-Vol1 ‘JANUARY, 1943. = No.1 - FARLOWIA. A JOURNAL OF Cryprogamic BoTany Eprrortat Boarp a5 DAVID H. LINDER, Editor E. V. SEELER, Ja, Mg. Ed. . R. Bartrotomew - -G. H. Ripa __ F. Verpoorn - 'W. H. Weston, Jr. _R. M. Wuetpen : PUBLISHED BY : W LIBRARY AND HERBARIUM OF HARVARD UNIVERSITY / 90 Drvintry Avz., Campringz, Mass, FARLOWIA Published semi-annually. Four numbers = one volume of eppromimarcly five hundred pages. Subscription price $5.00 per volume. Single numbers $1.50 each. Subscriptions and remittances should be addressed to FARLOWIA, 20 Divinity Ave., Cambridge, Mass., U.S. A. . Numbers lost in the mails will be replaced if reported within a reasonable period. CONTENTS OF VOL. I, NO. 1 PORMWORO. OVS) GUIE dt LUO ues os orca vn ck ook bs ae ee 1 Tue Arms or Furure ReskarcH in THE Atcan. By V. J, Chapman........ 5 Notss on New ENGLAND Augan, III. By Roy M. Whelden................ 9 Apive Mossus rrom New Guinma. By H. N. Dicon........ Pee A RO Mossss or Papua, New Guinzga. By Edwin B. Bartram. Sara SS Seay meget 41 Tue Genera KickxEa, MaRrenseia, AND ComMANsIA (Phycomycetes). EY PAU Pes SONOS obs hae y GV cate he bane Nae aE) CP aE OES 49 Basipium ForRMATION anp Sporn DiscHareH IN GrunosroraNerum Nipus- Avis. By Alton Bi Prince. 7 2 5a er aus. See onee 79 Tan Genus PeciicunaRta (Thelephoraceae). By Donald P. Rogers....., oa 06 Soman Funeicotovs Fouwnat. By Rdgar ¥: Beeler, IR; «sep See sek ove wok 119 STupius IN THE Gmnvs cua TIT ( Discomycetes). By W. Datprence White. 135 FARLOWIA _ A JOURNAL OF CRYPTOGAMIC BOTANY VOLUME 1 — 1943-1944 | ee ar PUBLISHED BY FARLOW LIBRARY AND HERBARIUM OF HARVARD UNIVERSITY : 20 Divinity AvVE., CAMBRIDGE, Mass. 4 te i Ny ey Me FARLOWIA A JOURNAL OF CRYPTOGAMIC BOTANY Vou. 1 JANUARY, 1943 oT NOt) FOREWORD The question may be raised: Why have a journal for taxonomic cryptogamic botany? To those who query the desirability of inaugurat- ing a periodical of this description, the following brief answer is offered. Those who have been interested in cryptogamic botany in its taxonomic aspects have long recognized the need of monographic studies, either taxonomic, morphological, cytological, or ecological, which might con- tribute to the systematic arrangement of the many and diverse groups of organisms which constitute the non-vascular cryptogams. Likewise it is evident that taxonomy is the keystone of all biological studies since it provides the investigator in other fields of endeavor a label for the specific unit of plant life that is being studied, thus making it possible for future students to confirm or to amplify the previous investigations. The stability of the name employed to label the organism depends to a great extent on the thoroughness with which the special group has been studied, how carefully the older and often imperfect records have been evaluated, how widely acquainted with related groups is the writer, and how well he has incorporated the information that may have been pro- vided by morphology, cytology, genetics, ecology, chemistry and other related fields. Unfortunately, our knowledge is still meager, but on the basis of that which we do possess, it may be stated that accurate tax- onomic investigations not only borrow from but contribute to related branches of knowledge. At the same time, it may be said that this discipline is a prime necessity for the validation of results, either from a scientific point of view or, since all knowledge may be of eventual prac- tical value, from the viewpoint of its contribution to human welfare. Frequently the systematist, through his special knowledge of the organ- isms that come to his attention, is in a position to save a considerable amount of time for the chemist, the physician, the physiologist, or the bio-physicist since he is able as a result of his taxonomic studies to suggest a single species or a group of species that will satisfy the needs of the investigator in the other fields of research. This contribution is almost wholly forgotten because it is less dramatic than are the results not infrequently secured by those, who without any attempt to belittle them may be called test-tube biologists, who started their studies where generations of taxonomists left off. Many examples could be 1 2 FARLOWIA, VOL. 1, 1943 chosen from the broad field of cryptogamic botany which would demon- strate clearly the dependence of other branches of biological science on the results of the many years of labor of the collectors and the taxonomists and which would illustrate the value of the so-called useless information. Probably no better example could be chosen than the situation that has arisen as a result of the cutting off of America’s supply of agar-agar by the Japanese who with their cheap labor and favorable localities have a monopoly on the harvest and production of the algal material. With the cessation of imports, commercial concerns have turned to the algae of the American coasts for substitutes for the agar that they used in the preparation of hand-lotions, ice cream, and other products. Commercial needs are more easily supplied than are those of the bacteriologists, the public health officers, and the plant and animal doctors who require a substance of rather specific physical characteristics for the hardening of the culture media that they employ for growing the pathogenic organ- isms which they are studying in an effort to control diseases. It is im- mediately apparent that agar plays a very important role in the welfare and nutrition of this and allied countries. The acute shortage of this material has made it necessary that substitutes be found at the earliest possible opportunity. Fortunately, as a result of the activities of the collectors, many of whom were amateurs who made the gathering of representative algae a hobby, and of the taxonomists whose studies have made known the morphology and structure of the algae, there is present and available for use, either in the literature or in the various herbaria, a large amount of information that is of immediate value. Those who are today seeking a substitute for the Japanese agar have been spared the large amount of pioneering work that has been done for them, and can turn their attention without loss of time to the analysis of the more likely species. The above may be considered by some to be an excuse for the con- tinuation of taxonomy, but taxonomists directly or indirectly have made too many contributions to knowledge to apologize either for their exis- tence or their errors. Rather it is intended to prophesy how little credit will be given to the taxonomists and collectors and to emphasize the reason for the relatively small amount of support received by them today. It is this lack of support that has made it extremely difficult for American systematists to publish the longer monographic studies of which there is great need. Part of the difficulties arise from the fact that the papers submitted for publication are in competition with those dealing with current scientific fads or else those having more immediate practical value and they are accordingly judged to be of less importance. The other side of the story is that most journals are published by academies, societies or institutions which for the most part must limit the privilege of publication to their members or staffs, and since the membership is comprised of many individuals with almost as numerous FOREWORD 8 interests, articles must be limited in number and length in order that all may have a fair opportunity to make public the results of their investigations. To remedy this situation and to make possible the pub- lication of longer taxonomic papers, it has been decided to publish the present journal which for the time being will include articles up to one hundred pages in length and that deal with taxonomy, cytology, morphology and the related fields that contribute to the classification of the non-vascular cryptogams. The first volume will be published semi-annually over a period of two years, after which it is hoped that the