, K—, C blood-red, KC—; lower cortex about 15 p thick, black, pseudo- parenchymatous, cells about 3 & in diameter. Apothecia 15 mm. broad, 10 mm. tall, stalk 3 mm. in diameter, 7 mm. tall; exciple shallowly sulcate and scrobiculate, sparsely to densely isidiose, disc concave, imperforate, margin extending 0.5 mm. beyond the thecium, isidiose on the edge; amphithecium 40-175 p thick near the margin, of the same structure as the thallus with an algal layer under the hypo- thecium as well as next the upper cortex; hypothecium about 25 y thick, of slender, periclinal, gelified hyphae, appearing almost amorphous, hyaline; thecium 40 р tall; paraphyses slender, dichotomous, coherent, tips not thickened in the pale brownish epithecial gel; asci clavate-cylindric, 30 X 13 p, tips thickened, proto- plasts mamillate when young, 8-spored; ascospores distichous, long-ellipsoid to subfusiform, unicellular, hyaline, about 12 X 4-5 р Our specimens differ in several respects from the description of H. des Abbayes; the dimensions of the thallus and isidia are greater, the exciple is more scrobiculate, and the ascospores narrower. In some respects it is intermediate between this species and Parmelia tinctorum Despr. from the Canaries. Sierra Leone: Sefadu (Gbense), on trunk of Elaeis guineensis, P. Adames, comm. F. C. Deighton M4753. PaRMELIA subciliaris Dodge, comb. nov. Parmelia nilgherrensis Nyl. v. subciliaris Vainio, Hedwigia 37:(40). 1898. Type: Uganda, Ruwenzori, 0° 5' S., 2900—3200 m., С. Е. Scott-Elliott 218. Thallus foliose, about 4 cm. in diameter, pale olive buff, lobes about 1 cm. broad, margin crenulate, upper surface K yellow, smooth to slightly impressed, rugulose, minutely rimose-areolate in the older portions, cilia about 2 mm. long, simple or once dichotomous, black, margins of some lobes capitate-sorediose, soralia about 1 mm. in diameter, rarely subconfluent; under-surface rugulose, black, somewhat lighter at the margin; upper cortex 40 и thick, a palisade of pseudoparenchyma, cells about 6 p in diameter; algal layer 25 p thick, cells proto- coccoid, 7 p in diameter; medulla 140 р thick, of interwoven, thick-walled, mostly periclinal hyphae 6-7 p in diameter, K—, C pink, KC—; lower cortex about ot thick, black, pseudoparenchymatous, cells 4 p in diameter, rhizinae about 60 p in diameter, 3—5 mm. long, relatively few and confined to the middle of the thallus, twice or thrice dichotomously branched. Sterile. : Uganda: Ruwenzori, Western Province, Toro District, ridge forest on Nyina- bitaba, 2500 m., on roots of Tridactyla bicaudata (Lindl) Schltr., H. A. Omastin [Vor. 40, 378 ANNALS OF THE MISSOURI BOTANICAL GARDEN PaRMELIA Sovauxu Müll. Arg., Linnaea 63:32. 1880. Type: Angola, Pungo Andongo, Soyaux. Thallus very large (two pieces available for study, 10 X 7 cm., and 11 X 7 cm.), 130—150 y thick, pale olive buff to olive buff, К yellow, lobes crenate and crisped, up to 2 cm. broad, suberect, surface smooth, rimulose in the older portions, white-reticulate; under-side black, minutely rugulose; rhizinae very few, simple or with branched tips forming а holdfast; upper cortex 15 y thick, of thin-walled pseudoparenchyma about 2 cells thick, heavily incrusted with minute yellowish crystals; algal layer about 30 y thick, continuous, cells protococcoid, 5-6 и in diameter; medulla 80 p thick, of loosely woven, mostly periclinal very thick- walled hyphae 3 м in diameter, more densely woven next the lower cortex, K—, С blood-red, KC—; lower cortex 12—15 № thick, of septate, brown, conglutinate hyphae about 6 y in diameter, cracking away and leaving the lower closely woven hyaline hyphae of the medulla to form a new pale buff cortex. Apothecia perforate, which is almost completely gelified with very few minute yellow crystals; hypo- thecium 30 y thick, almost completely gelified but showing traces of periclinal hyphae; thecium 65 џ tall; paraphyses conglutinate; dichotomous above, tips not thickened in the very pale brownish epithecial gel; asci broadly clavate, very thick- walled at first, tips remaining thick until the spores are mature, 8-spored, 35—45 X 14-18 и; ascospores ellipsoid, unilocular, hyaline, 11-12 X 6-7 и. Spermogonia oblate-spheroid, up to 130 № in diameter, 105 р tall, immersed in the medulla, neck about 15 џ long, 25 м in diameter, wall wholly carbonaceous at maturity, pseudoparenchymatous; spermatiophores simple or dichotomous near the base, about 20 X 1 p; spermatia cylindric, straight, 16-18 X 1 y. After the thecium disintegrates, the hypothecium functions as a cortex, leaving a slightly rugulose surface concolorous with the thallus, but not white-reticulate. Müller Argau evidently had only young apothecia ("parvula"), as young apothecia only 2-4 mm. in diameter are also present in our material In habit, our plants suggest Parmelia latissima Fée, but internal structure is quite different. Sierra Leone: Picket Hill (Colony), 740 m., T. S. Jones, comm. F. C. Deighton 4592. USNEACEAE Thallus fruticose, erect, Prostrate or pendent, attached to the substrate by a hapteron, corticate with longitudinal, conglutinate hyphae in Alectoria, Oropogon and two species of Ramalina, otherwise fastigiate, of compact, subvertical hyphae, only somewhat pseudoparenchymatous; algae protococcoid (Trentepohlia in Usnea 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 379 sect. Roccellinae) ; medulla compact to arachnoid, often with а chondroid axis or strands of thick-walled, conglutinate, parallel hyphae, variously disposed. Apo- thecia circular, sessile or stipitate; amphithecium well developed; азс! 1—8-spored; ascospores hyaline or brown, unicellular to muriform, relatively thick-walled. 1. Ascospores unicellular 2 1. Ascospores bi AE cortex oir n^ E 1. Ascospores mu ‚ lar porou on. Th. 2. Medulla d a i sili Aabar of НЫ of — tNUE н... Шт мев Zahlbr. 2. Medulla uniform, either DH or compact and h 3. Cortex of conglutinate, periclinal Бур Alectoria rod 3. Cortex rne ate, of sul ical hyphae 4 4. Medulla of loosely tangled hyphae 5 4. beg ot periclinal hyphae "d 3. 6 s Thallus bo bollo, dni sia ee flattened Evernia Ach. Tul more or less inflated Dactylina Nyl. us poderi iform, E ort, th Endocoena Crombie 7, E P loos ven hi ads а КЕШ чөй ra este AE егы id st ae Thamnolia Ach. 7. Medulla either Haas artilaginous, or inn i diio entiated as a chondroid axis................ 8 8. Thallus podet form, relatively short, ew aliad apothecia unknown.............. Siphula Fr. 8. Thallus fruticose, usually tn longer 9. Thallus flattened, whole gue As саре, algal yin lacking O o oaae Everniopsis Nyi. 9. Thallus cylindric or ет See Па loose surroun hei ex ANE S icm: HH ve 10. Specialized m cal tissue not ACE ааа aziera Mont 10. Specialized HM tissue highly graec Ыы cdi en flattened; метал ia vule Ramalina Ach. USNEA Usnea Hill, Gen. Nat. Hist. 2:85. 1751; Wigg., Primit. Fl. Holsat. 90. 1780. Type: Lichen floridus L. Thallus fruticose or filamentous, usually of several compound branches, dicho- tomous, rarely sympodially branched, or unbranched, 1 cm. —7 т. long, erect, prostrate or pendent, attached to the substrate by a hapteron; branches thick at the base, thinning uniformly or abruptly toward the apex, 0.2-7 mm. thick, terete, angled or longitudinally sulcate or foveolate, smooth, verrucose, papillate, tubercu- late or spinuliferous, continuous, areolate or annulate; soredia farinose or isidioid; cortex thin and fragile, or thick and cartilaginous, of densely woven, thick-walled, conglutinate, vertical hyphae, evanescent on the primary branches in a few species; algae usually protococcoid (Trentepohlia in the Roccellinae) ; medulla of loosely or compactly woven longitudinal hyphae; chondroid axis single, of thick- walled conglutinate, longitudinal hyphae, usually solid, rarely lacerate on the sur- face (hollow with a few arachnoid hyphae in the subg. Ewmitria). Apothecia lecanorine, cup-shaped, plane or somewhat irregular, lateral (appearing terminal when formed near the tip of a branch, bent sharply and partly fused with the apothecium, the tip then resembling a long cilium), margin thin, smooth or scrobiculate, nude or ciliate; paraphyses conglutinate, septate, somewhat branched; asci subcylindric, 8-spored; ascospores unilocular, ellipsoid, hyaline. Spermogonia rare, immersed in the cortex, wall pale or slightly darkened; spermatiophores spar- ingly septate; spermatia straight, one end often slightly thicker. [Vor. 40, 380 ANNALS OF THE MISSOURI BOTANICAL GARDEN Of the six subgenera, only Eumitria and Euusnea occur in our area. In pre- paring the following key, I have followed Motyka, Lich. Gen. Usnea Stud. Monogr. 1-651. 1936—1938, giving his geographic distributions for the various species. 1. Algae алые thallus grayish green, rigid, 5—10 cm. tall 2 5 gae protoc 3 2; е wid soredia and AME а axis with some blackened Baral a abundant, 3—5 mm. in diam anganyig ea у: U. per erbispidella Steiner 2 Thallus rt soredia o or La axis at a narrow central em apotheci cia rare, iec. chtensteinii Steiner 3; Chondroid axis chile in vb eke Крыва partly filled pies a = loosely w hypha pothecia usually ciliate, rarely eciliate; thallus ooth or egat Aia Жс scrol eulie nor tubércalates soredia isidiose; subg. ПОМНИ, 4 3. Chondroid axis solid, rarely somewhat Бани: never with a distinct cavity; subg. EUUSNEA.....9 4. Thallus distinctly angular (at least papillae in rows), smooth between the papillae white, rather thick and lax; Mozambique. U ped M . Thallus terete, not angled nor papillae in rows 5. Thallus almost eramulose, with a few irregular ra muli, HOMI ashy green, branches perpendicular, straight or irregularly flexuous, 2 mm. in diameter, tapering gradually to the apices; sterile 5. Thallus typically ram mulose, branches divergent, farinose : mt of ot narrow; thallus olive green; “Angola U. Теме Mot Cavity of axis broad; thallus glaucous; Tanga elata Mot. 2 Thallus а not sore pes branches somewhat Dem. apothecia 0.7-1 cm. in dan. r, disc pruinose; Uga U. firmula Mot. 7 Thallus rov » soris, sad y green, often prui 8 8. Thallus osos ашы apr ачат Tanganyika.......... U. Baileyi ee Mot. E але. 112 omm n diameter, ا‎ aa below, "A were Las Helena Е ee ee T im plici № ч”, Mot. 9. Thallus folate or scrobiculate, not terete, epapillate, flaccid, чае not blackened; E кыы iu о 10 9. Thallus doe or acutely 6 usually more rigid. 15 allus 15—20 cm. long, not fruticulose, very flaccid 11 p Thallus 3—9 cm. tall, Е шр. а. 12 11. Thallus ipa. ag aee jJ RV NN TAL U. terrestris Mot. 11. Thallus n rediose, partly somewhat fuscous, not con torted; Camer а$...... U. versicolor Mot. 1. Thallus эзен fno 3 cm. tall; fertile; not sorediose; ree а ie PRE НОВИ U. pela tansy (Mill. Arg.) н 12. ТҺа ed, sorediose ................ 13. Thallus 9 cm. tall, distinctly aa, wrinkled; Abyssinia to Kenya U. corrugata Nix. 13. ree 67 « cm tall, not acu неу w: rinkle libe E R E MEC E ВЫ U. pu drei ts (Müll. Аг) Мог. 14. Мы. sparingly = eg ramulose; soredia сы on ridges; cortex larger branches not areolate; French Guine ры U. нон Мог. 15. т” articulate with orc s not papillate, Mem cm. long; medulla very FMIGULATAR ава 16 15. Thallus concentus or articulate without pseudocyphellae, often fruticulose, papillate { СС ТАНИНИ а ИО кошо ту ин оет 16. Thallus ed cm. long, regularly riole Tanganyika U. flavescens E hal mulose or with an occasional puel us, 25—40 cm. IONE erinra 17: Рети elevated, farinose; thallus 2 . long, 2 mm. in diam and СООО ee ee ee U езен Аг Р-У -— n (0 , 4 20. Th lus ue алача eous or mi green, not changing color in the herbarium steri ri a pasan Stirton 20. Thallus not foveolate, becoming lurid fuscous in the herbarium; apothecia 5 mm. in diameter; Abyssini U. rugosa Mot. военное ни, 1953] DODGE—LICHENS OF TROPICAL AFRICA 381 21. Thallus never ashy green, either рерин stramineous, volley у ог becoming fuscous in the иу; living plants with a thin papery cortex, o thicker, very smooth.......... 