uncertainties brought about by the war will be removed and that the support of the journal by cryptogamic botanists will make possible the establishment of FaRLOwIA as a quarterly journal. Thus those of us who have enjoyed the fruits of his contributions to the develop- ment of American cryptogamic botany by his teachings or through the use of the facilities of the herbarium and library that he had brought together and dedicated to the advancement of this special field of botany, may fittingly acknowledge in 1944 the centenary of the birth of William Gilson Farlow. Davip H. LInDER, Curator FarRLow HERBARIUM HarvarRD UNIVERSITY 1(1): 5-8 FARLOWIA January, 1943 THE AIMS OF FUTURE RESEARCH IN THE ALGAE V. J. CHAPMAN Since the beginning of this century very considerable progress has been made in our knowledge of the algae. Up to then interest centered, as was perhaps natural, around morphology and systematics, and at the same time compilation of flords proceeded apace. Now, more atten- tion is being given to the ecology, physiology and reproduction of the algae, but these branches are still in their infancy. This change in outlook does not imply that no future progress can occur in morphology and systematics, but it does mean that the greatest advances of the future can be expected to be made in these younger branches. For this reason research ‘in them should be encouraged, though the older aspects should not be neglected since they still offer many problems that remain to be solved. Morphology and systematics are an essential background to any botanical subject, and the more complete our knowledge of them, the more likelihood is there of sound and substantial advances in other branches. Today one of the most important needs in algal systematics would seem to be a continuation, with greater intensity, of the attack being made upon the smaller microscopic algae, especially the marine forms. In recent years much has been achieved in this direction. for soil and fresh-water species and Pringsheim with his cultures (Biol. Reviews 16: 198. 1941) has indicated the best method to be adopted. Here one may perhaps express a hope that someone will soon publish a book describing all these forms with the provision of a key for their identification. There are still, especially in tropical and sub-tropical marine habitats, a number of species which require investigation and description. One may mention here the encrusting species of all groups: these have been studied fairly thoroughly in Europe and to a somewhat lesser extent in North America. Because they are so inconspicuous and often are not found in fruit, relatively few phycologists have studied them and many are regarded as rare. Persistent search in England has shown that this is not correct and that the majority have a far wider distribution than has been ascribed to them. This will no doubt be found to be true in other countries. Our knowledge of such forms in tropical and sub-tropical regions is exceedingly scanty and the position is not much better in the southern hemisphere. There are certain genera, such as Enteromorpha, which I have been studying for some years, and Cladophora, which require to be investi- gated with a view to monographic treatment. At present it is quite impossible to identify with any degree of certainty some of the Clado- phorae, more especially the tropical and sub-tropical forms. Valuable studies of this character have been undertaken by Kylin (Lunds Univers. Arssk. 20: 1924) for the Delessariaceae in the Rhodophyceae and by 5 6 Fartowia, Vou. 1, 1943 Parke (Hartley Bot. Lab. Pub. 9: 1933) for the Mesogloiaceae. Even the common species of Fucus are in need of further investigation in spite of excellent work upon them by the earlier phycologists, e. g. Sauvageau. The salt marsh forms of this genus are particularly difficult to determine and yet they play an important part in the algal ecology of salt marshes. Numerous varieties of Fucus species have been described and it is only by collecting and examining forms from many different localities that their status will finally be determined. If we turn from systematics to morphology a principal need would seem to be a study of the developmental stages among members of the Fucales. In recent years Delf (Journ. Bot. Lond. 75: 273. 1937, and 77: 129. 1939) and her school have done much towards elucidating the development of the reproductive organs in the lesser-known species of this group, but the young stages have been largely neglected, a fact which can be ascribed to the difficulties of identifying them in the field or with growing them in culture. It is important that this problem should be overcome because it is of very considerable significance in ecology. A very valuable start in this direction has been made by Nienburg (Wiss. Meeres. Abt. Kiel 21: 51. 1931). Culture studies of cer- tain algae have recently forced us to realize that we have not yet deter- mined all those species which have very different forms for their sexual and asexual generations. It has recently been shown (J. Behlan, Beih. Biol. Pflanz. 27: 221. 1939) that Chlamydomonas variabilis and Carteria ovata are alternate generations of the same species and also (P. Korn- mann, Planta 28: 464. 1938) that the same applies to Halicystis ovalis and Derbesia marina. These are comparable to the older classic example of Cutleria and Aglaozonia. With the improved culturing technique we must re-investigate the life-histories of many species with a view to ascer- taining whether they behave similarly. Bliding (Svensk. Bot. Tidskr. 27: 233. 1933) has shown by such means that two well-marked forms of Enteromor pha prolifera are simply two generations of a single species. The investigations of Knight (Trans. Roy Soc. Edin. 56: 307. 1929), Schussnig and Kothbauer (Oest. Bot. Zeit. 83: 81. 1934) and Papenfuss (Bot. Gaz. 96: 421. 1935), which show that Ectocarpus siliculosus may have a different life cycle in different parts of the world, lead us to the conclusion that we must not be satisfied with a single study of a life history, however complete it may appear to be. The Chaetophorales is an order for which it is very desirable to have some advance in our knowledge of life cycles and chromosomal conditions. Such knowledge will prove to be of the greatest value phylogenetically. In physiological studies the greatest advances will probably come from biochemistry, especially studies concerned with the pigments. The classification of the algae is primarily based upon pigmentation and hence the discovery of any unusual features, e.g the phaeophycean pig- ment fucoxanthin in Polysiphonia violacea, in Zygnema pectinata and CHAPMAN: RESEARCH IN ALGAE 7 the Chrysophyceae (P. W. Carter, I. M. Heilbron, and B. Lythgoe) (Proc. Roy. Soc. 128: 82. 1939), may lead to far-reaching systematic con- clusions. Apart from this the réle of growth substances in the algae has been but little studied and further investigation may show that they play as important a part in algal development as they do in the phanero- gams. Most phycologists would now subscribe to the existence of ‘sun’ and ‘shade’ algae, but there is not the same agreement upon the criteria that determine to which group any given species should be assigned. Further work upon this problem is necessary in order to arrive at a definite working basis. Since ecology is such a young branch of phycology there are innumer- able problems, any one of which would repay study. Some, however, are more outstanding or of greater urgency than others. So far there has been too little attention paid to the operation of critical levels. The existence of these was first discussed by Colman (Journ. Mar. Biol. Ass. U. K. 18: 435. 1933) and since then by David (Ph.D. Thesis, Aberystwyth. 1941). There is no doubt that such levels are of prime importance but further studies are essential before we can really understand their significance. It is also probable that in different localities the critical levels will vary and this should provide a clue to the dominant factors. More or less associated with this problem is the non-tidal exposure factor. The importance of this on salt marshes has already been emphasized (V. J. Chapman, Journ. Ecol. 27: 160. 1939) but it has not yet been studied for the rocky litoral. It is believed that the incidence of these periods, when no inundation occurs, has a profound effect upon the occurrence of many species, especially when the periods operate during the germling phase. Whether the effect is due to actual lack of water or to loss of water or to some other cause has also to be determined. These remarks particularly apply to the fucoids and here we can appre- ciate how important it will be when. we can distinguish with certainty in the field from their very earliest stages, the germlings of the different species. Up to the present, largely because of this difficulty, insufficient attention has been paid to the germling phase. In the case of Ascophyl- lum, for example, this would seem to be all-important because it is absent from areas exposed to strong wave action: young plants are not to be found in such places so that the inhibitory effect takes place at a very early stage. Studies of recolonization after denudation or on new surfaces, such as those by Rees (Journ. Ecol. 28: 403. 1940), David (1. c.) and Boken- ham (Ann. Nat. Mus. 9: 1938), will lead to important conclusions and help us to understand the interrelations between different species on the shore. Such work has already shown that colonization by Fucus is commonly preceded by the development of an Enteromorpha mat. It has been suggested that this mat provides the necessary humidity for the Fucus germling, but it may equally well provide a suitable substrate 8 FaRLowIA, VOL. 1, 1943 for attachment. In any case the function of this mat in promoting the attachment of germlings requires investigation. The part played in the different communities by microscopic and encrusting species is not prop- erly appreciated, though in this case it is partly due to a lack of adequate systematic knowledge. Yet again studies of areas associated with peculiar environmental conditions, such as that of the Baltic by T. Levring (Ph.D. Thesis, Lund, 1940), cannot help but lead to important results. The abnormal conditions induce morphological changes or ecological varia- tions which can often be correlated with the operation of a single factor. Such areas are not common, but it is important that they should be investigated wherever they occur. Studies of geographical distribution may be regarded as a part of ecology and here there is almost a virgin field. Some work can be carried out with the use of existing floras, but in many areas the floras cannot be employed because of the unsatisfactory condition of their systematics and uncertainty of synonymy. Plant geographical studies of the phanerogams have led to extremely important conclusions, and there is no reason to suppose that similar studies of the algae will not do likewise. In fossil botany the principal interest, so far as the algae are concerned, would seem to be with that new group, the Nematophytales (W. H. Lang. Phil. Trans. B. 227: 245. 1937). It is to be hoped that further material will be discovered that will provide decisive proof as to whether they were algae or not. If an algal nature is confirmed then they will take their place as perhaps the most interesting algae of all time. GONVILLE AND Catus COLLEGE CAMBRIDGE, ENGLAND 1(1): 9-23 FARLOWIA January, 1943 NOTES ON NEW ENGLAND ALGAE Ill. SOME INTERESTING ALGAE FROM MAINE? Roy M. WHELDEN Casual observation of collections of freshwater algae from the numerous waters of Maine reveals: an algal flora of surprising richness; more careful attention shows it to be well worth intensive study to which the following notes may be considered only as an introduction. The collections on which these notes are based have been selected from several hundreds of vials of algal material gathered in every month of the year through seventeen years. Beyond the statement that no collec- tion noted here was made from any large body of water, either lake or river, no description can be made to include the many habitats from which collections were made. One might stress the fact that several separate collections were made from what appeared to be the least promis- ing places, such as a shallow rain pool formed in a foot-path. Despite what seemed to be a sterile habitat, a sample nevertheless was taken and later found to contain an abundance of Closterium macilentum, a large percentage of which was in some stage of conjugation. This and other habitats to be mentioned subsequently seem to justify the statement that no place should be passed by too carelessly, since surprising discoveries may be completely missed by such lack of attention. As originally planned, collections were made as frequently as feasible from as wide a variety of habitats as possible: from densely shaded brooks or pools; from cold water and some from shallow pools in full sunlight when the water at the time of collecting was extremely warm; from crystal clear waters to some of deepest brown color. Final selec- tion of material to be considered is due in part to its rarity, to the oc- currence of anomalous forms, to noteworthy variability, and to stages of reproduction. MYXOPHYCEAE The presence of blue-green algae was frequently noted in a large ma- jority of the collections; in no case did these algae occur in any quantity. One of the most interesting of these was Eucapsis alpina Clements and Shantz (Fig. 1) which was found in abundance in a single station, a vegetation-filled pool in which the water was only a few inches deep, with several feet of slimy brown ooze beneath. The Eucapsis colonies varied in size from small ones having only sixteen cells to large com- pound groups of more than five hundred cells. The first specimens ob- served were collected in early June; later they were found in abundance up to mid-November. Many colonies were examined and the component cells carefully measured. When mature the cells were of strikingly uniform diameter. Different colonies showed a rather wide range of 1Contribution from the Laboratories of Cryptogamic Botany and the Farlow Herbarium, Harvard University, no. 221. : 9 10 Fartowia, Vou. 1, 1943 cell size: many were formed of cells 2-3 » in diameter — var. minor Skuja; others, found with the small ones, were 3-4 y» in diameter. Very many more had cells 6—6.5 » in diameter; often the large-celled colonies were composed of the largest number of cells. The presence of this alga was observed at this station from 1927 to 1940. Not once, despite repeated search, was it found in any other of the extremely numerous habitats, some very like this one and only a few rods away. Frogs and a variety of wading and swimming birds are frequent here; yet it would seem that they have never successfully transported this alga elsewhere. Occurring in the same marsh with Eucapsis, and in many other places throughout the southern half of the state is Glaucocystis nostochinearum Itzigs. This alga seems always to occur sparingly. In one shallow marsh, specimens of Anabaena were found in small numbers, always occurring singly and devoid of any obvious sheath. The spherical cells comprising each filament were 1.5 » in diameter or slightly less; the heterocysts were also spherical and 2—2.5 » in diameter; akinetes were always remote from the heterocysts and were cylindrical, 4.5 » in diameter and 18.5-22 » long. While somewhat smaller in all dimensions, this seems to be Anabaena minutissima Lemm. The only specimens found were from this single station. Anabaena flos-aquae (Lyngb.) Bréb. is found rather frequently throughout this region. Its var. minor W. West was found in one place in large amounts. The numerous filaments were of cells 2.7x 2.0 » in diameter. The heterocysts were 2.8-4 », diameter, and the akinetes, present in large numbers, varied from 5.9x 12 uw to 84x 22 p. The many other species of blue-green algae found in this region may never