22 21. азе ashy gree rarely r "o r if NET papillate or ki na si idom becom- g brown in аы P* rbar 39 225 "Thallus long, pendent, subarticulate or moniliform; STRAMINEAE 23 22. Thallus Dn ne ulose; GLAB 30 23. Thallus pale neous or pale yellow, з= changing color in the herbarium, sub- ht most no ramuli; A 24 articulate 23. Thallus Aoao; fuscescent in the а залее ramulose; EUSTRAMINEAE; saxicole U odis Mot. 23. Tum" stramineous, soon fuscescent in the herbarium, sulcate and angled, Aas with abundan t ramuli SULCATAE 24. Thallus 0.6 mm. in diameter, very flaccid, medulla К reddening; Uganda a U — мс епуа 24. Thallus 1-2 mm. in diameter, rigid 25. Ramuli abundant, branches 2 mm. thick below, very slender mm Kenya nost. xasperata (Müll. Arg.) Mot 5. Ramuli confined to upper portion of branches in small cluster A s Helena......U. pulchella Mot. 25. Ramuli absent or very rare, not in clusters; thallus moni T Ane medulla iae dense; Kenya southward U oniliformis "n 26. Thallus less deeply and distinctly sulcat 26. Thallus very жрт lo ваат. su rp 27. Thallus 0.5 mm. in diamet compressed; East Africa U. e 27. Thallus 1.5 mm. in diameter; Kenya sout ard usca Mot. 28. Дене s2m ore in diameter, Mey angled, becoming quite ы апа rr coward ee tips; U. decipiens Mot. 2% Branches about 1 mm. in diameter, less acutely angled, ramuli few and aar a "M VEA 1 cm. in dia 29; Thallus 13, i long, ramuli very few and slender, up to 10 mm. кен exciple foveolat апда U. flaccida (Müll. Fuse Mot. 29. Thalia 40 cm. lon muli coarse, 5 mm. long; exciple smooth; Ug кыз quatoriana Mot. 30. Thallus subartieulate, sparingly ramulose; medulla lax; Mere P OCLADAB а. 31 30. Thallus subarticulate or continuous, with pco voe ramuli; A dila lax; fertile; ILIIFERAE 35 30. ds ш сылы; ramuli or spinules very crowded; medulla lax; fertile or | ediose; SCÃBRIDAE .............. 30. Thallus Eu o n when fresh, very dark fuscous in the he РЕ med wie ‚ compact; OsssoLucas; thallus 8.5 cm. tall, 3 mm. in — ‚ fru orediose; Tanganyika ......-..- еее e. Май. А) Mot. 31. Thallus. 12- 15 cm. long, ا‎ EA flaccid, irregularly branched ramulose; sored: in punctif soralia 31: Thallus 19 cm. tall, flaccid, more or less e RE t 32. Thallus very indistinctly papillate; Tanganyika............... hispidula ди Arg.) Za ^ia 32. Thallus with distinct wrinkles but cay pg БС pact Tanganyika | d southwar 33. Medulla К red, ramuli quite stiff, not crowded; thallus 7-10 cm. tall; Kenya ею. дне Laur. A3; Modulo Es o ELL TL LG аа — 34. Thallus 10 cm. tall; soralia in distinct user. Kenya and Uganda........ U. Haumanii Mot 34. Th per Ba e tall, soralia in indistinct, ewhat blackened tubercules; Abyssinia Co 20 اا پک ر‎ MM T U. ra et Mot. 5 Ramal ae but relat dd aps; mm in dim Si ce, Mot 35. Ramuli iow тозо р ре ud ктү 36. Thal iy sparingly b пас. branches с ог сваи im inflated; Pi iui 4 о QM HUNE 5 36. ا‎ distinctl pA Se ge epeatedly bra BE EE у GU OL STO NU EE 37. Thallus 3—5 cm. tall, Y indes, cu olivaceous in o» herbarium; apothecia small; Camer: vi "CN fuscous in the herbarium; e ish green becoming deep i ; T > sige Teia) Mot. а ARRA a cilia енга Usada lera po unn n d с ; spinules m тыйы us 9 em ^ whieh ian ы UNA yellow fun ноба. я deep red; deeem Tan kk southwar o кар УУ НИИ dulata Stirton sngany а southward sa ins Mes dense; medulla K yellow then s s y P TIR M г; Congo to Rhodesia......U. perspinosa Mot. ечиб red; apothecia 2—3 mm. іп diamete [Vor. 40, 382 ANNALS OF THE MISSOURI BOTANICAL GARDEN 39. Thallus мр —Á— thick and loosely woven, irregularly bi, apices short and thick, not filiform, smooth or чау tuberculate; DENDRI 40 39. Thallus dice а edu illa thin, compac 48 40. Thallus without pseudocyphellae, ev sparse; fertile; CLADOCARPAE....... 5 20. 41 40. Thallus so ge seudocyphellae, ри tly ramulose, usually ний лева Гал RS 44 40. E etel wit oe тоос yphellae, almost era sérii articulate, rugose with farinose ARIAE 46 41 Thallus à sine pe ndent, at least 1 mm. thick, usually thicker, tips rather rer and d thick, арау BÎ bb اا‎ rather lates medulla somewhat К reddeni in dia — = ika U. e ue мш 41 „Thallus dos diu а. cm. tall, usually much shorter, i وچ‎ not sore dios Thallus 5 cm. tall, dirty Зы olive green; apotheci mm. in di leniter, exciple papillate; West Africa U. scutata Mot. 42. Thallus 7—15 cm. tall; ес ир to 10 mm. in diameter; thallus ashy fuscescent e herbarium, denna ely ramulose and papillate 43 43. Thallus 8 cm. tall, medulla 600 д E en loosely woven; apothecia cupuliform, exciple ooth; ramuli inflated; Nige gola U. hispida Mot. 43. Tha idis 7 cm. tall, sodalis 400 | я chick apotecia plane or deformed, exciple scrobic- ulate; ramuli thick, not inflated; U. Le diewil Mot. 44. Thallus pale fuscous, white cet н firm, caespitose, branches subparalle Abyssinia U накі IRE 3 44. 'Thallus not wh otted 5; llus both fertile see sorediose; Abyssin J. abissinica Mior. 45. Tha Бо 45. Thallus fertile, not sorediose, smooth, етта cortex firm; Abyssinia to vnil RUM U. obtusata Mot. 46, Thallus coh green, medulla 900 м thick; Cameroons southward......U. Bornemuelleri Steiner 46. апа Thallus sepia to fuscous or almost — in the herbarium, branches articulate ibn medulla 600 u; Abyssin 47. Branches 2-3 (-5) mm. in dian mete er, iodi isidiose in ridges U. Eese kibr. 47. Branches 2 mm. in diameter; apothecia rare, 10 mm. in diameter, exciple smooth........ U Flotowii "bk serpentaria Mot. 48. ~~ sti thin, white, never rose; thallus fruticulose, sparingly branched, 12(-20) cm. tall, densely ramulose; SETULOSAE 49 48. Medulla ° oe rote-color or if white then thallus over 25 cm. long, pendent; 49. Thallus каре green, 3.5-10 cm. tall; DENSIROSTRA --50 49. Thallus with red or red-variegated cortex, 4—12 cm tally R BIGI -52 н llus 5 cm. tall, etd n ely ramulose throdéhotit; Hye branched; apo- a 5 mm. in iam ng U. me ме 50. Thallus irregularly ê part rtly papillate or ver 51. Thallus 3.5 cm. tall, er and tips coralloid publ: еее griek ы с oclada — Mot. 51. Thallus 8—10 cm. tall, ramuli unbranched; apothecia 2 cm. in diameters ен southward . J. picta 4 (Steiner Mot. 52; таси 20 cm. long, — face subarticulate, not densely p yellow then red; Kenya and Uganda Aag nicer SCENS pom Mot. 52. Thallus fruticulose, 4—12 tall, rigid. densely — атн уа ова 53. Thallus i: e tall, apothecia di Tanganyika southward............ U. 7 (Zahlbr.) Mot. 53. Thallus cm. belt. oedi = зеге 5 54. Sorli br oad, eroded; мы 12 cm. tall; bicolorata Mot. 54. T small; thallus m. tall, densely branched tips — у e ee less mspicuous than in Pd species of the section; Kenya to 1Јвапйа........................ ML IU ld Ч J. eri (Stein) Mot. 55. Thallus relatively thick, more or less docu but not angular or ые ne pale green, frequently: brésiched; Cuna TAE OSE ET Luan ea peat. ben 56 55. Thallus thick, e ar in cross-section, infrequently branched, green; GONIODEs..........——- 59 55. Thallus slender, te , infrequently branched, grayish green; LONGISSIMAE............ ase 61 56. — rose; re fe 6 cm. eee rather denis tamulose, pale green, axis ы 0ш seid lacerate in middle; Congo southward.................... anthera Vainio ewhat 56. мех wie thallus over 25 cm. long, pendent, наў and irregularly e Td 57 57. gem at са enti reddening or fuscescent; papillae irregularly arranged, adult bran abou in diameter, cortex iS: Congo southward RM ML SN o e U. distensa Stirton 37. Medulh YX— i ds odas ctun saccus nm ыйы сла T a M 58 1953] DODGE—LICHENS OF TROPICAL AFRICA 383 58. Thallus 60 cm. long, axis 500 д in diameter; ramuli close, larger, dichotomous; apothecia 10—20 (—30) mm. in diameter; Angola to Tanganyika........ U. amplissima Stirton 58. Thallus 25 cm. long, axis 250 и in diameter with some fuscous hyphae; ramuli dis- nt, short, tips farinose-verrucose; sterile; St. Helena U. um i à rd; Congo 59. Thallus rarely over 2 mm. in diameter, quite soft 60. Thallus 50 cm. long, almost without ramuli, folds at angles elevated; Uganda........ U. chloreoides (Vainio) Mot. 60. Thallus more than 100 cm. long, regularly ramulose, very long, quite soft; Nigeria southwar U. africana Mot. 61. Cortex well developed even on the primary branches, continuous or areolate 62 st. te. 64 61. Cortex of larger branches evanescent or shredded, mostly ecortica 62. Branches 0.9 mm. in diameter, ramuli sparse and irregular, 1—3 cm. long; St. len: U. Lyngei Mot. 62. Branches up to 0.8 mm. in diameter or partly thicker and fasciate-flattene 63 63. Thallus partly fasciate-flattened, axis solid; Cameroons U. livida Mot. 63. Thallus terete or slightly angular, deformed in larger branches; axis solid or wi yellow hyphae; regularly ramulose, quite rigid; Kenya . contorta Jatta 64. Thallus pale green, 1 mm. in diameter, ramuli 1-2 (=3) cm. long, rare; Congo........ U. arguta Mot. 64. Thallus pale stramineous to ashy green, wholly ecorticate, 0.4—0.6 mm. in diameter, d ramuli 0.5-1.5 cm. long, abundant; Congo to Tanganyika.............. U. trichodeodes Vainio UsNEA LEPROSA Motyka, Lich. Gen. Usnea Stud. Monogr. 106. 1936. Type: South Africa, Lydenburg, arboricole, Willms. Thallus 7 cm. long, subpendent, closely divergently branched, closely ramulose below, almost without ramuli above, very flaccid, soredia pale ashy green; base attenuate, expanding rapidly and repeatedly branching, 1.3 mm. in diameter, usually distinctly articulate, more or less terete but partly foveolate, smooth, farinose from confluent soredia, ramuli close, filiform, 0.5—5 mm. long, straight, the thicker again ramulose and sorediose. Apices quite long, sparingly branched, eramulose, flexuous and sorediose; cortex 100 p thick, soft; medulla 500 р thick, loosely woven, white, K—; axis 250 р in diameter. Apothecia 5 mm. in diameter, exciple smooth or with spinules; cilia few, short, thick, obtuse, sorediose; disc plane ог convolute, densely farinose. Soredia isidiose on minute ridges, variable in size. The above is based on Motyka's description, as our material is only a fragment, about 3 cm. long, with 5 apothecia and part of the cortex torn away in collecting. Such characters as are observable agree with the above description. Sierra Leone: Sugar Loaf Mt., 600 m., on rocks in forest of upper slopes, F. C. Deighton 4441. USNEA HISPIDA Motyka, Lich. Gen. Usnea Stud. Monogr. 589. 1938. Usnea barbata v. florida £. australis Vainio, Cat. Welwitsch Afric. Pl. 2:396. 1901. Type: Cameroons, Huea, arboricole, Lemperch. Thallus about 15 cm. tall, rigid, becoming dark ashy fuscous in the herbarium, the smaller branches somewhat paler, opaque, base short, rigid, closely annulate, repeatedly sympodially branched at right angles; branches straight or slightly curved, articulate, joints inflated up to 3 mm. in diameter, papillae quite dense, irregular, conic, acute, the larger joints partly paler and rarely almost. epapillate; cortex 100 p thick, hard, slightly fuscous; medulla about 600 р thick, loosely [Vor. 40, 384 ANNALS OF THE MISSOURI BOTANICAL GARDEN woven, white, K—; axis 500 pu in diameter; ramuli close, perpendicular, 2-10 mm. long, attenuate at the base, thicker in the middle, apices smooth, the larger tubercu- late and bearing apothecia. Apothecia numerous, rarely beyond 10 mm. in diameter, cupuliform; exciple smooth or with some large tubercles on the larger apothecia; cilia few, usually 4—8 per apothecium, broad at the base, conic-attenuate above, smooth; disc concave to plane, chalky white-pruinose; ascospores hyaline, uni- locular, 11 X бы. Sierra Leone: Sugar Loaf Mt., 650—775 m., F. C. Deighton M4440. Cameroons: [Victoria], Mrs. G. Thomson. BLASTENIACEAE Thallus crustose, indeterminate or effigurate, attached to the substrate by hyphae of the prothallus or of the medulla, heteromerous, rarely homoeomerous; ecorticate or with a fastigiate cortex which rarely appears pseudoparenchymatous; algae protococcoid; sometimes corticate below in Gasparrinia and Kuttlingeria. Apothecia round, sessile or immersed, biatorine, lecideine or lecanorine, usually with an algal layer under the hypothecium; epithecium granular or powdery, usually containing chrysophanic acid (producing a purple or violet color with potassium hydroxide); paraphyses simple, septate, tips usually characteristically thickened; asci normally 8-spored, fewer in some species of Bombyliospora; ascospores hyaline, thick-walled, usually polari-2—4-locular (unilocular in Protoblastenia and Ful- gensis, plurilocular with rounded protoplasts in Bombyliospora). There is no agreement among lichenologists on the number of genera to be recognized in this difficult family. Bombyliospora is so distinct that it is generally recognized as a genus, although some might not include it in this family but leave it in the Lecideaceae. Some would recognize only Caloplaca or one of its synonyms for the other genera, while I am inclined to recognize all of the genera in the fol- lowing key. Since many species have been described in one genus and transferred elsewhere, depending on the number of genera recognized by an author, I have prepared keys to all the species described from tropical Africa and sorted them into the genera which I recognize, but without making new combinations in order not to add to the synonymy. The whole family is badly in need of revision, especially the temperate species. 