have been reported as occurring there. However they are of such common occurrence there and elsewhere that their presence may be expected. For the present at least they remain unnoted. CHLOROPHYCEAE Among the green algae one finds many of particular interest. One of these was a species of Closteridium (Fig. 7) which occurred rather abundantly in a small marsh pool. The majority of the solitary cells were semi-circular in shape with one side straight or only slightly curved. Each pole bears a single pointed spine which may be straight but fre- quently is distinctly curved. The cell wall is thin. The single chloro- plastid fills nearly the entire cell and contains a single pyrenoid and also a single nucleus, the latter approximately centrally located in the cell. These cells vary considerably in size, being usually between 75 and 95 » long (including spines) and 30-40 » in maximum diameter. The spines are 11-13 ». long. The smallest cell measured was only 63 p long and 23.5 » in diameter. Among the many cells seen were a large number apparently in various stages of division. The most noticeable fea- ture of these was the presence of two distinct pyrenoids, which probably re- WHELDEN: New Encianp AcaE. III 11 sulted from the division of the single pyrenoid normally present. Once di- vided the two pyrenoids seem to move apart subsequent to which there ap- pears a distinct division of the chloroplastid, leaving a single cell with two distinct plastids. Several of these were seen. Further stages showing that actual cell division did occur were not found. Cells in which these stages occurred were 100-105 y» long but with diameters of the same magnitude as had the normal cell. Despite the rather larger dimensions - and coarser habit this alga seems to agree quite well with descriptions of C. siamensis (W. and G. S. West) Wille, reported in this country by Prescott. Occurring with the Eucapsis considered above and apparently also restricted to the one pool, is a species of Tetraédron which has appeared regularly in collections made throughout many years. The thick-walled cells are triangular with sides generally strongly convex, or occasionally straight or slightly concave. Each angle bears a single stout, curved, straight or rarely slightly hooked spine. The single chloroplastid con- tains no pyrenoid. The length of the cells, spines included, varied from 102-118 », without spines from 77-84 »; the breadth, without spines, from 50-62 ». The length of the spines is 11.3-17 uy. W. and G. S. West in 1895 described, without any figure, as Tetraédron tortum, a species which has much in common with that considered here. Their species was relatively narrower, 44 » being given as maximum breadth, twisted, and with sharp spines about 30 » long. The specimens herein described are much plumper, in end view generally presenting sub-circular outline; they rarely, if ever, show a shape that could be de- scribed as twisted, and they have much shorter spines. This combination of characters seems sufficient to justify describing this as a new species, Tetraédron mainensis n. sp. (Fig. 4). Tetraédron cellulis magnis, tumidis, trigonis, lateribus convexis, vel rectis vel rarissime plus minus concavis; anguli spinas crassas rectas, vel arcuatas vel plus minusque uncinatas ferunt; membrana crassa glabra; cellulae cum spinis 102-118 p. longitudine et sine spinis 77-84 yp. longitudine, 50-62 y. latitudine, 40-51 yp. crassi- tudine; spinae 11.3-17 p longitudine. The desmids have always received much attention, partly because of the striking variety of their many species and their beautifully symmetti- cal shapes. No less: interesting are the many problems they offer — problems of distribution, of habitat, of variability, of reproduction have frequently drawn attention. In the region considered in this paper, desmids are one of the most abundantly encountered groups of algae, and certainly are not devoid of interest. One of the commonest species, occurring throughout the region, is the ubiquitous Netrium digitus (Ehrenb.) Itzigs. and Rothe. Attention was drawn to the numerous observed specimens by the wide range in size: the smallest specimens were 80-85 » long and about 30 » broad, and a well graduated series existed which varied from these smallest specimens 12 FARLOWIA, VOL. 1, 1943 up to those that are 300 » and more long and 90-100 » broad. The cell outlines varied from sub-cylindrical to fusiform, with broadly rounded to truncate ends. Closterium occurs in abundance throughout the region. Collections may frequently be made in which a single species occurs, but more frequently specimens of this genus are found scattered among other algae. Zygospores are not infrequently found, developing at all seasons of the year. One pool gave Cl. rostratum Ehrenb. zygospores in April; in July, zygospores of Cl. Ralfsii var. hybridum Rabenh. were found; and in September Cl. setaceum Ehrenb. was conjugating freely. Usually only a small part of the specimens are conjugating, but occasionally one finds instances in which large numbers are seemingly forced to conjugate at the same time. The Closterium macilentum Bréb. mentioned earlier offers such a case. In this collection many of the zygospores departed considerably from the spherical shape described by Brébisson for this species, and were much larger. The example in Fig. 3 is 53x57 w and rather vaguely angular in shape. The cells contributing to this were 420x 17 » and 460x15 ». The zygospores of Cl. Ralfsii var. hybridum Rabenh. were spherical and 70-80 » in diameter. Many interesting problems other than zygospore formation present themselves here. Sigmoid forms occur very frequently; examination shows that in some cases the cells which appear as sigmoid are really of spiral form. In like manner the striae or ribs so characteristic of many species may depart from the normal arrangement in which they are parallel to the optical margin and run in spirals around the cell. More frequently the spiralling is restricted to the apical portion of each semi-cell. Another feature in the cells of this genus is the gypsum particles found in the terminal vacuoles. Variation of these granules is often great, as in a collection of Closterium angustatum Kitz., in which some cells had vacuoles containing as many as sixteen long slender granules, others 6 to 10 ovoid particles, others two to six broadly cyl- indrical granules, and many but a single large nearly spherical one. The granules in the two ends of a single cell might be quite different. Cells from this same collection showed many with spiral ribs. Some are shown in Fig. 5. Many other species have a similar range of variation in granules. Abnormally shaped cells are not infrequently found in collections of this genus. Already mentioned are the sigmoid and spiral forms. Occasionally more abnormal forms appear. The two examples of Closterium striolatum Ehrenb. shown in Fig. 8 are from a collection in which many such cells occurred. In some the apical portion was almost at a right angle to the cell axis; all degrees of bent cells were found between these and quite normal cells. No cell was found in which both ends were abnormal. In a few cases cells were found in the later stages of division, with the younger semi-cells of the two daughter cells showing the same type of bending. WHELDEN: New ENGLAND ALGAE. III 13 Specimens of Spinoclosterium curvatum Bernard (Fig. 6) have ap- peared in several widely separated parts of Maine. The cells vary some- what, especially in the degree of curvature and the shape and orientation of the apical spines. The latter are usually straight and from 17-21 p long, but occasionally curved spines are found. Many specimens showed terminal vacuoles containing 8-10 small granules. Cells were frequently found containing specimens of a parasitic fungus (Olpidium sp. ?) so frequently found in Closterium and other desmids. Occasionally one finds a collection of desmids which is practically pure, containing but a single species, in the most unexpected places. One such was gathered from a shallow sand pit into which a small turf of Polytrichum moss had slid. No algae were apparent in the clear water some ten inches deep, yet examination showed the mosses covered with masses of Euastrum oblongum (Grev.) Ralfs. Most of the cells were very uniform in appearance, 155-175 » long and 82-90 yp broad. One specimen merits notice, and is shown in Fig. 2 (right). First ob- servation indicated that a zygospore was present; examination showed it to be a cell in which division had gone to the formation of a large somewhat spherical mass 108 pw in diameter with a coarsely irregular surface and dense contents of which the nature could not be determined. The specimen shown in Fig. 2 (left) presents certain troublesome points: it is 161 » long and 80 » broad; in one semi-cell the upper lateral lobes are rounded, suggesting EL. humerosum Ralfs; in the other the correspond- ing lobes are subquadrate with retuse margins and are characteristic of E. oblongum. Anomalous cells result from arrested divisions in other desmids as well as Euastrum. Fig. 10 shows two such cells of Cosmarium bioculatum Bréb., together with a normal cell. In both abnormal cells shown and in several other similar ones seen, specimens of a parasitic fungus occurred. Cosmarium subturgidum (Turn.) Schmidle is a rare species described from India. A smaller forma minor has been found in Asia, Africa and Australia. It is interesting to note that a form should appear in Maine, where it occurred in considerable numbers in a small pool in a Sphagnum bog near the coast. The cells were large, from 2.6—3.7 times as long as broad, distinctly constricted, sinus obtuse, angled; semi-cells ovate with truncately rounded ends; cell-wall smooth except in apical portion where many coarse irregularly distributed pores appear. Vertical view circular. The chloroplastids are parietal longitudinal bands, 7—8 per semi-cell, and of these 3—4 are visible in face view. The cells are 130-145 » long, and 46-48 », in maximum diameter. The isthmus is 38-44 p» broad. These cells are distinct from the species and form by their narrower shape and by the presence of pores at the apex, and so are described as a new form of this apparently rare species. (Fig. 9.) Species of Micrasterias are among the commonest of the desmids of 14 FarLowIA, VOL. 1, 1943 this region, twenty-four species having been found, mostly in abundance and widely distributed. All species show considerable variation in the nature of the lateral lobing. Anomalous forms seem however to be relatively rare. Two such cells of Micrasterias rotata (Grev.) Ralfs are shown in Fig. 15. These occurred in a shallow slow-flowing brook of clear cold water in which there were thousands of cells of this species. The few anomalous forms were all of the same nature, represented by individuals in which one semi-cell had the lobes much distorted and sometimes nearly at right angles to the main axis of the cell. No evi- dence of any disturbing factor could be discovered. Another collection of Micrasterias proved to be of more than passing interest. It was made from a small pool in a brook flowing very slowly through a swamp. Few indeed were the cells found on examination of the material—a few specimens of Closterium, one or two specimens of Staurastrum muticum Bréb., and a somewhat larger number of Micrasterias specimens. All the latter were of a single species, but no two were found to be exactly alike, nor did any cell have its two semi-cells alike. In several cases one semi-cell was quite definitely Micrasterias pinnatifida (Kiutz.) Ralfs; in others a semi-cell suggested M. oscitans Ralfs. By far the greater number of the semi-cells were not easily assigned to any species. Varia- tions were particularly noticeable in the lateral lobes, which appeared now broad, now narrow, nearly entire or provided with prominent apical spines. The variations of the polar lobe were less frequent and often less pronounced, and included an apical margin which varied from straight to strongly convex, and with ends merely angular, or with one tooth or two. Separating the lateral and apical lobes the incision varied from extremely narrow to very broad. In many cases the two sides of a semi-cell varied somewhat, though usually they were of the same general pattern. No noticeable variations could be observed in the other species of desmids present. Four specimens of this Micrasterias pinnatifida (Kiitz.) Ralfs are shown in Fig. 16. The cells are 58-69 pu long, 63-71 y» broad, and the isthmus is 8-12 » broad. Four species of Xanthidium are here found widely scattered and in abundance. These are X. armatum (Breb.) Rabenh., X. antilopaeum (Bréb.) Kiitz., X. cristatum (Bréb.) Kiitz., and X. tetracentrotum Wolle. Other species such as X. subhastiferum W. West and X. pseudobengalicum Gronb. also occur, but infrequently. The commoner species hold par- ticular interest because of the extreme variability which seems to be exhibited. The specimens of X. tetracentrotum shown in Fig. 14 serve to illustrate this very well. All these are from a gathering in early August from a small pond. At the time of collecting, the plants were actively dividing, filaments of 4-8 cells being found in abundance. All cells were of normal size for the species, being 38-45 » long, 35-45 p» broad without spines, and 52-65 p» with them, and 24-26 p» thick. The isthmus was 10—-12.5 » broad. The cells bore a yellowish brown thickened area WHELDEN: New Encuianp Atcae. III tis in the middle of the lateral face; the scrobiculations thereon sometimes formed a semicircle or crescent, but more frequently a complete circle. Within the circle a few accessory scrobiculations often appeared. Great- est irregularity attached to the spines, both in their number and their disposition. The smallest number found was in those cells in which each lateral angle bore a single spine. Other cells bore a pair of spines in each angle, or eight spines for the entire cell. All variations between these were found, the paired spines being in a horizontal plane. One of these is shown in vertical view in Fig. 14, the semi-cell having two spines on one angle and one on the other. The other semi-cell has un- paired spines. In many specimens there were two spines to each angle, but they were in a vertical plane. Often unpaired spines were found above these, the cell shape then becoming subhexagonal instead of ellipti- cal. Many cases were observed in which a prominent papilla appeared instead of a spine. Some specimens presented a decidedly asymmetrical appearance as a result of the very irregular disposition of the spines. All the variations or irregularities noted were cells which could be con- sidered mature, the central ornament being deeply colored or the cell actively dividing. Xanthidium antilopaeum (Bréb.) Kiitz. is much more widely distrib- uted in this region than is X. tetracentrotum Wolle, several distinct varie- ties being found. One of these is Irénée’s var. quebecense. Many speci- mens of the latter are much larger than his, the largest measured being: 100 » long (with spines), 92 » (without spines), 106 » wide (with spines), 90 » (without spines), and the isthmus 16—23 » broad. The central scrobiculations varied greatly in different specimens, but most frequently these were arranged in a broad ellipse within which many others formed a second ellipse or were most irregularly disposed. Sev- eral zygospores were found in the same collection, but none with empty semi-cells attached. They were spherical and about 60 » in diameter, without spines. The surface bore many simple straight spines 11-15 p» long. It seems quite possible that these are zygospores of the numerous var. quebecense cells present. A pool of stagnant water in a large granitic ledge contained large numbers of another form of Xanthidium antilopaeum. The many cells examined are of very uniform appearance and represent a distinct variety. They are of medium size, about one and one-quarter times longer than broad, sinus at first linear, then divergent to a rounded