1. Apothcecia biatorine, but algae ma ay occur beneath the hypothecium........1.. eorr tttm et tene 2 1. Apothecia lecideine; thallus crustose, indeterminate; ascospores ет иеа Dodge & = 1. Apothecia lecanorine 2. unilocular Protoblastenia pr 2. Ascospores: р 3 2. Ascospores 3—4-locular . атріа 2. Tg dei gee pee protoplasts ыы) or rounded.............. Big DNI 3: » indeterminate, usually ec -Blastenia M. 3; Thallus ique usually corticate ee Trev. 1953] DODGE—LICHENS OF TROPICAL AFRICA 385 4. Ascospores unilocular, thallus effigurate............................. sss .Fulgensia Mass. & sti 4. Ascospores Hao bilocular 4. Ascospores 3 (—4)-locular; ta ed КСЕ еШ ior. Mox onc ds i 5. Thallus indeterminate, usually ecort mia Mass. 5. Thallus effigurate, usually corticate геи above and below... abest Tornab. BOMBYLIOSPORA BoMBYLIOSPORA DNtrs. in Mass., Ricerche Autonom. Lich. Crost. 114. 1852. Psorothecium sect. Bombyliospora Stzbgr., Ber. Thatigk. St. Gall. Naturw. Ges. 160. 62. Lecania sect. Platylecania Müll. Arg., Flora 65:327. 1882. Type: Lecidea tuberculosa Fée. Thallus epiphloeodal, indeterminate; cortex decomposed, of periclinal hyphae; algae Trebouxia or protococcoid; medulla of periclinal hyphae, rather compact. Apothecia sessile, constricted at the base, large, margin persistent; amphithecium absent; parathecium light-colored, not carbonaceous, pseudoparenchymatous from radiating thick-walled hyphae; hypothecium hyaline or darkened, of periclinal, slender hyphae; paraphyses slender with clavate to pyriform tips; asci 1—8-spored; ascospores large, hyaline, ellipsoid to oblong, septate with rounded or lenticular protoplasts. 1. Ascospores under 46 р long ....... 2 1. Ascospores o dris a at ng 5 2. Ascospores 4-locular, 30-46 X 14-22 p; asci 4-spored; RS rather dark colored; São Thomé thomensis (Му 1.) — cospores 6—8-locular ......- = 3. Азсозрогез 28-30 X 11-12 и; asci 8-spored -...... 3. Азсозрогез 20-30 X 9-13 u; Tanganyika...... inguensis v. intermedia Sones Arg.) йй. B. dom 4. па Вузів thecium 65 ш tall; = dark fuscous, margin orange; thal В. “nigeriens сені Dodge o 80 u thick; Nigeria 4. Ес ат wish; "thecium 90-95 ш tall; disc orange or tawny, margin paler; thallus 100 д thick; Tanganyika ..........—....——-——m B. cerinella — m pale X E S 22 Е ча мл . Hypothecium dark ........... лее een eren tenente tenente 6. Thallus cervine — isidio-tartareous; disc very dark brown; ascospores solitary, ET BO X 20 д; Tanganyika Mem . Meyeri Stein 6. "Thallus grayish olive e, мтят verrucose; disc orange, margin white; asci 3—4-spored; : ascospores 67—70 X 12-14 ш; Nig OLE ИНЕС ИРИ E Thoroldi Dodge 7. Thallus pale glaucous, CoE he nente моей + Ms ite M 7 dated. изн, u roba solitary, about "apart е к ure a d aiam в „В. endoleucites (Stirton) Zahlbr, Ў: ; Thallus pale erre rugulose granulose, fa: arinose; disc pruinose thea f MEM PT thec а а fulvous to rufo-fuscous; ascospores 90 X 23 p; asci (2—) 4-spored; Tanganyika ненна at oc миин ные не .B. pruinata ы Arg.) Zahlbr. Juan 7. Thallus. din green, rugulose and rper granulose; disc fuscous black; fuscous; г Bomsy.iospora nigeriensis Dodge, sp. nov. Type: Nigeria, Ondo Province, Ipetu, on Theobroma, C. A. Thorold 106. Thallus epiphloeodes, rugulosus vel laevis, continuus, ad 80 p crassitudine, ob- scure olivaceo-alutaceus; cortex 15 р crassitudine, decompositus gelifactusque, hyphis periclinalibus; stratum algarum 20 p crassitudine, cellulis Trebouxiae, 3—4 p diametro; medulla ad 45 p crassitudine, hyphis conglutinatis periclinalibus inter- [Vor. 40, 386 ANNALS OF THE MISSOURI BOTANICAL GARDEN textis. Apothecia solitaria vel aggregata, rare 3—4 concrescentia, basi constricta, 0.7-1 mm. diametro, margine aurantiaco, integro, elevato dein disco aequante; disco brunneo; parathecium 120 р crassitudine, strato extero 15 p crassitudine minutis cum crystallis aurantiacis, aliter hyalinum, hyphis radiantibus, pachy- dermeis, conglutinatis, septatis, pseudoparenchyma formantibus; hypothecium hyalinum, 40 p crassitudine, hyphis periclinalibus, 3—4 u diametro, pachydermeis; thecium 65 и altitudine; paraphyses tenues, pachydermeae, super ascos dichotome ramosae, apicibus clavatis cum crystallis brunneis tectis; asci ellipsoidei, 50—55 Х 15 и; ascosporae octonae, hyalinae, ellipsoideae vel subfusiformes, protoplastis lenticularibus, 8-loculares, 27-30 X 12 y. Thallus epiphloeodal, rugulose or smooth, continuous, up to 80 y thick, deep olive buff; cortex about 15 џи thick, decomposed and gelified, of mostly periclinal hyphae; algal layer 20 u thick, cells of Trebouxia, 3—4 и in diameter, closely packed or more scattered; medulla up to 45 p thick, of conglutinate, interwoven, mostly periclinal hyphae. Apothecia solitary or aggregated, rarely 3—4 con- crescent, constricted at the base, 0.7-1 mm. in diameter, margin orange, entire, elevated at first, then level with the flat, amber-brown disc; parathecium 120 p thick below and on the sides of the thecium, the outer 15 u orange with abundant aggregates of minute crystals, the rest hyaline, of radiating, conglutinate, thick- walled, closely septate hyphae 4—8 p in diameter, forming a pseudoparenchyma; hypothecium 40 u thick, hyaline, of thick-walled, mostly periclinal hyphae 3—4 в in diameter; thecium 65 y tall; paraphyses slender, thick-walled, dichotomously branched above the asci, tips slightly clavate, obscured by masses of brownish crystals to a depth of 15 y in the epithecial gel; asci ellipsoidal, 8-spored, 50-55 X 15 p; ascospores hyaline, ellipsoid to subfusiform, protoplasts lenticular, 8-locular, 27-30 X, 12 y. This species belongs in the group with B. cerinella Zahlbr. from which it differs in its thinner thallus, orange margin, fuscous to almost black disc, lower thecium, hyaline hypothecium, and more septate ascospores. B. thomensis (Nyl.) Zahlbr. from São Thomé differs in having asci about 4-spored, ascospores larger, 30-46 X 14—22 р, only 4-locular. Nigeria: Ondo Province, Ipetu, C. A. Thorold 106, type; Oyo Province, Iseyin, C. A. Thorold 107; Moor plantation near Ibadan, C. A. Thorold 105; all on Theobroma. Bompy.iospora Thoroldi Dodge, sp. nov. Type: Nigeria, Oyo Province, Iseyin, on Theobroma, C. A. Thorold 108. Thallus epiphloeodes, rugulosus, verrucosus, continuus, 40 p crassitudine, griseo- olivaceus; cortex 15 p crassitudine, hyphis tenuibus periclinalibus conglutinatis; stratum algarum 12-15 y crassitudine, cellulis Trebouxiae, 3—4 p diametro, dense dispositis; medulla 10—13 н. crassitudine, hyphis tenuibus dense contexta. Apothecia 0.5—0.8 mm. diametro, basi constricta, margine tenui, albo, disco plano, cin- namomeo-alutaceo; parathecium hyalinum, 60-85 p crassitudine sub hypothecio, 1953] DODGE—LICHENS OF TROPICAL AFRICA 387 ad 40 р superne tenuescens, hyphis radiantibus, pachydermeis, 4 и diametro, pseudo- parenchyma formantibus, hypothecium hyalinum vel pallide flavum, 12 y crassi- tudine, hyphis tenuibus periclinalibus contextum; thecium 105 џи altitudine; paraphyses pachydermeae, conglutinatae, apicibus clavatis vel pyriformibus; asci leptodermei, apicibus non incrassatis; ascosporae ternae quaternaeve, 10-loculares, cellulis intermediis longioribus, protoplastis subcylindricis, 67-70 X 12—14 y. Thallus epiphloeodal, rugulose, verrucose, continuous, 40 р thick, grayish olive; cortex 15 p thick, cells of Trebouxia, 3—4 и in diameter, closely packed but some- times interrupted by strands of medulla; medulla 10-13 y thick, of slender, closely interwoven hyphae. Apothecia 0.5—0.8 mm. in diameter, constricted at the base, margin thin, white, disc plane, cinnamon buff, both watery white when moist; parathecium hyaline, 60-85 p thick below the hypothecium, thinning to 40 p thick above, of radiating, thick-walled hyphae about 4 и in diameter, forming a pseudoparenchyma; hypothecium hyaline or pale yellowish, 12 p thick, of slender, mostly periclinal, interwoven hyphae; thecium 105 y tall; paraphyses thick-walled, conglutinate, tips clavate to pyriform, ending in the pale yellowish epithecial gel; asci 3—4-spored, thin-walled, tips not thickened; ascospores hyaline, 10-locular, middle cells longer, protoplasts cylindric with corners rounded, septa and wall thick, 67-70 X 12-14 р BLASTENIA BLASTENIA Mass., Atti I. В. Ist. Veneto II, 3:append:101. 1852; Flora 35:575. 1852 Type: B. sinapisperma (Lam.) Mass. For discussion, see Dodge & Baker, Ann. Mo. Bot. Gard. 25:611. 1938. Thallus crustose, indeterminate, powdery, granulose or rimose, attached to the substrate by the hyphae of the prothallus or of the medulla, homoeomerous or heteromerous; ecorticate or with slightly developed cortex; algae protococcoid. Apothecia round, immersed to sessile; parathecium well developed; amphithecium absent, epithecium granular or powdery, K violet or purple; hypothecium hyaline; paraphyses simple, septate, capitate; asci 4—16-spored; ascospores hyaline, ellipsoid, polari-bilocular. Spermogonia immersed, spherical; spermatiophores septate; spermatia short, cylindric, straight. In the following key the species with black apothecia probably belong in Huea, but in the absence of material, I have not wished to make the transfer. Blastenia Brebissonii у. microspora evidently belongs in Xanthocarpia. “1. Ascospores 3-locular, 20-24 X 12-13 и; Angola......B. Brebissonii v. microspora (Vainio) эн. 1. Ascospores 2-locular Le ниетине г ry: таги nm granulose; tpothecis жн — 22 9908,00 тавлар tm aD dé. AZ IN "i h , 0.3—0.6 mm.; ИЕ 12-14 X Ис кч. ochres apoi dens Enan a ae B. ر‎ e) Zahlbr. black when dry, 0.2-0.3 mm.; ascospores e Era ee ty vivement ac /5 у Ут An 2. Thallus white ا‎ ы [Уот. 40, 388 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. Disc ferruginous-fuscous or dark rufous, blackening; saxicole 4 3. Disc flesh-yellow; apothecia 0.5 mm.; ascospores 8-11 Х 4-7 ш; Ilha Ргіпсіре............ albidopallens (Nyl.) Zahlbr. is, Кепуа сы laetebunda (Hue) Zahlbr. ‚ Disc orange tawny; apothecia 0.2-0.4 mm.; ascospores 14—16 X 6-8 д; Mauretania... B. sordida (Hue) Zahlbr. 4. Thallus granulose; apothecia 1 mm.; ascospores 14—16 X 7-8 и; Cormoro Archi- pelago B. cormorensis Lindau 5 B - Disc orange-red; apothecia 0.5-1.2 mm.; ascospores 15-22 X 8—12 w w 4. alus rimulose areolate . Apothecia fuscous black, plane then convex to subspherical, pedicellate; St. Helena.... B. lactescens (Leight.) Zahlbr. 5. Apothecia fuscous, plane, margin black; apothecia 0.5-0.8 mm.; ascospores 15—1 6 и; Ilha Principe B. albidofusca (Nyl.) Zahlbr. ‚ Apothecia cinnamon fuscous, plane, margin black, 0.4 mm.; ascospores 11—13 X 6—6.5 и; Kenya B. polioterodes (Steiner) Zahlbr. мл л PYRENODESMIA PYRENODESMIA Mass., Atti I. В. Ist. Veneto 1L 3:119, 1853. Callopisma DNtrs., Giorn. Bot. Ital. II, 2:198. 1847, non Martius, 1827 (Gentia- naceae). Caloplaca 'Th. Fr., Lich. Arctoi, 218. 1860, pro parte. ` Type: For discussion, see Dodge & Baker, Ann. Mo. Bot. Gard. 25:619. 1938. Thallus crustose, attached to the substrate by the hyphae of the prothallus or of the medulla, without rhizinae, uniform, mostly yellow and K purple; hetero- merous, ecorticate or nearly so; algae protococcoid; medulla of loosely woven, thin-walled hyphae. Apothecia round, appressed or sessile, seldom immersed, lecanorine with a well-developed amphithecium containing cortex, algal layer, and medulla; epithecium granulose to powdery, usually K violet or purple; hypo- thecium hyaline, lying above the algal layer; paraphyses simple, septate, capitate; asci 8-spored; ascospores hyaline, ellipsoidal to fusiform, polari-bilocular. Spermo- gonia immersed, with a hyaline wall; spermatiophores closely septate; spermatia short, straight, cylindric. In the following key C. is the abbreviation for Caloplaca. 1. Apothecia immersed or nearly ѕо 2 1. Apothecia emersed to sessile and constricted at the base; disc orange, or fulvous 4 2. Thallus dark olive, subsquamulose; apothecia 0.3—0.5 mm.; ascospores 10-14 X SNL LS re: dg ОО C. asmarensis Jatta 4-7 р; 2. Thallus white or nearly so, chalky, areolate; disc black; ascospores 14-16 X 7-8.5 u; auretania zx 3. Thallus margin ciliate, areoles all about the same size and shape......C. concinnata (Hue) Zahlbr. es very variable in size and shape....C. inconcinna (Hue) Zahlbr. 4. allus fuscescent, disc ferruginous, blackening; ascospores 11—14 Х 3.5 и; Ango C. benguellensis (Nyl.) — w 3 = g a H 5 9 pee 5 8. = = э LÀ ч 8. 4 М batt iain til cere ak ЕН 4. -Thallus white to dp i 1. $$ — 4. Thallus ferruginous е orange; apothecia and ascospores small 6 ow Ач d 5. Thallus granulose, disc ochraceous orange; Abysinia TT C. Odoardi (Bagl.) Zahlbr. 5. Thallus subsquamulose but not effigurate, disc deep orange; ascospores 12-14 us Mauretania li (o E S P. conglobata (Hue) Dodge 6. Thallus rimose-areolate; Mozambique. Т" C. zambesica (Müll. Arg.) Zahlbr. 6. Thallus powdery OTAD IL UL UU eee . exasperata (Bagl.) Zahlbr. 