basal angle. Apex usually straight, less frequently slightly convex, in- frequently faintly retuse, the lateral margins straight or slightly convex. Angles rounded, each furnished with a pair of stout upwardly curved (rarely straight) spines frequently yellowish-colored. In a few instances a third spine may exist between or lateral to one or both pairs of basal spines. Central thickened area brownish-yellow and circular or ellipti- cal, and furnished with numerous scrobiculations which are variously 16 FarLowliA, VoL. 1, 1943 arranged. Sometimes these are very faintly visible or even completely lacking. Vertical view elliptical with truncated ends, each bearing a pair of parallel or slightly converging spines. Side view of semi-cell subcircular. Length 74-80 »; breadth (with spines) 87-95 p, (without spines) 60-68 y; thickness 45-48 y», breadth of isthmus 24-36 »; length of spines 13-16 ». Distinct from other species and other varieties in relative dimensions, in its straight apical margin, and the disposition of the spines. This plant seems to have been first described in 1888 by W. H. Hastings, presumably in his local newspaper, as Xanthidium antilopaeum var. truncatum. Soon after W. West described the same plant as X. Tylerianum, collected from Maine. About two years later Hastings raised his variety to species rank, citing at the same time West’s X. Tylerianum (misquoted as X. Taylorianum) as synonymous. It would seem better to keep this desmid as a variety of X. antilopaeum; therefore it becomes Xanthidium antilopaeum var. Tylerianum (W. West) Whel- den comb. nov. (Fig. 17). Arthrodesmus quadridens Wood is one which has frequently appeared in collections from Maine and has always attracted considerable attention, primarily because it seemed so different from all other species of the genus. Recently Prescott and Scott, with good reasons, have removed this alga from that genus, and established for it a new genus, Spinocosmarium. They note that the species S. quadridens shows “a great variation in the degree to which granules and processes are developed”. This is indeed so. However, collections vary greatly in this respect. Frequently one may examine scores of specimens in a single collection and find no obvious variation. In another collection from a habitat closely re- sembling the first, every specimen seems distinct in some way. The most conspicuous variation occurs in the spines which are so striking a feature of the lateral margin. These may be simple or once-furcate; between cells in which all spines are simple and those in which all are furcate, every conceivable combination of simple and furcate spines occurs. In the furcate spines the two forks are usually nearly equal in size, but instances do occur in which one fork may be reduced sometimes to a degree where it becomes merely a minute tooth on one side of the spine. Infrequently spines are borne at the apical angles and are usually much smaller than the lateral ones. Generally the spines occur singly, although occasionally two may be produced on an angle. All specimens I have examined are alike in having a single large rectangular tubercle just above the isthmus. The many other tubercles vary greatly both in prominence and in number, especially in the lateral parts of the cells. In rare cases coarse pores occur in abundance and irregularly distributed in the wall between the tubercles. Generally this wall appears smooth. The cells were 33-42 » long, 51-57 » broad (with spines) and 33-40 p» without; the isthmus was 9-13 » broad; the cells were 24-26 w thick. Three species of Staurastrum merit notice because of their zygospores. WHELDEN: NEw EncuLanp ALGaeE. III 17 On three different occasions zygospores of St. punctulatum Bréb. have been found in abundance, once in mid-May and twice in late summer. The most noticeable feature of these zygospores is the broad range in size, from 36 p» to 62 p» in diameter, the processes included. (Fig. 11.) In several shallow pools Staurastrum Rotula Nordst. (Fig. 12) is a species encountered frequently and generally in large numbers through- out the warmer months, becoming particularly common in August and September. During the entire period the rather large cells show a certain amount of variation; the number of processes may be either eight or nine. These rays usually are quite straight, those of the two semi- cells being parallel. Not infrequently one finds specimens in which the processes are curved outward. The margins of the processes are in most instances coarsely crenulate, but occasional specimens are found with perfectly smooth processes. The verrucae which are so character- istic a feature of this species also vary, being usually rather large and bidentate, in other cases small and conical, and occasionally completely lacking. Some of the specimens have larger dimensions than those usually given, being 42-48 » long, and up to 124 w broad (processes included). Zygospores of this alga were found in very large numbers during August and early September of 1929. They were spherical bodies 36-43 p. in diameter, with many straight or slightly curved processes 28-32 yp long. In some zygospores the apices of all processes were armed with two or rarely three curved teeth; in other zygospores the ends of all processes were simple and rounded; in some, processes of each type occurred, The body of the zygospore was deep brown to black in color. Fig. 12 shows three zygospores with different types of processes, and two cells, one with crenulate and the other with smooth processes. In- frequently zygospores of this species were found infected with a small undetermined fungus. In these the processes were generally much shortened (10-15 » long), coarser than normal, or even unciform. Such infected zygospores were easily recognized as abnormal, even when the parasitic body was not distinctly visible. Staurastrum Johnsonit var. depauperatum G. M. Smith (Fig. 13) oc- curs quite generally in this region, sometimes in abundance, together with the species. Occasional specimens of zygospores of the present species were found. They were spherical bodies of brownish color, 28-32 » in diameter and provided with 12-15 simple straight processes 16-20 yw» long. Disregarding purely taxonomic problems and those dealing with the life-histories of algae, interest in this group of plants has centered around three main questions, their distribution in both a local and a broader sense, the variations that they exhibit and possible explanations of them, and the formation of sexual stages, with some attention being given to the periodicity of their existence. The appearance of sexual stages and especially of zygospores in the 18 Fartowia, VoL. 1, 1943 desmids, is often a rare occasion. To record their occurrence, therefore, seems ever desirable, the more so when notes on any unusual circum- stances attend this; with the accumulation of such cases an understanding of some of the causes leading thereto may be had. Certainly my own observations on zygospores, far more numerous than those recorded above, lend no support to any statement that any one season favors their appearance. Frequently one finds a collection in which many different species of the algae present are fruiting freely, thus suggesting that this event is a consequence of external factors. The collection in which the zygospores of Staurastrum Rotula were so abundant is of this sort, for in this ten different algae, including eight desmids, were forming zygospores. In other cases, large numbers of algae may be present, and only one fruiting. Collections in which a dozen species of Closterium are abundant may show only one forming zygospores. It might then be suggested that an internal factor might be operating. The question of occurrence, or favorable habitats, may be briefly mentioned. Everyone who has given serious attention to collecting algae knows that there are problems involved, in deciding just where collecting may be best. Nor can any rule be given for this. A collection may be very poor in numbers and quantity, yet show rich variations in the material that is present. Apparently identical habitats may show entirely different results, one spot yielding abundant material while one nearby is almost barren. Variations of this kind may be found in two collec- tions made in a monotonously uniform habitat and only a foot apart. It would seem that oftentimes the algae maintain a remarkably fine adjustment to some factor or factors in their environment or within themselves. All too frequently algae, especially desmids, may disappear from some place with startling abruptness, not a trace remaining in a habitat which twenty-four hours before gave very rich collections. The question of variations in algae and the causes of them has at- tracted considerable attention. Jacobsen (1875) was one of the first to consider the nature and causes of variations so frequently observed in desmids. He advanced several possible explanations and noted the wide geographical range of the variations. De Wildeman (1887) paid at- tention to this question, brought together may cases of variations noted by previous workers and concluded that . . . “ce ne sera que par une étude approfondie faite sur de nombreux matériaux de diverses prov- enances que l’on pourra arriver 4 donner, a chaque espéce, la place qui lui convient dans la série et 4 supprimer toutes les espéces dont la creation est basée sur der caractéres secondaires.” Later de Wildeman (1895) noted the frequency of occurrence of variations in the genera Euastrum and Micrasterias. Cushman (1907) found the species of Micrasterias to be very variable, and remarked that “as many of these so called varieties are often seen forming one semicell of a specimen, the other semicell of which is typical, it seems hardly wise to recognize them as WHELDEN: New Encianp Atcag. III 19 good varieties.” Johnson (1894), Stange (1908), Playfair (1910), Ducellier (1914, 1915, 1917, 1918), Woronichin (1926) and Huber- Pestalozzi (1931) are among the many who haye given considerable attention to the occurrence and extent of variations under natural condi- tions, and offered various explanations as to their causes. More recently Lefevre (1939) has studied the problem both in nature and more especially in cultures. His extensive experiments on several genera and species of desmids lead him to conclude that anomalies and monstrosities appear much more frequently in cultures than in natural habitats. He is led to state that “Observations based on material grown in culture can never replace material observed in nature, but will serve only to supplement these and may give proof to matters otherwise left very doubtful.” The notes on variations recorded above are based on materials from natural environment. They lead to certain observations. Monstrosities such as those in Euastrum and Cosmarium do occur infrequently and in many cases at least seem to be accompanied by, if not indeed a con- sequence of, attack by a parasitic organism. It is quite possible that in such abnormal forms as those of Micrasterias (Fig. 15) a parasitic organism is present, but in a stage the presence of which is not yet ap- parent. For certain it is that this genus is frequently infested with parasitic organisms. Possibly such abnormal forms occur only if the parasite enters just before division of the host begins, or in its early stages. It has frequently been stated that the many variations in size, form, ornamentation, and cell contents occur much more frequently in certain genera. Undoubtedly this is true. It is also a fact that in many cases the very nature of the cell makes it difficult to observe variations — small smooth species of Cosmarium cannot offer as much in the way of variety as ornate ones. Where obvious variations do occur it is seldom possible to ascribe to them definite causes. Many factors outside the cell come to mind. Abrupt changes in temperature, and large fluctuations in the amount of substances dissolved in the water have been frequently mentioned. Yet when one observes what happens in two small nearby pools, or examines two collections from points near one another in a single pool, where great variation appears in some of the species in one and as great uniformity in the species in the other collection, one con- cludes that such a factor cannot always operate. Nor do they seem adequate to explain all observations. Many things indicate that there is within the organisms themselves something which may lead to the appearance of variations, and that. this together with other external factors operate together perhaps in very subtle ways to bring about the morphological changes which one observes. Only through observa- tions accumulated slowly, and carefully supplemented whenever possible with observations on controlled cultures, can a solution of these prob- lems of algal behavior be hoped for. Harvarp UNIVERSITY CamBripcE, Mass. 20 Fartowia, VoL. Ll, 1943 LITERATURE CITED Cushman, J. A. 50. Fig. 2. Transverse section of the outer edge of a hard pine test block showing ad- vanced stages in fungal hydrolysis of the cell wall caused by Ceriosporopsis halima. The enzymatic activity of the hyphae is almost exclusively confined to the central layers of the secondary wall. The more highly lignified inner and outer layers of the secondary wall are relatively unaffected at this stage of decomposition. Specimen number 1l. X 420. Fig. 3. Transverse section of a spruce test board attacked by Halophiobolus longi- rostris. Note the long neck of the deeply embedded perithecium. Specimen number 22. > 50. Fig. 4. Radial longitudinal section of the same wood specimen shown in Figure 3. Note the complete destruction of cell wall substance in the region surrounding the perithecium. Within the tracheids of the wood are spherical inflated cells produced by hyphae in the lumina. Specimen number 22. 50. Fig. 5. Transverse section of hard pine test block attacked by Ceriosporopsis halima showing early stages in the breakdown of the cell wall by hyphal action. The area shown is two growth rings in from the outer exposed surface of the wood. Note the restriction of hyphal cavities to the central layers of the secondary wall. Specimen number ll. X 230. 467 MARINE FUNGI BarcHoorn & LINDER Pirate VII 1(3): 469-481 FARLOWIA January, 1944 STUDIES IN THE GENUS ZYGOSACCHAROMYCES I. TRANSFER OF PELLICLE-FORMING YEASTS TO ZYGOPICHIA 1 WALTER J. NICKERSON 2 That the taxonomy of even the sporogenous yeasts is in an unsatis- factory state is a statement admitting of little debate. The important work of Stelling-Dekker (1931) served to bring considerable order out of chaos, but the need is still felt for monographic treatments of the vari- ous genera of yeasts. In considering only the organisms of which cul- tures were available to her, Stelling-Dekker followed a practice which has much in its favor. Practically, however, it results in almost com- plete neglect of species not included in the monograph by all except specialists in a particular group. One function of a monographer is the critical consideration of as many of the forms assigned to a group as is possible. Since it is manifestly impossible for one person to have access to cultures of all the species in a large genus, the monographer is forced, by necessity, to make decisions based solely upon descriptions in the literature. However, the scanty descriptions in some of the older literature are frequently valueless, and the names should be cleared from the books as nomina nuda, nomina dubia, etc., as the case may be, unless an authentic culture is at hand for a redescription. The yeasts are a poorly defined assemblage of fungi, chiefly Ascomy- cetes, with a monophyletic standing in the Endomycetaceae favored by some workers, and a biphyletic treatment embracing the Exoascaceae sponsored by others. It seems best to think of a “yeast-state” much as one today thinks of a “colloidal-state” rather than of “a colloid.” This “no-man’s land,” which does not even grant its workers a distinctive name (!), is one cultivated chiefly by