7. Thallus areolate, subeffigurate at margin; saxicole; Mozambique [may belong in Gaspar- Zahlb Е e m Aime ee о ICT ONLUS IDIBUS MOE S Тула, ü ы 3 E 7. Thallus squamulose Е о а ол — 8 1953] DODGE—LICHENS OF TROPICAL AFRICA 389 8. Squamules 0.2-0.3 mm. on black hypothallus; apothecia 0.2—0.5 mm.; ascospores 12-16 X 6-8 и; Uganda C. citrinella (Jatta) Zahlbr. 8. Squamules 0.4—0.7 mm. on pale hypothallus; apothecia 0.4—0.5 mm.; ascospor 15-16 X 6—6.5 u; S. Sahara C. asekremensis (Hue) Zahlbr. PYRENODEsMIA conglobata Dodge, comb. nov. Lecanora (Pyrenodesmia) conglobata Hue, Mém. Soc. Bot. France 30:7. 1917. Caloplaca conglobata Zahlbr., Cat. Lich. Univ. 7:110. 1930. Type: Mauretania, Boulanouar, on bark of Acacia tortilis, Chudeau. Thallus epiphloeodal, about 1 cm. in diameter, whitish, areolate, areoles quite convex, margin indeterminate, white-arachnoid on bark, but black, about 0.1 mm. wide where growing over the margin of Ionaspis ascidioides; cortex 16—20 p. thick, pseudoparenchymatous, heavily nubilated with minute yellowish crystals; algal layer 30-40 м thick, of closely packed, protococcoid cells 5—6 p in diameter; medulla scarcely differentiated, but closely septate, much branched hyphae pene- trate the cork cells and more or less disorganize them to a depth of 550 м. Apo- thecia sessile, somewhat constricted at the base, urceolate when very young, becom- ing plane, the margin not elevated at maturity, disc burnt sienna, flat, margin flame-scarlet; amphithecium 135 м thick below the margin of the thecium, thinning to 65 p at the top, of the same structure as the thallus, algae in large irregular colonies, 40-55 м in diameter, extending to the top of the thecium when young but soon compressed to a zone below the margin of the thecium when mature by the expanding, hyaline parathecium which is about 15 р thick below, expanding to 65 р thick above, the outer 8 и heavily nubilated with orange granules, of con- glutinate, slender, thick-walled hyphae and forming a stipe below reaching the bark, 135 y in diameter; hypothecium 40 и thick in the center, thinning to the margin, of closely woven, slender hyphae becoming subvertical just below the thecium; thecium about 100 y tall; paraphyses slender, conglutinate, repeatedly dichotomous above the asci, tips moniliform, 3 д in diameter, heavily nubilated with orange crystals to a depth of 15 р; asci 55 X 15 p, ellipsoid-clavate, 8-spored; ascospores 12 X 5.5 m, protoplasts spherical, terminal, 3 д in diameter, connected by an isthmus, hyaline. At maturity this species might be mistaken for Blastenia, as the algal zone is confined to a ring around the base of the thecium, while all the tissue on the sides of the thecium is parathecial. Nyasaland: Kasungu Hill, 1100 m., corticole, L. J. Brass 174582, Vernay Nyasaland Expedition, 1946. GASPARRINIA GasPAnRINIA Tornabene, Lichenogr. Sicula 27. 1849. Ampbiloma Koerb., Syst. Lich. Germ. 110. 1854. Aglaopisma DNtrs. in Bagl., Mem. Accad. Sci. Torino II, 17:396. 1856. Туре: С. murorum (Нойт.) Tornabene. (For discussion, see Dodge & Baker, Ann. Mo. Bot. Gard. 25:622. 1938.) [Vor. 40, 390 ANNALS OF THE MISSOURI BOTANICAL GARDEN Thallus crustose, effigurate or lobed and subfoliose, mostly yellow, K purple, heteromerous; corticate on both surfaces, cortex pseudoparenchymatous, cells thin- walled; algae protococcoid; medulla arachnoid, of thin-walled hyphae. Apothecia round, appressed or sessile, lecanorine; amphithecium containing cortex, algae, and medulla; epithecium granulose or powdery, usually K purple or violet; hypothecium hyaline, lying above the algal layer; paraphyses simple, septate, capitate; asci 8- spored; ascospores hyaline, ellipsoidal, polari-bilocular. Spermogonia immersed with a thin hyaline wall; spermatiophores closely septate; spermatia short, straight, cylindric. In the following key C. is the abbreviation for Caloplaca. 1. Thallus vitelline to мән yellow 2 1. Thallus vae зын гед 3 2. Tha m hin, ы narrow lobed, center Yerencifocm; ascospores 11—18 urorum ы gremliformis (in) Zahlbr. 2i Thallus thin, linear lobes with —— ie. apothecia - ospor 6 maritime rocks; Angola C. fla on чал N yi L) Za 2. Thallus viteline, lobes crisped, margins contiguous; Vani 1 Ы C. crispicans (Nyl.) Zahlbr 3. Apothecia unknown; thallus margins tuberculate, miniat Apothecia iL. 4. Areoles convex, small; Socotra vus granulifera (Mill. we Zab 4. Areoles plane, larger; Somaliland oc ditatus — rg.) Z 5. Apothecia innate or nearly so, small disc miniate; POR RU ا‎ mpbiloma debanense pd 5 sr a emersed to sessile Corticole; apothecia small; Abyssinia C. Beccarii (Bagl.) Zahlbr. ет 7. Азсозрогез 7-10 X 2-3 и; apothecia 0.3—0.5 mm., adnate; Eritrea C. delicata Jatta 7. Ascospores 6—8 u broad br ms 8. Ascospores 8-11 x f и; ненин 0.7—0.9 mm.; Angola......C. elegantissima (Nyl.) Zahlbr. 8. Ascospores 12 X 7 д; Socotra C. deplanata (Müll. Arg.) Zahlbr. 8. Ascospores 12—14 X pt. шз apothecia 0.5-1 mm.; Socotra....C. Balfourii (Müll. Arg.) Zahlbr. BUELLIACEAE Thallus crustose to squamulose, simple or effigurate, without rhizinae, attached to the substrate by hyphae of the hypothallus or of the medulla; cortex dern evanescent in some species; algae protococcoid; medulla loosely woven, of t walled hyphae; sometimes sorediose. Apothecia round, immersed to sessile, lecideine or lecanorine; paraphyses simple or branched above; asci normally 8-spored; asco- spores smoke gray becoming brown, 2—4-locular or dwarf-muriform by a longi- tudinal division of one or more of the middle cells, usually with a thick wall but without a halo as in Rhizocarpon. Spermatia short, straight. Usually the family is divided into two genera; Buellia with lecideine apothecia (parathecium highly developed and carbonaceous), and Rinodina (parathecium only slightly developed and hyaline, amphithecium well developed). Only Buellia is represented in our material. BUELLIA BUELLIA DNtrs., Giorn. Bot. Ital. II, 1:1:195. 1846. Type: Of the three species originally included, we may eliminate B. canescens (Dicks.) DNtrs., as it belongs in sect. Diploicia which may deserve generic rank. Of the two remaining, Clements & Shear (Gen. Fung. 323. 1931) have chosen B. parasema DNtrs. 1953] DODGE—LICHENS OF TROPICAL AFRICA 391 Thallus crustose, simple, margin sometimes effigurate (sect. Difloicia), at- tached to the substrate by hyphae of the hypothallus or of the medulla, without rhizinae; cortex fastigiate, often evanescent, rarely pseudoparenchymatous; algae protococcoid; medulla of interwoven thin-walled hyphae; occasionally sorediose. Apothecia appressed to sessile (immersed in sect. Melanaspicilia), lecideine, blac unless the disc is pruinose; amphithecium absent; parathecium entire or dimidiate, carbonaceous; paraphyses capitate, epithecium dark; asci usually 8-spored; asco- spores brown to black, ellipsoid, bilocular (in sect. Eubuellia) or 4-locular to dwarf-muriform (in sect. Diplotomma), with thick walls but without a halo (distinction from Rhizocarpon sect. Catocarpon). In sect. Melanaspicilia, the apothecia are immersed in the thallus but its well- developed carbonaceus parathecium and small bilocular ascospores clearly relate it to ВиеШа rather than to Rinodina where it was placed by Zahlbruckner. 1. Apothecia innate or эн: ^ saxicole; MELANASPICILIA 2 1. Apothecia sessi PA pi rats 4 2. Thallus wh perta neos 12 X 6 p; Socotra B. leucina Müll. Arg. 2. Thallus Даны " veta clay imose-aerolate. 3 3. Apothecia 0.3—0.4 mm.; ascospores ei X 5.5—6.5 ш; thallus margin ee Ocotra innata Müll. Arg. 3 Apothecia 0.3—0.4 mm.; ascospores 10—15 X 6-7 ш; thallus pî EIS peaini B. subimmersa Müll. Arg. 3. Азсозрогез 10-12 X 6-7 д; thallus K yellow, black-margined; Angola В. spuria v. ferruginea eee Thallus indistinct, evanescent or not described Thallus subcaerulescent; ascospores 10-12 X 5-6 и; spatial Mauretania.................. Chaudeauiana sree Zahlbr. 4. Thallus rose-color; apothecia 0.8-1 mm.; ascospores Pa inh X 5-6 ш; cortic B. Tobleri — B Usambara 4. Thallus olive fuscous; saxicole 4 Thallus — to clay color м ь allus white to ashy ог glaucous greenish 11 5. Medulla paths ascospores 14-18 X 6-9 д; corticole; Angola....B. disciformis у. rhodina Vainio Medulla white 6 6. Disc white-pruinose; thallus K—, C red; saxicole; Abyssinia B. Caldesiana v. subpruinosa мо. мл Disc not = позе : 7. Геге жуз 0.3 mm.; ascospores 14—16 X 7.5—8.5 Us gnicol 4 isciformis т, oblong (Müll. Arg.) Zahlbr. 7. Apothecia 0.7 mm.; ascospores 24—34 X 12-16 и; ; ramulieole Kenya : : : i isciformis v. pachyspora Zahlbr. 8. Apoth 0.16—0.25 mm.; ascospores 10—13 X 7-8 и; Soc SEE I B. Нана f. A sni = Arg. 8. Apothec .; ascospores 10 Ж 5-5.5 и; Mozambique............-.------ üll. Arg. i Tents Lx. velles; rimulose, black-margined; реки s X "yug ui gres pacbyospora Mass. Th "Um ll hy; h- ose, margin contorted; corticole; Socotra............ m 4 disciformis v. contorta (Mall, Arg.) данне. хо = 9. Thallus ochraceous to clay-color 10. — 0.25—0.3 mm.; те 8—11 X 4.5-5.5 ш; saxicole; Nyamnyam чот, ые Sudan ngo. Mu CI OH, T т, afra Vainio 10. 0.3 es ii 5 n; corticole; Mozambique...... ът —0.6 mm.; vane аа Vainio 11. Thallus gray- green to pale greenish olive 11. Thallus white = Lus dabis MANCA An Е 12. Ascospores 24 5 X 10-11.2 и; thallus ashy green to green, granulose, prothallus ashy black; pis чы 0.3—1 mm., disc bluish-pruinose; corticole; Usambara........ , B. crassa Riehmer 13 12. Ascospores smaller, 11-15 X 5-7.5 № [Vor. 40, 392 ANNALS OF THE MISSOURI BOTANICAL GARDEN 13. Thallus subsquamulose, greenish gray; apothecia 0.5—0.75 mm.; terricole; Ruanda........ B Wet Zahlbr. 13. Thallus —Ó Mans grayish olive; apothecia 0.5 mm. or aggregated in pact gro n diameter; corticole; var Le B. wi oan Dodge 13. Thallus s pia ott SR green; apothecia 0.16—0.25 mm.; кА substigmatea Е Arg. 14. Metalls cinnabar red; thallus of tumid, bullate areoles, chalky ai е. уегу со eer ruinose; saxicole; Abyssinia a p ^ Me sdulla 15 Lipsio Pe poorly developed; Woe 0.3 mm.; ascospores 14-16 B. disciformis v. oblongata (MS УЗ Б 15. Corticole 15. Saxicole i 16. Asci 12-spored; ascospores 15.5-17 X 5—6.5 д, each locule often чен» ы then appearing falsely 4-locular; thallus black-margined; Е < Snir hn: v. pata is Vainio 16. Asci 8-spored 1 17. Ascospores eth х сн и; Mozambique.................. B. americana v. palmensis (Vainio) es 17. Asc is 18. «ы magn EL disc black; ascospores 13 X 7 и; == gs е, хаваны ЛЫ "Mall. A 18. era ies АСЕ 19. Ascospores 9-12 X 4—5 ш; thallus rimose, areoles convex, «уно К огап St. Helena approximant 75) Zahlbr. 19. Ascospores 12-15 X 5-5.5 д; thallus areolate, whitish; Sier T ее. oni Dodge 20. ptem black, areoles white, angular, scattered or partiy contiguous; Sec white, Са at first; parathecium dimidiate; ascospores in B. po Malt Arg. 20. Нуна» Dicke eise d Reread the assimilative thallus; Бы 0.3 m В. prosperens "Mill. My 20. YÎ ea or m a very narrow black margin 21. Ascospores 8-9 X 4-5 u 2 21. Ascospores 8—13 X 4—6 д; thallus areolate, white, К yellow; apothecia small; Abyssini B. italica v. еле is M. 21. Ascospores 11—16 21. doner 14-16 X 7. d 5 д; thallus rugost-areolate, тинч ~ greenish-pruino: v. subaeru, uginosa "Mall, Arg. 21, Ade 15-17 X 7-8 ш; thallus ashy, er K К yellows Анка, TM B. dis v. de rr Bagl. ispers 22. irons 0. Fe : mm.; thallus white, K slightly yellowish, ish a narrow black stellulata E subtilis Vainio ч Jatta 22, Poetics Meg з mm.; thallus ashy; Eritrea 25. oe 0, 5 mm.; thallus white, rimulose with а narrow black margini as B. delaevata yh) Zahlbr. 23. Арин б 0.4—0.8 mm.; thallus chalky white, essaie" without a conspicu gin; Angola B. subalbula (Nyl.) "Mill. Arg. ВовмлаА Adamesii Dodge, sp. nov. Type: Sierra Leone, Sefadu (Gbense), on trunk of Elaeis guineensis, P. Adames, comm. F. C. Deighton M4755. Thallus epiphloeodes, rimoso-areolatus, ca. 130 p. crassitudine, pallide griseo- olivaceus; cortex non bene evolutus; stratum algarum 25 y crassitudine, filamentis cylindricis dense dispositis Trenfepobliae?, 4—5 р diametro, cellulis cylindricis aut apicibus rotundatis inter hyphas verticales, 2 и diametro; medulla ca. 105 p crassi- tudine, hyphis verticalibus tenuibus, plus minusve granulis inspersis, magis peri- clinalibus et compactis in strato inferno 15-20 p crassitudine. — Apothecia orbicularia, sessilia, basi constricta, solitaria 0.5 mm. diametro aut aggregata in gregibus ad 2 mm. diametro, nigra; parathecium inferne 80—95 p crassitudine ad 1953] DODGE—LICHENS OF TROPICAL AFRICA 393 10 р superne tenuescens, pseudoparenchymaticum, cellulis leptodermeis, radiantibus, 6-7 p diametro, brunneum; hypothecium 25 y crassitudine, brunneum; thecium 105—120 д altitudine; paraphyses tenues, subdichotome ramosae, cellulis terminali- bus subsphericis nigro-brunneis; asci subcylindrici, 55 X 14 и; ascosporae octonae, imbricatim monostichae, biloculares, brunneae, 14—15 X 5-6 p. Thallus epiphloeodal, rimose-areolate, about 130 д thick, light grayish olive; cortex not differentiated; algal layer 25 и thick, of closely packed, vertical fila- ments of Trentepohlia? 4—5 y in diameter, cells cylindric or with rounded ends, between vertical hyphae 2 u in diameter; medulla about 105 м thick, of slender, mostly vertical, loosely woven hyphae, more or less inspersed with granules, be- coming compact and periclinal in the lower 15-20 p. Apothecia round, 0.5 mm. in diameter when solitary, often aggregated into groups up to 2 mm. in diameter, black, constricted at the base; parathecium 80—95 и thick below the hypothecium, thinning to 10 д at the top of the thecium, of radiating, thin-walled pseudo- parenchyma, cells 6-7 и in diameter, brown, the outer 15 р very dark brown; hypothecium about 25 p thick, dark brown; thecium 105-120 y tall; paraphyses slender, not dense, somewhat dichotomous, terminal cells dark brown, subspherical, 3 p in diameter; asci 8-spored, subcylindric, 55 X 14 р; ascospores imbricately monostichous, bilocular, deep brown, not or slightly constricted at the septum, 14-15 X 5-6 p. The systematic position of this species is somewhat uncertain. The algae seem to be filaments of Trentepohlia, unless they are protococcoid and so distorted by mutual pressure as to appear filamentous, a condition occasionally seen in some species of the Verrucariaceae. If the filaments are Trentepohlia, the lichen may be related to Catinaria Vainio, which has hyaline ascospores. The often-elongate and variously curved (sometimes lirelliform) apothecia of the crowded aggregates closely resemble those of Encephalographa (with Palmella algae) and are formed in the same way as those of E. cerebrinella (Nyl.) Zahlbr. from Kerguelen. The loosely arranged paraphyses are also very unusual in Buellia. The first apothecium is round and solitary, about 0.5 mm. in diameter. After the ascospores are shed, the thecium disintegrates. From several points near the margin, or more irregularly scattered over the disc, new apothecia proliferate until the parathecia touch but do not fuse, and by mutual pressure assume very irregular shapes until the aggregate, seen from above, somewhat resembles a member of the Chiodectonaceae such as Sarcographa. In some of the larger groups (not sectioned) there is a suggestion of a third proliferation. There is also a possibility that the species might be referred to Rinodina, as 1 have seen an occasional algal cell in the outer portion of the tissue which I have called the parathecium, clearly apothecial in origin, not thalline. The ascospores more closely resemble those of Buellia. [Vor. 40, 394 ANNALS OF THE MISSOURI BOTANICAL GARDEN Bue. Deightoni Dodge, sp. nov. Type: Sierra Leone, Njala (Kori), on Ampbimas pterocarpoides, F. C. Deighton M4334B. Thallus epiphloeodes, albidus, subverrucosus, margine indistincto, usque ad 55—65 p crassitudine; cortex decompositus, gelifactus, 8—16 д crassitudine, superne granulis inspersus; stratum algarum 27—40 p crassitudine, cellulis Trebouxiae 5—6 p diametro; medulla hyphis 2 и diametro. Apothecia 0.3—0.4 mm. diametro, nigra, margine integro, tenui, subelevato, disco convexo, nigro, nudo; parathecium dimidiatum, 16 д crassitudine, hyphis pachydermeis nigro-brunneis; hypothecium centro 55 p crassitudine ad marginem tenuescens, hyphis brunneis, 4 и diametro; thecium 55—65 и altitudine; paraphyses tenues, super ascos dichotome ramosae, apicibus capitatis, cellulis apicalibus sphaericis, obscure brunneis, cellulis penultimis pyriformibus, pallide brunneis; asci cylindrico-clavati, 30 X 12 p; ascosporae octonae, brunneae, biloculares, anguste ellipsoideae subcylindricaeve, ad septum non constrictae, 12—15 X 5—5.5 y. Thallus epiphloeodal, margin indeterminate, slightly verrucose, up to 55—65 p thick above the outermost bark cells, whitish; cortex decomposed, gelified, 8—16 p thick, outer portion with minute granules; algal layer 27—40 y thick, of colonies of Trebouxia, cells 5—6 u in diameter, separated by strands of vertical hyphae 6-8 р wide; medulla of hyphae about 2 p in diameter, blackening and disorganizing the outermost bark cells, then forming a hyaline layer about 55 д thick, of loosely interwoven strands of hyphae between fragments of disintegrating hyaline bark cells, with an occasional whole cell or small group of bark cells, some strands of hyphae penetrating between the brown bark cells to a distance of 120 д. Apothecia 0.3—0.4 mm. in diameter, black, margin thin, entire, slightly elevated, disc nude, black, becoming very convex; parathecium dimidiate, 16 u thick, of dark brown, thick-walled hyphae, carbonaceous in thick sections; hypothecium about 55 p thick in the center, thinning to the margin, of dark brown hyphae 4 р in diameter; thecium 55—65 и tall; paraphyses slender, not crowded in the thecial gel, dicho- tomous above the asci, tips capitate, terminal cells spherical, dark brown, penulti- mate cells pyriform, pale brown; asci cylindric-clavate, 8-spored, 30 X 12 p; ascospores brown, bilocular, narrowly ellipsoid to subcylindric, not constricted at the septum, 12-15 X 5-5.5 и. Dimensions are quite variable in different portions of the same thallus and apothecium. I have given maximum dimensions, the minimum being about half those given. After the thecium disintegrates, the base remains white-powdery, so that parts of the thallus appear soraliate. It is not clear whether the thallus con- sists of small areoles, discrete and rounded on an inconspicuous hypothallus, bearing 1-3 apothecia each, or whether the thallus was originally continuous over a large area and the intervening tissue has weathered away. 1953] DODGE—LICHENS OF TROPICAL AFRICA 395 PHYSCIACEAE Thallus foliose, deeply lobed, appressed to the substrate or with ends of lobes erect (rarely completely erect in some species of Anaptychia); rhizinae usually present; corticate above and below (except in some species of Anaptychia); algae protococcoid. Apothecia circular, sessile, lecideine, pseudolecideine or lecanorine; paraphyses simple; asci 8-spored; ascospores brown, septate, usually 2-locular, rarely 4-locular or dwarf-muriform, with thick walls. Spermatiophores septate; spermatia short, straight. PYXINE PyxINE Fries, Syst. Orb. Veg. 267. 1825. Phragmopyxine Clements, Gen. Fung. 84. 1909. Type: Lecidea sorediata Ach. Phragmopyxine was based on P. Eschweileri Tuck. Thallus foliose, deeply lobed, attached to the substrate by rhizinae; upper cor- tex pseudoparenchymatous from conglutinate vertical hyphae; algae protococcoid; medulla thick, of mostly periclinal hyphae; lower cortex often not sharply dif- ferentiated, of periclinal hyphae; soredia common. Apothecia lecideine, at least black and more or less carbonaceous at maturity; epithecium K violet; hypothecium dark; paraphyses simple, conglutinate; азс! 8-spored; ascospores brown, 2(-4)- locular, thick-walled. Spermogonia immersed, mouth dark; spermatiophores sep- tate; spermatia lateral, short-cylindric, straight. Thallus isidiose, o wrinkled, K yellow, medulla x yellow fulvescen P. retirugella f. porrige Mäll. Arg. . corticole; 1. Thallus Thallus without isidia or soredia 5 2. Medulla yellow to pote or tawny 3 2. Medulla w 4 > . Medulla and marginal soredia light yellow; saxicole; Abyssinia "i m P. Meissneri v. sorediosa Müll. Arg. 3. Medull d 11 rficial soredia orange; corticole; Kenya ulla and small superficial soredi g TON raso ЕТ "m 3. Medulla tawny, small soredia white to tawny; lobes 0.5 (-1) mm. sibi: apothecia 0. mm.; ascospores 14—20 X 5-7 y, 2(—4)-loc ular; Perg. Angola......... .P. chry saute Vainio 4. Lobes 3-6 mm. ve eee soralia 0.5—1.5 mm., soredia greenish forem: apothecia 0.5—0.8 ascospores 16—22 X 6-8 ш; corticole; eras s ELE JA. dimorpha Hue 4. Lobes 2—3.5 mm нй; а уе loe; soralia spherical, 2—2. inn м s gr sang ти 2d — " 15-19 7—8 corticole; Frenc est Tro; peer 0.7 1 mm.; ascospores x Bi : Chevalier АИ rficial, white; apothecia not M corticole; 4. Lobes narrower soredia superficial, whi pot aa е} oras наду jd Ass, Thallus is to yellowish gray а Thallus white 6 Fm e color, habit of P. Cocoes; medulla C orange-red; sterile; P alpen ki M мл +6 6. Th ‚ under-side black, е рия apothecia Е allas yello ovi n 220) -X x as 5 м; saxicole; Ann durs adi eh devertens (Ny) Vainio ad thi s a white; sterile; ra 6. Howe olive buff, under er black, without rhizinse, me E n fut ole; Ascospores 17-21 д long; thallus with habit of P. Meissneri, medulla K—; cortic Abys эы ‚Р. endoleuca (Mall. Аг.) Ie M nanm sad [Vor. 40, 396 ANNALS OF THE MISSOURI BOTANICAL GARDEN 8. Lobes plane, much wider than typical P. Cocoes; corticole; Congo....P. Cocoes v. congensis Stein 8. Lobes convex 9. Epithecium greenish, lobes very convex; corticole; Usambara......P. Cocoes v. convexior Müll. Arg. 9. Epithecium fuscous, disc convex, chalky pruinose; lobes slightly convex but never plane or concave; corticole; Socotra P. convexa Müll. Arg. PyxiNE Adamesii Dodge, sp. nov. Type: Sierra Leone, Sefadu (Gbense), on trunk of Elaeis guineensis, P. Adames, comm. Е. C. Deighton M4756a. Thallus foliosus, olivaceo-alutaceus, centro subcrustosus, laciniae subimbricatae, lobis marginalibus subflabellatis, dichotome ramosis, apicibus ad 0.5 mm. latitudine, rotundatis, subpruinosis, 100 д crassitudine, К, С—; soralia elevata, rotundata, ad 0.7 mm. diametro, sorediis flavido-viridibus, granularibus, K—, C—; cortex 16 u crassitudine, pseudoparenchymaticus, hyphis verticalibus, 3 и diametro, cellula supera granulis inspersa; stratum algarum 15 y crassitudine, cellulis protococcoideis; medulla alba, К, C—, ca. 50 y crassitudine, hyphis periclinalibus, 4—5 и diametro, dense contexta; cortex inferior 8 u crassitudine, hyphis nigro-brunneis periclinalibus. Sterilis. Thallus foliose, olive buff, subcrustose in the center, laciniae subimbricate, marginal lobes subflabellate, dichotomous, apices 0.5 mm. or less wide, rounded, slightly white-pruinose, 100 u thick, K—, C—; soralia elevated, 0.7 mm. in diam- eter, soredia yellowish green, granular to subisidioid, K—, С—; upper cortex 16 p thick, pseudoparenchymatous, of vertical hyphae 3 p in diameter, upper cell in- spersed with minute granules; algal layer 16 p thick, cells protococcoid; medulla white, K—, C—, about 50 д thick, of densely woven periclinal hyphae 4-5 p in diameter; lower cortex about 8 p thick, of very dark brown periclinal hyphae. Sterile. The thallus had been overgrown by Parmelia lobulascens у. isidiosissima and most of the algal cells had disintegrated, leaving lacunae in what is obviously the algal layer. PHYSCIA Рнузста Vainio, Etude Lich. Brésil 1:138. 1890. Type: The early history and synonymy of this genus are very involved, and I will postpone their consideration to a later time. Thallus foliose, attached to the substrate by rhizinae, prostrate or outer portions of lobes erect, laciniae narrow, branched, margins sometimes ciliate; upper cortex pseudoparenchymatous from conglutinate vertical hyphae; algae protococcoid; medulla rather loosely woven of mostly periclinal hyphae; lower cortex of con- glutinate, periclinal hyphae. Apothecia lecanorine, amphithecium well developed; parathecium scarcely differentiated (unless the thick black disc under the hypo- thecium represents a modified parathecium in sect. Dirinaria), disc deep reddish brown to fuscous and black, epithecium K—; paraphyses simple, usually septate; asci 8-spored; ascospores 2(—4)-locular or dwarf-muriform, brown. Spermogonia immersed or nearly so; spermatiophores septate; spermatia lateral, long cylindric or somewhat curved in a few species. 1953] DODGE—LICHENS OF TROPICAL AFRICA 397 1. Hyaline below the hypothecium; EuPnvs 2 t HE zo to black layer below the иы ue uat DIRINARIA 8 Thallus at least n pruinose, те apothecia unknown 3 allus not sorediose; apothecia presen 4 3; тыы je? ashy, и K i d above, medulla K—; soralia elevated; P. reti culata Vainio 3. тый sapra white, fuscescent, K—, medulla K-—; soralia 0.5-1 mm. in diameter г; епуа Poncinsii Hue 4. TRIN margins setose, apices of laciniae yellow; Usambara........ P. setosa f. duis Mall. Arg. 4. Harm subcrustose in center, radially wrinkled, ma sian lobes not ciliate; apothecial gin crenulate, шг dark [beau saxicole: GUINEA ае Parmelia incisa n Thallus not as abov 5. Ascospores 25-30 X dirum №; apotheci gin entire, disc dark fuscou pruinose; thallus white, smooth or partly wrinkled, pale Ко, АЙЫ. seed dilatata Nyi. 5, Ascospores 21—24 8—9.5 ш; apothecia 1.5 mm., margin vertically sulcate, disc then black, not рен; thallus m^ to olive buff; "n eco ed ІР. Poncinsii Hue pieds uito" 5. و‎ res not over 22 р, mostly s . Ascospores 16-18 8 X 9-11 u; apot eec 0.4—0.7 тт., margin € crenulate, ur dark fuscous; thallus olive, subcrustose in center, дона short, smooth; Usambar P abbreviata Mall. A 6. Ascospores 12-15 X 7-8 и; apothecia Ven € vega Socot 6. ији Абер шз apot othec argin didis. “tise dark rufous, uinose; — үөнү not pruinose, $ “ie un afra Hue 7. Thallus dark olive g rella Müll. АН 7. Thallus olive fuscous P. obs "eaae у. fue Ma Arg Thallus rier К yellow, medulla white, K—; ascospores (13—)16. 4 X 6-6 Ivory Coa isi СИ bá Abb. Thallus not is isidios 9 9. Medulla s some ade of deep red; nuls and reference qoem doubtful as coloréd médullae а and Рухт 10 9. Medulla ipe 11 10. Medulla pur rple- red; һи у. coccinea Мі 11. Arg. 10. Medulla blood-red; duda pale gray, white-pruinose; Kenya.......... Crocynia baematina Stein 10. Medulla red; Congo Crocynia Leopoldi Stein 11. Thallus гек ic sterile, and reference here doubtful 12 11. Thallus white or grayis 12. Laciniae convex, narrow, soralia circular, soredia isidioid; Ascension Island...............- Р Аа cium Arg. 12. Laciniae broader, similar to n varieties of P. picta Ре intense yellow ingly sorediose above; Ken P. picta v. rti. Müll. Arg. 13. Thallus ashy 14 13. Thallus pure white 16 14. Аз 12-15 X 5-7 и; thallus ashy to lead-color, fuscescent; Angola............ E pos " 4 ‚Р. africana Müll. Arg. 1 14. Ascospores 13-18 X 5-8 Ms thallus ashy-glaucous 14. пенен 15-23 X 7-9 и; disc ashy-pruinose, dark disc under the hypothecium uch thicker than the e wan Be part ni р, per ides hei the _thecium; thi llus similar to P. stellaris v. acrita; corticole; üll. Arg. 15. Thallus K yellow; Ilha Principe P. palmarum Vainio 15. Thallus к; А г mere Vainio Ae SE sor са 9—1 —6 м; h li acr [ 16. ‘hints ospores 15—20 x 6—8 ш; lobes wide; corticole; Kenya.............. i рожа Hue . — sers , pulverulent ре: Guinea.............. 16. Ascospores 18-22 X 6-8 u; valde crenu ate pul A т ho v] 16. Ascospores 15-23 X 7-9 и; dark disc Ld o thecium d much thinner than edia, K— ow; saxicole; Abyssinia ESA thecium; lobes with pulvinate sor y AL MEN чун bagl. Рнү$сїА Poncrnsn Hue, Mém. Soc. Bot. France 28:10. 1916. Type: Kenya, Tika River, southeast of Mt. Kenya at Blue Post, 45 km. from Nairobi on road to Fort Hall, 1500 m., Poncins. [Vor. 40, 398 ANNALS OF THE MISSOURI BOTANICAL GARDEN Thallus foliose, not closely adnate, more or less circular, up to 2.5 cm. in diameter, laciniae 1-2.5 mm. wide, radiating, white to olive buff, margins and surface of lobes soraliate, soralia up to 1 mm. in diameter, convex, and coarsely granular; upper cortex 20—40 д thick, pseudoparenchymatous from vertical thin- walled hyphae, cells mostly 6-8 p in diameter, upper portion heavily nubilated with grayish granules; algal layer 20—40 p thick, of protococcoid cells 5—8 p in diameter, continuous and closely packed; medulla 100—200 pu thick, of loosely woven, periclinal hyphae 3—4 и in diameter, heavily nubilated with minute brownish granules; lower cortex 20—40 p thick, of conglutinate thick-walled hyaline hyphae, the outer 6-8 и nubilated with dark brown granules; rhizinae 60-75 p in diameter, long, white, finally darkening. Apothecia elevated, cupulate, constricted at the base to substipitate, about 1.5 mm. in diam- eter, margin smooth, then vertically sulcate, inrolled, becoming completely isidio- sorediose, disc rufous at first, becoming black, not pruinose; amphithecium of young apothecia 185 д thick, of the same structure as the thallus but the algal layer 100 р thick on the sides of the thecium, only 60 p thick below with cortex thickened to 60 р; parathecium not clearly differentiated; hypothecium 15 р thick, of dense subvertical hyphae; thecium 105 м tall; paraphyses conglutinate, dicho- tomous above, tips capitate, terminal cell brownish, subspherical, 3—4 р in diameter, nubilated with brown crystals; asci clavate, thick-walled when young, 8-spored, 65 X 16 p; ascospores bilocular, constricted at the septum, brown, imbricately monostichous, 21-24 X 8-9.5 p. Spermogonia flask-shaped, immersed in the medulla and the lower half of the algal layer, wall hyaline, scarcely differentiated, venter about 50 y in diameter and 40 y tall, neck 20 р in diameter and 55 p tall; spermatiophores erect, septate; spermatia bacilliform, straight, about 4—5 X 1 p. I hesitated to refer my material to this species, as the thallus is not pruinose, the medullary hyphae are heavily nubilated, and the dimensions of the mature thallus are twice those given by Hue. The thalli are very fragile and have been removed from the substrate. The structure of the thallus and of the soralia agrees with that given by Hue. The apothecia in the Cameroons material are immature, as only young asci are found and soralia are just beginning to form on the margins, while the Nigerian material is quite mature, ascospores varying from brown to almost black. The dimensions of the microscopic E are identical. Cameroons: near Kumba, on Theobroma, C. A. Thorold 137. Nigeria: Majeriko near Ibadan, on Theobroma, C. A. usd 138. PHYSCIA ABBREVIATA Müll. Arg., Bot. Jahrb. [Engler] 20:260. 1894. Type: Usambara, corticole, Holst 1016 p.p. Thallus foliose, small, closely appressed, smooth, 120 и thick, margin very ir- regularly lobed, lobes rounded, 0.5—1 mm. wide, center subcrustose, deep olive; upper cortex 20—25 y thick, hyaline, pseudoparenchymatous; algal layer 55 p thick, of closely packed protococcoid cells 5-6 и in diameter; medulla 25—40 p thick, of compactly woven subvertical hyphae, with occasional algal cells nearly to the lower cortex; lower cortex 20-25 y thick, of coarse, conglutinate hyphae, the outer ones brownish; rhizinae short, about 15 и in diameter, once dichotomous 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 399 near the tip. Apothecia 0.4-0.7 mm. in diameter, sessile, not constricted at the base, margin entire to subcrenulate, concolorous with the thallus, disc opaque, fuscous black; amphithecium 55 д thick, of the same structure as the thallus; hypothecium 15 y thick, of deeply staining, subvertical hyphae; thecium 105 p tall; paraphyses slender, walls gelified, dichotomous above the аѕсі, the upper 3 cells spherical, brown, 3 и in diameter; asci 50 X 12 p, thick-walled and tip of the protoplast thickened when young, 8-spored, clavate; ascospores brown, bi- locular, not constricted at the septum, protoplasts ellipsoid, connected by an isthmus when immature, 16-18 X 9-11 y. Only a single thallus about 5 Х 10 mm., with four apothecia, was available for study, on a twig associated with Ionaspis ascidioides, Pyrenodesmia conglobata, and Pbyscia afra Hue. e apothecium sectioned is rather immature; the asco- spores still very pale brown, 13—14 X 6-8 p. Nyasaland: Kasungu Hill, 1100 m., on living bark, L. J. Brass 17458a. Pryscia AFRA Hue, Mém. Soc. Bot. France 28:11. 1916. Type: Kenya, Tika River southeast of Mt. Kenya at Blue Post, 45 km. from Nairobi, on road to Fort Hall, 1500 m., corticole, Poncins. Thallus white-opaque but not pruinose, K—, rounded, 15 mm. in diameter, laciniae 1—1.5 mm. wide near the center, narrowing to 0.3—0.6 mm. near the 2-3- fid tips, slightly convex, white below with white rhizinae; upper cortex 30—40 p thick, pseudoparenchymatous, of vertical hyphae 8—10 џи in diameter; algal layer 40 p thick, cells protococcoid, 6—10 p in diameter; medulla 50—80 џи thick, of compactly woven, very thick-walled hyphae 4 д in diameter; lower cortex 20 p thick, of hyaline, periclinal hyphae; rhizinae short, 60 и in diameter. Apothecia 1 mm. in diameter, constricted at the base, margin thick, prominent, entire, disc dark rufous to black, pruinose; amphithecial cortex 30 м thick above, 40-90 p thick below; parathecium hyaline, 12—15 и thick above to 40 y thick below de thecium, of periclinal hyphae above, intricate and forming the hypothecium below; paraphyses dichotomously or corymbosely branched above, tips capitate, dark rufous, 2—3 р in diameter; ascospores brown, bilocular, distichous, (15—) 18-20 (-22) p. I have had a single thallus with only one apothecium which was not sectioned. The macroscopic and the microscopic characters of the thallus agree with the above description which was translated from the original. Nyasaland: Kasungu Hill, 1100 m., on living bark, L. J. Brass 174582, Vernay Nyasaland Expedition. ANAPTYCHIA ANaAPTYCHIA Koerb. in Mass., Mem. Lichenogr. 33. 1853. Borrera Ach. Lichenogr. Univ. 93, 496. 1810, p.p. min. Heterodermia 'Trev., Atti Soc. Ital. Sci. Nat. Milano 11:613. 1868. Pseudopbyscia Müll. Arg., Bull. Herb. Boissier 2:append. 1:10. 1894. Type: not designated; A. ciliaris and A. leucomelaena treated, either of which [Vor. 40, 400 ANNALS OF THE MISSOURI BOTANICAL GARDEN would conserve the genus in its present usage. Воттета was based on fifteen species, three now included in Anaptychia, the rest now in the Usneaceae, Parmeliaceae and Teloschistaceae, and the name has been abandoned. Heterodermia Trev. was based on seven species, all now included in Anaptychia. Pseudophyscia Müll. Arg. was based on Lichen speciosus Wulf Thallus foliose to fruticose, but always dorsi-ventral, prostrate to more or less erect, laciniae relatively narrow, branched, margins often ciliate; upper cortex of conglutinate periclinal hyphae; algae protococcoid; medulla rather loosely woven of mostly periclinal hyphae; lower cortex of conglutinate periclinal hyphae, often absent except at the margins of the lobes. Apothecia lecanorine, covered by the amphithecium until nearly mature, margin usually lacerate or lobulate; amphi- thecium well developed; parathecium scarcely differentiated, disc fuscous, often pruinose, epithecium K—; paraphyses simple, usually septate; asci 8-spored; asco- spores bilocular, brown. Spermogonia immersed or nearly so; spermatiophores septate; spermatia lateral, short-cylindric, straight. Most species have unusually wide geographic ranges and no species have been described based on tropical African material. AwaPTYCHIA Adamesii Dodge, sp. nov. Type: Sierra Leone, Sefadu (Gbense), on trunk of Elaeis guineensis, P. Adames, comm. Е. C. Deighton M4752, growing over moribund Parmelia (Amphigymnia) lobulascens Stnr. v. isidiosissima Dodge. Thallus foliosus, dichotome vel subtrichotome ramosus, laciniae ca. 1 mm. latitudine, soraliis capitatis terminalibus, 150 џи crassitudine, olivaceo-alutaceus, inferne ecorticatus, superficie arachnoidea, flava; rhizinae nigrae, nitentes, 2 mm. longitudine, dense squarroso-ramosae, marginales; cortex superior 55—65 p crassi- tudine, gelifactus, hyphis periclinalibus dense contextus, strato extero 15 p crassi- tudine, crystallis minutis, flavo-brunneis nubilato; stratum algarum 15—25 p crassi- tudine, cellulis Trebouxiae, plus minusve angulosis pressione mutua, 6-7 и diametro; medulla 60-80 y crassitudine, hyphis pachydermeis, granulis griseis nubilatis, laxius intertextis et brunneis in strato infero, 15 u crassitudine. Apothecia immatura, stipi- tata, cupularia, margine inflexo dentato; cortex amphithecialis inferne 250 р crassi- tudine ad 35 р in margine inflexo tenuescens, gelifactus, hyphis superficie perpen- dicularibus, in dimidia parte extera granulis brunneis nubilatis, intus griseis; stratum algarum 105 y crassitudine circa thecium, ad 80 и in margine tenuescens; medulla 40 р crassitudine circum thecium; superficies thalli sub apothecio cortica- tus, cortex 135 u crassitudine, structurae corticis amphithecialis similis; hypo- thecium 80 p crassitudine, hyphis tenuibus subbrunneis, dense contextum; thecium са. 110 p altitudine; paraphyses tenues, dichotome ramosae, apicibus clavatis, 3 p diametro; asci immaturi clavati, pachydermei, 95 X 20 р; ascosporae non visae. Thallus foliose, dichotomously and subtrichotomously branched, laciniae about 1 mm. broad, mostly terminating in large capitate soralia 150 p thick, olive buff, ecorticate below, surface arachnoid, yellow becoming tawny in the older portions; rhizinae black, shining, about 2 mm. long, densely squarrose-branched, marginal, 1953] DODGE—LICHENS ОЕ TROPICAL AFRICA 401 close set, giving the appearance of laciniae on a black hypothallus; upper cortex 55—65 p thick, highly gelified, of interwoven, mostly periclinal hyphae, the outer 15 м nubilated with minute yellowish brown crystals; algal layer 15-25 и thick, more or less continuous, of closely packed colonies of Trebouxia, cells more or less angular from mutual pressure, 6-7 p in diameter; medulla 60-80 y thick, of inter- woven, very thick-walled hyphae about 5 м in diameter, nubilated with grayish granules, brownish in the lower 15 р, more loosely woven below. Apothecia im- mature, cupulate, stipitate, margin inflexed, dentate; amphithecial cortex 250 p thick below, thinning to 35 и thick over the thecium, highly gelified, hyphae perpendicular to the surface, forming a pseudoparenchyma, heavily nubilated with granules, brownish in the outer half, grayish within; algal layer 105 џ thick around the thecium, thinning to 80 u thick above it, outer surface of the layer dentate; medulla about 40 д thick around the thecium, not extending above it; under-side of the thallus corticate below the thecium, cortex 135 y thick, similar in structure to that of the amphithecium; hypothecium about 80 д thick, of densely woven, slender, slightly brownish hyphae; thecium about 110 p tall; paraphyses slender, ichotomous, tips capitate-clavate, 3 и in diameter; immature asci clavate, about 95 Х 20 p, thick-walled; ascospores not seen. FUNGI Mytilidium sp.—A few hysterothecia, shaped like а clam-shell, were found on twigs of Bauhinia tomentosa from Njala (Kori), Sierra Leone, Е. C. Deighton 4307, which do not seem referable to a previously described species, but the material is so scanty that I hesitate to describe it as new. The ascospores are brown, fusiform, 4-locular, slightly curved, about 32 X 10 p, uniseriate in the ascus. The hysterothecia are erumpent through a lichen thallus covering the bark, but I have found no connection between the fungus and the lichen thallus. Were such a connection evident it might be referred to Sclerographis, from which it differs greatly in habit. The paraphyses are slender, somewhat dichotomous above the asci, but not branched and anastomosing to form an epithecium as in the Ope- graphaceae, to which Sclerographis belongs. GENERAL INDEX ТО VOLUME XL New scientific names of gos and the final members of new combinations are printed in bold-face type; synonym italics; and all other matter in ordinary type. and page numbers having reference to figures and plates, in A Acanthographis, 311 Acantho Pucon 312 caspora Pene re A € Leprieurii, 335 Africa, нер, Some lichens of, 271 Aglaopisma, 9 Agriculture, hes system of, in Yucatan, 51 A 379 303 Almanac, Bluegrass pasture, 1 Alyxoria Vr ачен 389; debanense, 390 Amp. йота, 99; Adamesii, 400; ciliaris, Anderson, Edgar: The analysis of suspected hybrids as illustrated by Berberis X glad- ан 73; and William т Brown: popcorns of Turkey, 33 ымын, Henry N., and Sergius Н. Mamay: tween bluegrass pastures and, 1, 24, 26, Anomorpha, 312; turbulenta, 312 Anthracothecium, 273, 285; cinerosum, 285, 286; Doleschali, 283; sect. Euanthra- cothecium, 285; euthelium, 285; guineense, 285, 286; lugescens, 285; sect. Porinas- trum, 285; punctuliforme, 285; vitel- linum, 285 Antrocarpum, 341 nzia, 372 Archaeological: and ethnological races of studies of the Maya civ- New Менен А contribution to the lichen flora of, 6 Arthonia, 301, 303; ыны ч calo- spora, 305; ا‎ 304; cinnabar- ina, 304, у. dendritica, 304, v. ases ER 04, v. orbicella, imata, 304, v. rythrocarpa, 304, v. somali, 304; fuscescens, 308; gregaria у. ritica, 304; ilicinodes, 304; i ы 305, 306; leptogram- modes, 305, 306; leptographoidea, 304, tusariella, iv somaliensis, 304; sub- caesia, 304, Arthoniaceae, 01, 303, 338 Arthoniopsis, set © Artbopyrenia = нё 292 Arthopyreniella, Arthotheliv m, 301, 302, 205 $26; dicty- ophorum, 302; Bos Ascidium, 342; be hats а rhodo- Asiatic 1 races 6 maize, cob morphology of, 7 des tion to ethnology and archae- olo vm Asterisca, P p ii 335 Astrotheliaceae Astrotheliu Aulacographa, 312 Aulaxina, 307 В Васїйа, 352, 357; abessinica, 358; amylo- 358; araneosa, 358; с 358; і ade 357, 358; exiguella, 3 58; a, 358; а оз рее а 358; leucotrypta, 358; nigeriensis, 358, 359; nigrocincta, 358; subg. Scolicio- sporum, 274; Scottii, 358; socotrana, 358; stellatis. 358; sublecanorina, 358; M 358; trichosperma, 358; peir 358; sect. Weitenwebera, 357, В hybridization of, 75; relationships two species and their putative hybrids, 74, 76, 77 Bathelium, aire 278, 280, 286, 288, 296; positum, 289; d m f. simplicius, n Т, masto ideum, 289; papillosum, Ar 289, 290; porosum, 290; subalbens, 290 але 296 Belonia, 273 403 404 Berberis X gladwynensis, An analysis of 73 24, 77,78; : list “ eating p: fruit, 218; Wildwood, 2 Bent $66, 587; albidofusca, 388; albido- ens, 388; Жейсон v. mic ra, 7; cormorensis, 388; lactescens, 388; laetebunda, retaniae, 387; 8 maurula, 387; pertenuescens, 387; polio- terodes, 388; sinapisperma, 387; sordida, 388 Blasteniaceae, 384 Blastodesmia, 275 Blenotballia, 551 Bluegrass: pasture almanac, 1; pastures, 24- Bombyliospora, 584, 385; cerinella, x3 vag ominguensis v. intermedia, 385; eee te Meyeri, 385; фен, 85; pruinata, 385; thomensis, 385, 386; Thor 385, 386; togoensis, 385 Borrera, 59 9 Boris 286, 290; sect. Anthracothecium, ass, L. J., lichens collected by, 271 reise, 342 кё rnt ont 361; argentea, 361; Mariae, 561; tricolor, 361; tristis, 361 Brookhill "Фано: observations on bluegrass pastures made at, m L., Edgar Anderson and: orns 3 Turke А Bryopbagus, 348; Gloeocapsa, 34 ВиеШа, 355, 390; Adamesii, 392; afra, 391; ам 392; бе: Diploicia, 390; sect. Diplotomma, 391; disciformis orta, 391, v. oblongata, y 392, v. pachyspora, 391, v. rhodina, 3 391; sect. Менаві 391; veg 391; pachyspora, 391; parasema, 390, v subaeruginosa, sak paupercula, 392: йы. регепѕ, 392 ginea, 391; stellulata f. robali 392; balbula, 392; subdives v. mozambica, immersa, 391; io KE rata 391; E Rer obfuscata, 391; ТоЫей 391; tonini- шы. 392 [Vor. 40, ANNALS OF THE MISSOURI BOTANICAL GARDEN Buelliaceae, 390 Bunodea, 280 e Callopisma, 388 Caloplaca, 384, 388; arizonica, 67, 08; asma- rensis, 3 88; вонь 3 89; Balfouri, 39 Ta * exasperata, flavorubens, 390; granulifera, 390; gein liformis, 390; inconcinna, 388; murorum, 389, 390, v. granulifoemis; 390; ochrace- =» 390; Odoardi, 388; zarabetieé; Сне, eg chen from, 271 Candelaria, Candelariella, s> Carlosia, 30 Catillaria, “eo subternella, 358 Catinaria, 338, 339, 393 Chapsa, 342; набй, 343 ісогу іп pastures, 13, 2 Chiodecton, 301, 333, 337; album, 338; antyloplacoides, 337; amylóplacum, 337; Brunnthaleri, 338; subg. — 301, 337, 338; circumscissum, sub Enterographa; 337; subg. E rt intercede ens, 338; irregulare, 337; 3 minutulum, 337; plis bicum, 337; nanum, 337; palmensis, 3 rotundatum, 337; sanguineum, 337, seriale, 337; socotranum, 337; oiii 337 Sora REE. 333 Chiog Chlore phela, 363 Chromosome knobs on Turkish popcorns, grong d positions of, 44 Chromosomes: of Delphinium, 132, 134, 164, morphology of, 136, 147; of Dio- spyros, 213; of Petunia, treated with 1 : Clathroporina, 273, Coal Pura foci of кел found i in, 185 among certain archaeo- baud à and сока races of maize, 29 Coccotrema, 273: 287 Coe — 350; Deigh- toni, 349; ium 1953] INDEX Collema, 351; aggregatum, 351; ascaridio- Blenothallis, 351; = — w ә fasciculare, 351; flaccidum, iha 3517 ре 351; nigrescens, 51, var. minutum, 351; rupestre, 551; scotinum, 351; OH. 351 Collemaceae, 351 Coniocbila, 541 Contribution to = 2 flora of Arizona exic n, Coscinedia, 342 Creographa, de Perte 327 ems t 7; haematina, 397; Leo- Ua: мт Cryptothecia, 30 302; caesioalba, 302; geniflexum See ‘nigeriensis, 302; Stirtoni, 302; Баа lans, 302; Thoroldi, 302, 303 yphelium, 300 Cyphella, 36 pir acri data of plants see ie effect of x-rays, 261; methods used in study of Delphini hiex, 134; study of ais, ах morphology and systematic rela- tionships of Delphinium Х Belladonna ort. ex Bergm., 113 D Dactylina, 579 Darrow, Robert A., Lichen collection of, 63 cement relationship in speci ТЭР; 1460, 1 hromosomes of species, 132, 164, sr hybrids десе ней Ьу Рго- rived hexaploids, 150; history of, 113, 132; hybrids of the Bellsdonna type, 113, 121, 172, 174, leaf types in, 113, pollen studies of, 124, 130; morphology of nat- ural ne 113; satellite expression in hybrids, 138; seed germin ability am 155; taxonomy of ا‎ cies, Delphinium X Belladonna Hort. ex Bergm., The cytology, арни gy, and system- atic relationships of, Delphinium, 113; ajacis, on 117 132; Barlowii, 120, 130; Belladonna, 113, 405 115, 119, 125; Belladonna type, hybrids of, 117, 118, 146; X Belladonna 126; Bellamosum, 140; “В 121, 124, 130; "Capri", 119; cardinale, 117; cardiopetalum, 132; chamissonis, 116; cheilanthum, 113, sum, 113, var. ; ren me elatum, eit 121, 124; subg. Eudelphinium, 115; 115, 117, 118, 130, 168; “Grandiflora,” 117; a 114,115, 117, 119, 123, 126, 130, var. chinense, 118, 121, 133, 136, 170, “Blue Butter fly”, 121, 122, 137; gypsophilum, 133; Hanseni, 133; Hendersoni, 115; 120; 125, 166, “Lamartine”, 116, 124, 125, 130; sect. Kolobopetala, 115; Middendorffi, 116, var. parviflora, 116; oerheimi”, 119, 124, 125, b TA nudicaule, 117, чү: vereerqum, mosum, 115; Киузи, 133; snb" ава, id. 124, 126, 140, 168; Staphisagria, 114, 132; sect. Staphis- agria, 114; “Summer Skies”, 124, 176, X grandiflorum, 123; tatsienense, 115, ; "Tom Thumb", 124; varie- Desmaziera, 379 CIEN HIE 312; turbulenta, 312 Dimerella, Diorygma, 325; tinctorum, ax Parone 211; chromoso: of, 213; his- ‚ 211; discolor, 214; kaki, 214; bo (215 texana, 214; virginiana, 211 214 Diospyros срива І., The distribution of, 211; clonal types, 218, 224, 225; dispers a by animals and birds, 217, 9 =, oo 220; in mix 225; local distribution, 214, 2I de facto affecting, 212, 214; range of, 211, Diplogramma, үш yen Diplograpbis, 312 Diploschistes, 312 406 Di f ES p Dione virginiana L., 211 Dithelops Bolen sean | етай Some lichens of opical Afric оен, 312; а 314 Е Ecology: Bluegrass pasture almanac, 1; Dis- tribution of Diospyros virginiana L., 211; Wildwood, a study in historical, 227 Ectograpba, 526 Ectolechia, 342, 343 Elmo ore, Mr. and Mrs. Francis, lichen col- preliminary survey of 333 pra i дыв races of maize, variation in cob ology among certain, Etter, Alfr ed G.: Bluegrass pasture almanac, 1; veces udy in histosital colog кийре, Es Microglaena, 275 Ever ани 379 Farriola, 300 Fissurina, 312; Dumastii, 312; incrustans, Florissant, Mo., history of an estate near, Flower abnormalities of plants treated with x-rays, Forests in the St. Louis area, 1817-18, 230 beu of Medullosa, 183; of persimmon, Кон. p Fulgensia Fungi of ded Africa, 401 G Gage, Marilyn Amy: The cytology, morph- ology and systematic relationships of Del- — Х Belladonna Hort. ex Bergm., Gasparini, хм, 389; murorum, 389 Glaucinaria, 323 ст Glyphis, Ж, 334; cicatricosa, 334, v pulvinata, 334; labyrlathice, 334, 335; tricosa, 334 Gophers їп bluegrass pastures, 3, 30 Graphidaceae, 311, 333 [Vor. 40, ANNALS OF THE MISSOURI BOTANICAL GARDEN Grapbidula, 318 Graphina, 311, 312, 318, 323; abstracta, 324; aethiopica, 324; DB SUA 324; arthothelioides, 324, 325; нра «07у 324; carneum, 323; chlorocarpa, 312; ra, 324; incrustans, 312; javanica, 318; Pelletieri v. macrior, 324; grammina, 326; sect Platyé 311, 325; sect. Platygraphina, 326; pli- Poitiaei, 327; Неа реле diella, 324, 328; straminea, 324; sub- лен 324; sidan 323; ulcerata, 3 25; varian а 308, 311; Аве, 312, 314; albo- notata, 312; aperien ruinosa, 313; stern, 314; betuligna, 512; ым: E hinat, 303, leuca, 323; hyalinella, 313; increbrior, infida, 313; inusta, 323; Junghuhnii - : ОТ; > 1 327; pulverulenta, 312; raddacensis, 323; roseotincta, 312; و ;312 ا‎ Hs 318,327; г арта 313; su colaeo- spora sect. Glyphis, 334; Ава, 311, 312; 1 var. cus- ans, ; superans, 313; tenella, 314; Thoroldi, 313, 336; timidula, 312; аса 313; turbulenta, 312; ulcerata, 3 Grazing habits of RE effect of, on grasses, sect. Я 346; blurilocularis, 348; 348 Secoliga, Gymnographa, Gyrostomum, 340, 346 H Haematomma, 331, 365, 370; Brassii, 370; dactylopholis, 370, 371; similis, 370; sub- Hagenia picta v. rupicola, 397 1953] INDEX Harlan, Dr. Jack, ge of Turkish by, 3 Helminthocarpon, 307; нато ay Hemitbecium, 323; chrysenter ron, 3 orman, hybrid De i in Heterangium, 198 Heterodermia, 399 Heterotbecium, 361; Berteroanum, 361 Holotbelis, 273 Horse grazing, effect of, on bluegrass pas- 7 4, 87 Hybridization in the evolution of natural populations, Hybrids: An analysis of rg as fo trated by Berberis X gladwyne ‚ 73; of Delphinium Belladonna creed ex Bergm., 115; o Potenti; treated with x-rays, 25 Hy menodecton Fr pedi tat ien неа, 337 Hysterina, 30 Icmadophila, 365 Imbricaria, 372 Ionaspis, 346, 347; ascidioides, 347 Irradiation, effect of, on Petunia, 259 Kansas: coal- eo fossils found in, 184, 189 Kuttlingeria, L Larkspurs, Garland, see Delphinium um, 35 6, 296; astroidella, 05 28 296; elegans, 296; madreporiformis, 296; marginat 6; nigeriensis, 296, 297; ochrotidcoder: 296; pauperrima, 296; velata, Leaf abnormalities of plants treated with x-rays, 260, morphology of, 265, 268, 270 Lecanactidaceae e, 338 , 331, 338, 339; amylacea, 339; ni te use deducens, , 365; sect. e 385 ec 346 Lecanora, 355, 365; aequinoctialis, 367, 368; Aspicilia, 348; — endo carneo- lutea, 348; carneosulphure £- lobata, 389; corallina, 367$ s ies ee 407 er. ecoronata, 367; elatina, 370; elmorei 5,66; s ubg. Eulecanorea, 366; fibrosa, 367 ; кш аыл 367; flavidocarnea, 366; flavochracea, 367; granifera, н granu- lescens, 367; beemaiomina, 37 ; homalo- placa,367; ltericola, 3675 seman’ lateritica, 366; lepros у melaleuca, 367; triguttulata, 367; tropica, 367; PIE REE is, Lecanoraceae, 365 Lecidea, 352, 353; albocincta, 354; aman- iensis, 354; angolensis, 354, v. orienta cili. 354; ‘argillicola, 354; aut Biatora, 353, 09 P m M. .6 = © — & 8 4 سا‎ м > ©з nd су ч © zi RA v a ON "O = уро- , 354 рене $52. Leigbtonia, 288; in y M ; pruinosum, 318; puncti- 8; sericeum Leireun, n 18, 327; Lyell, 318; tar- атеит, ее ТЫ еа, 351; turgidum, 351, 353; vesiculosum, 351 Leptorhapis, 273 Leptotrema, 340 Letbagrium, 551; criteri 351; con- lomeratum, 351; sciculare, 551; rupestre, 551; oiv. 351 Letharia, 3 Leucogramma, carneum, 32 6; confertum, 65 кы. 323; "us cense, 323; serpentarium, 325; turgidum, 323 408 ma crispus, 351; floridus, 379; haemo- tomma, 370; lepadinus, 341; гүн, 339; pallescens, 369; puniceus, 370; ru мн rugosus, 308; лазе. 372; nme. ‚ 312; speciosus, 400; ventosus, 370 Arizona an ew Mexico, n 6 Lichens of tropical Africa, Some, Limboria, 312; constellata, 312; нета 312 glaucophaeoides, 361; lecanorinum, 361; ewtonianum, 361; nigro obrunneum, 561, 363; pezizoideum, 361; sepiaceum, 361 Loxospora, 370 M McQuade, Henry A.: Some x-ray effects in Petunia, 259 Maize: Archaeological remains of, in North America, 84, 86, in South America, 87; cob morphology of, 79; milpa system of culture as practiced by ‘the M of Northern Yucatan, 31; Popcorns of Turkey, 33; present-day races of, in Asia, 87, in N merica, 83, in South Ame erica, 87; significance i r ology among certain archaeological ind ethnological races of, 79 Malvern B. C oes Experimental Farm at Clarksville, Mo., атау, Sergius H., ud Henry N. Andrews: Some American Medullosas, 183 Manure, animal, in pastures, 2 Maya Indians, in northern part of ч» Маше culture as practiced Ьу, 5 Mazozia, 323 мони е anglica, 184, 197, уаг. var. thiesseni, 186; subg. REA а E distelica, 184, 186, 187, O3, 205, 209; elongata 6; ds centrica, 186, 188, 197; ' grandis 191, 1 0 ; эл 200, 205, 207, heterostelica, 183, 184, 187, 188, 190, 191, 196, 205, 206; киска, 198; noei, 191, 196, 200; olg. 198; primaeva, 186, 188; solmsi, 198; thompsonii, 184, 185, 188, 196, 202, 203, 20 Medullosas, Some American, 183 Мои, мт, E bysterina, 327 Meissneria, 296; varia, 296 Melampidium, 338 ANNALS OF THE MISSOURI BOTANICAL GARDEN 97 361; Deightoni, 361, 362; - [Vor. 40, Melanodecton, 337; indicum, 3 Melanotheca, 281, 286, 290, ds a hari iana, Milpa system of maize culture, as practiced y the Maya Indians of northern Yuca- tan, 53 Minksia, 533 Monoblastia, 273, 287 y: of anus x Belladonna variation in, among certain archaeological and et —— races of maize, 79 Mycoblastus, 352 Мусаны, э 27 Myriostigma, 302; ada — 342; album, ues woes му, T yxodictyon MENS. $n PS ordeo 302 New Mexico, Arizona ane, A contribution to the lichen чый of, Nickerson, Norton H.: ih riation in cob morphology баў сеггаїп archaeological and ethnological ма s maize, Nigeria, lichens from, ог merican races sf maize, 105; cob phology of, Nyasaland, lichens from, 271 Ocellis Осана, 340, 342; albescens, 343, 346; 345; subterebrata, 343; PRE 345; trypanea, 343, 344 oe vig 369; palmicola, 369 Лема collected by, 271 ont fee 308; alboatra, 309; cinerea, 309; astroides, 339; aterula, 309; cretacea, 309, 327; cylindrica, 308; deli- cata, 309; depressum, 327; diophora, 308; 1953] INDEX 409 Dracaenarum, 309; elegans, 309; sect. тооны, 5 Euopegrapha, 308; pene 308; ES pe armentari Е A cip 286 309, humilis, 309; Hi poeta 339; leidella, 309; leptographa, 309; fic hencides: 308; ا‎ 309; Lyellii, 318; medalini, Poitieei, 327; prosedoides, 309; Seer eii ата, 303; rufa, 309; s ай. 309; да 327; а 309; sireblocarbe, 347; e n a 309; subgraphidiza, 310; subnothella, 309; are, 2 ; Vainioi, 308; vermelhana, 309; vestit viridulat, 309 O эзе re 307, 333, Opbioparma, Oropogon, 37 8, Oxystoma, 308; к, 308 Р Pachyphiale, 346 Paleobotany, 183 375; eurycarpa, 375; anningtoniana, 375; Hüdenbréodtis, 375, v. ciliata, 375; isidiosissima, 374, 375, 377; Meist 75; Nenybarti, 375; ibodeits; 375; neitobieitil, 375; nilgherrensis у. sub- ciliaris, 377; nitens, 375, f. isidiosa, 374; oliv v. esorediata, 375, v. sorediosa, 375; orchidophila, 373, 374; pedotke ysc usambar ‚373; Doings | v. bryo 373; bcn T tunia, Some x-ray effects in, 259; cyto- logical data, 261; morphological abnor- cip 260, 268, 260, 270; pollen co Visus st: 312, 326; alata, 328, 332; + lfourii, 327; iilos 527; сені: ruino i Deightoni е fuscescens, 328; innata, 327, ; leptotremoides, 328, 331; leucophora, 327, 32 ontagnei, 7; 328; pezizoidea, 327; quassiaecola, 327; scalpturata, 318, 327; — ri 329; streblocarpa, 327; ultum, 3 eere 311, 312, 318; confluens, v Deightoni, 319,320; Баз ад riticella, 319; duolicans, iu flammula, 318; inusta, 323, v. radians, 319; leuco- E то 319; lignatilis, 319; Lvl 318; lynceodes, 284, 319, 322; medusulina, 318; micrograpta, 319; micrograptoides, 319; navicularis, 319; ochra racea, 319, 320; diosca. 319; paragrapta, 319; Phyllo- ul 319; diste, 3 19, 321; subdeve- lans, 319; subnivescens, 3 19; tecta, 319, 322; tigrinella, 319; tides. 512 40 3 Phragmopyxine, P Phyllophthalmara, 34 Phyllopsora, 364; E 364; Buett- neri, 364; — osa, 364; parvifolia, 364 Phyllopsoraceae, Physcia, 396; eas. 397, 398; a 397; afra, 397, 398; african a, 397; а 397, v. Bay 397; palmarum, 397; picta у. coccinea, 397, v. RE 397; Pon- insii E Hi 397; singularis, 410 Pbyscia, 372 у Pioneer Hi-Bred Corn Co., Turkish pop- corns grown at, 33 Platygramma,318; ray riticum, 318; Lyellii, 18; suffultum Plo 326, 327; мы 327 Pollen of Petunia plants, effect of x-rays on, 261 > Polyblastia, 274 Рае 272; 273, 274, 275, 280, 275; haematochroa, 275; нь Nen 276; pyriformis, 275, 276; sphaerica, 275, 277; tropica, 275 0 3 6, 48; — Turkish, 3 chromosome knobs 44; фа ка 40; list of types, 42; variations in, +, 38; on e race, 37, 39; Asiatic € 35, Porina, 273, 278 Precipitation, effect of, on distribution of Diospyros virginiana, 213 Preliminary s survey of the Milpa system of maize culture as ана by the y Indians of the northern part of the Yuca n Peninsula, 51 Protoblastenia, 384 goana, 2 duplex f. simplicior, ; «Шок, 280; за 280; sect. Hemithecium, 280; sect. 5 Homalothecium, 279; sect. Holotheciu Psoromidium, 364; Wellingtonii, 364 Psoro theciam: sect. Bombyliospora, 385 Ptychographa, 307, 333 Pycnographa, 333 Pyrenastrum, 297; erumpens, 298; para- thelioides, 298, Genk 297; pruinosum, 298; septicolare, 297 [Vor. 40, ANNALS OF THE MISSOURI BOTANICAL GARDEN Pyrenodesmia, 385, 388; conglobata, 388, 389 Pyrenopsis baemaleia, 251 Pyrenula, 273, 278, 280, 342; Acaciae, 281; Alni, 281; ананна. 279; аѕріѕгеа, 282; ceratina, 279; discolor, 279; ducoléreliá, 282; Gravenreuthii, 281, 292; heirs 282, 284, v. denigrata, 284, v. uscula, 282; Limae, 282; а оао 282, 284; орала, 281; subg. Melanotbeca, 292; mozambica, 281; neglecta, 279; adds: 281; obscurascens, 295; ан. 281; ocellata, 281; oculifera, 281; parva, ; subaperta, 281; trombetana, 281, 282; try pened; 344 Pyrenulaceae, 272, 333 Pyrgillus, 300 — 318; rere 318; javanica, medusulina, 8 Pyrrbogra pha, 318; ‘flammula, 318; javanica, medusulina, ы, "ex Adame зэ» s лерни 395; chryvantha, 3 Сосо 96; Роси сз 395; endoleuca, 395; Mei v. sorediosa, 395; reirugll f: SM 395; phurans, 395 R Racodium, 349 Rediation ا‎ on Petunia: cytological, 261; on flowers, 260, 270; on leaves, 260, 268, 270 Ramalina, 378, 379 Ramonia, 3 hizocarpon, 352 Riddlea, 286, ыл; рарШоза, 287 Rinodina, 3 90; darrovii, 69, 70; nim- phaeocarpa, 71 Rudolph, иене a to ew the e flora of. Arizona and N ехісо 5 Sagiolechia, 3 St. Louis area: animal population at Wild- species in, 232; Wildwood, 227 1953] INDEX Sarcographa, 333, 335, 393; Cascarillae, 335; rum, 335; sec тек а, 335; labyrinthica, t ri ; Thoroldi, 335, 336; tigrina, 335 Scaphis, 308; anfractuosa, 308; compressa, 308 Schismatomma, 338 Schistophoron, 300 Schistostoma, 341; о» 341 Sclerophyton, 3 eco igo дч jas; Deightoni, 348; pluri- locul Nani sect Melnotbece 292 emigyalecta, rcr ed A from, 271 д г, m of ospyros ea L3 Solinogriphe, Solenospora, b4 orina, 30 South American races of maize, 35, 86 Sphaerophoropeis, poropodiu Stesobolas, 342; Berkeleyanus, 343 198 Stock, grazing pom of, 2 Sutcli ia, Sw effec of excretory behavior on blue- Sy ion ий, 273; Bayrhoefferi, 273 abis ier seis 337; vri 337 echoblastu abl r a of е x Belladonna Hort. ex Bergm., T Temperature, effects 5 on distribution of Diospyros no 13 T balloloma, 3 Thamnolia, ite T becaria, 326; quassiaecola, 327 Thecograp a, 326; ceramia, 32 at вех fulva, 275; sect. meristosporum, э. 273; sect. T belopsis, 273 Thelococcus, 27 Thelographis, 326 Thelopsis, 273; flaveola, 273; inordinata, 273; rubella, д: ве1еповрога, 273,274; subporinella, T heloschisma, b Eschweileri, 318 411 аала: 510, 341; cameroonensis, he ; cavatum, 345; forat jn 4 СТАЯ 342; Pechuelii, 341; porino- ides, 343; schistcitumoldet. 341; subtere- brata, 343; terebrata, 344; sect. Tremo lopsis, 342; AUTE m, 341; SI. 42; M a Thelotremac c Theobr og of tropical Africa, lichens grow- ing on, Thorold, t jd lichen collection of, 271 Tomasellia Toninia, 35 Trees at Wildw n St. Louis Co.: early descriptions, 230; in 1946, 232 , 254; ages of, 234, di aie heights СИ к th rates, 234, compare witness trees, 237, density of, 239; каллы жїр 245 Tremotylium, 340; africanum, 340; ango- lense, Trichophyma, 303 TD em. 385 Trypetheliaceae, 286, 333 Trypethelium, 286, 290; aciculare, 290, 291; зу can 290, 291, 292; ponas É pusillum, 296; pustulatum, 290; sphaero- cephalum, es Sprengelii v. porosa, 290; subalbens, Turkey, а of, 3 pcorns, 33; diuidit knobs, n 33; variations in, : 42, Aegean race, 27, Asiatic race, 36, Tylophorella, 300 Tylophoron, 300; a 300; mod- modestius, 3 U Uganda, lichens from, 2 rine, animal, effect Tf on pasture grasses, 2, 9, 24, 26, 28, 30 Usnea, 379; equi 382; acanthera, 582; africana, > nt бы 412 Baileyi, 380; bicolorata, 382; Вогпе- muelleri, 382; cartilaginea, 381; chloreo- ides, 383; complanata, 380; contorta, 385; corrugata, 380; St ane 380; de- 82; leprosa, 380, 383; Liechtensteini, 38 0; livida, 383; Lyngei, 383; M 38 cyphellata, 380; pulchella, 381; pulveru- lenta, 380; rugosa, 380; tüvidesceri, 381; rüwetiorisid. 381; хаба: 381; scutata, 382; шр сиин, 381; speciosa, 380; 385; umbrata, 385; undulata, 81; usambarensis, 381; Vainioana, 381; r 454 vesiculata, 380; Wel- 38 7 Ustalia, 303; anguina, 323 У Variation: in cob morphology among сег- tain ЖИ апа KEKÎ races [Vor. 40, ANNALS OF THE MISSOURI BOTANICAL GARDEN of maize, 79; in Turkish popcorns, dia- grams нисан ‚ 38 iege aspistea, 318; mamillana, 284, ; guineensis, 285; parella, 369; porosa, a. t M ea, 344; subcaerulescens, 275 Volveris; 34 У Weeds: control of, in maize culture, 58; in bluegrass pastures, 13, 16, 2 1 Wildwood—A study in historical ecology, ages of, 235, sizes, 234; vegetation be- neath trees, 246 X-ray: effec Petun Some, methods a бер, 249: коя wi 0 Xylographa, 307 Xyloschistes, 312 Yucatan Peninsula, Maize culture as prac- ticed by the Maya Indians in the north- ern part of the, 51 Zea Mays, see Maiz Zwackbia, 308; ен 308 MISSOURI BOTANICAL GARDEN STAFF Emeritus Director JOHN S. LEHMANN Assistant Director EDGAR ANDERSON xecutive Assistant Leicester B. Faus SARON W. DODGE, ROBERT J. GILLESPIE, ogist Research анны ек E My euer JP GEORGE B. VAN ScHAACK, Curator е Her Honorary Curator of Grasses HENRY N. ‘Axons JULIAN А. STEYERMARK, Paleobotani Honorary Research Associate Rorra M. Tits N, ; Joun D. Dwyer, A on Eee of the Research бос erbari Ноєн C. аа. : Nett С. Horner, Curator Museum of Librarian and Editor Economic Plants : of Publications GERALD Urgicr Business Manager BOARD OF TRUSTEES President JOHN S. LEHMANN и DANIEL Second Viet Petru: EUGENE DUDLEY FRENCH GEORGE T. Moore Henry HITCHCOCK A. WESSEL SHAPLEIGH RICHARD J. Lock woopD ETHAN А. Н. SHEPLEY ROBERT BROOKINGS SMITH EX-OFFICIO MEMBERS ARTHUR Н. COMPTON, — STRATFORD Г. MORTON, Chancellor of Washington ; ; President of the Academy University - р Science of St. Louis RAYMOND В. "TUCKER CKER, : ARTHUR C. LICHTENBERGER, MNT Ta ge ; PE е RA ЗЕ © ERRHARD President бе шга of neat of St Los