techniques borrowed from bac- teriology and mycology as the need may arise. In the past, adherence to the International Rules of Botanical Nomenclature has only too fre- quently been neglected. And a somewhat exclusive adoption of bac- teriological techniques has resulted at times in the neglect of valuable taxonomic tools such as the range of variation, morphology of spores and of asci, geographical distribution, etc. In presenting this paper as the first in a series intended to constitute a monograph of the genus Zygosaccharomyces, the author is aware of some, at least, of the difficulties and limitations imposed by the very na- ture of the organisms involved. Use must be made of any distinguishing * Aided in part by a grant from the Faculty Research Fund of Wheaton College. ? The writer, now serving with the Army of the United States, wishes to thank Drs. D. H. Linder and W. L. White for their help on problems of nomenclature, Drs. Mrak and Wickerham for kindly supplying some of the cultures used, and Dr. H. W. Anderson, University of Illinois, for sending type slides of Z. Chevalieri var. Andersonii with which comparisons were made. 469 470 FarRLowl1A, VoL. 1, 1944 character available, be it morphological or physiological, for the organ- isms in question do not fall into clearly defined, discrete species, but present a picture of individuals more or less concentrated into groups (species) with gradations in all directions away from any given center of concentration. A character which shows a normal statistical distri- bution is in general of little separatory value taxonomically. For instance, if spore sizes within a genus fall along a normal distribution curve when graphed, then this particular character will not serve alone to differentiate groups. On the other hand, a character exhibiting discontinuous variation within a genus serves readily to divide a genus into two groups: i. e., those that have and those that have not. The “centers of concentration” men- tioned above grow out of the overlapping of the curves of discontinuous var- iation. Possibly no microbiologist could be found who would seek to con- ceal the fact that the species concept in regard to yeasts is a fairly intangi- ble thing; but for practical purposes, possibly to parallel or interweave with the development of a comparative biochemistry, it is essential that we be agreed upon principles of intra-generic classification. The inter-specific gradation which exists must be recognized as such and not serve to cause the creation of numerous “poor” species. The existence of strains of microorganisms is clearly recognized by all as a difficult question taxo- nomically; a strain is essentially any isolate of a micro-organism that is in any way distinguished from its “parent” species; usually it is distin- guished in some comparative way, as larger, smaller, more vigorously fermenting, etc. However, in the case of some micro-organisms, where parasitism is involved, one is able to distinguish clearly among forms identical with respect to the usual morphological and physiological cri- teria. See Leonian and Lilly (1943) for an interesting discussion of the species-complex problem in microbiology. Perhaps the existence of strains serves to indicate the direction in which the production of dis- continuous variation within a genus is progressing with time, and per- haps it is a result of an improvement in our ability to make distinctions. The subgenus Zygosaccharomyces Barker, following Klocker (1924) and Stelling-Dekker, is rather large, with the literature on it scattered. Approximately 60 species have been described since Barker (1901) founded the genus. To date, Stelling-Dekker’s treatment has been most complete with a consideration of some 20 species. During the past 5 years the writer has gathered together some 90 cultures including about 40 of the described species. From these cultures, a few stand out by reason of their not meeting the generic requirements of Zygosaccharo- myces. While the writer, in this paper, does not intend completely to delimit the genus, he is in thorough agreement with Klécker and with Stelling-Dekker in excluding all organisms that form films promptly in ordinary sugar-containing liquid media. Before taking up the problem of the relationships existing among species properly belonging to Zygo- saccharomyces, the writer hopes to simplify the problem by transferring NICKERSON: ZYGOSACCHAROMYCES 471 forms which do not meet the requirements of the genus. The remainder of this paper will be devoted to a discussion of 5 pellicle-forming yeasts that have been assigned at one time or another to Zygosaccharomyces, but which should be classified in Zygopichia. METHODS Sources of the organisms studied in this paper are listed under the species concerned in the section on taxonomy. Stock cultures were car- ried as stab-cultures in wort agar (Difco) with 1% added Bacto-agar. Culture studies employed wort and wort agar slants; for liquid cultures a synthetic medium described later in the paper was also used with the addition of 1.5 gm. Bacto-asparagine per liter. Measurements on vegeta- tive cells were made, using an ocular micrometer, after 48 hours growth; averages of the measurements are reported. Sporulation occurred read- ily by seven days on wort agar slants except in the case of Z, Chevaliert, H-175, which responded to culture on Sabouraud’s dextrose agar (Difco) with a 5% addition of Aspergillus filtrate prepared according to Nick- erson and Thimann (1943). Fermentation tests were performed with Durham tubes; if dextrose, fructose, and mannose were not fermented, no other sugar was investigated. Nitrate utilization was studied in a liquid medium using KNOs as the sole nitrogen source; a test for nitrite was also performed; a discussion of the value of examining solely for the presence of nitrite will be found later in the paper. Ethanol utiliza- tion was studied using the synthetic medium referred to but omitting glucose, peptone, and asparagine; recently boiled absolute alcohol was added aseptically after autoclaving to give the medium a final concen- tration of 2%. A discussion of adaptation and tolerance to increasing solute concentrations, including ethanol, will be found later in this paper. Gelatin liquifying ability tested with stabs and plates of wort gelatin (pH 4.8) and malt extract (Difco) gelatin (pH 5.8). Maximum tempera- ture for growth studied in wort and in the synthetic medium. EXPERIMENTAL In view of the findings of Baars (1930) on the sulfate reducing bac- teria, wherein he showed that Vibrio estuarii is merely a salt-adapted form of Spirillum desulfuricans, it was thought worthwhile to investigate some of the yeasts discussed in this paper. As will be pointed out, Zygosaccharomyces Guilliermondi is quite similar to Zygopichia Cheva- lieri, being distinguished chiefly by its reported ability to grow in media containing 25% NaCl. Since the possibility exists, at least, that Z. Guilliermondi is merely a salt-adapted form of Z. Chevalieri, the following experiments on adaptation of various members of Zygopichia were undertaken. To a basic medium consisting of: 3.0 g. (NH) 2SOx, 2.0 g. KH2POx, 0.25 g. CaCls, 0.25 g. MgSOu, 20.0 g. glucose, 2.0 g. Bacto-peptone, 0.10 g. 472 FarLowI1A, VoL. 1, 1944 yeast extract, and distilled water to make 1 liter, various amendments were added as indicated in the respective experiments. The usual meth- ods for quantitative measurement of yeast growth—counting of cells or weighing of cells—are difficult to use with these film-forming species; in most cases only a qualitative statement is given as to the extent of growth; in a few cases, quantitative measures of metabolic products— chiefly total acid production—have been made. All experiments were kept at a room temperature of about 20° C. In tables 1-3 are found data obtained from duplicate experiments on subjecting yeasts to various environments. All cultures were inoculated from vigorous growths on wort agar and observed at intervals for pellicle formation; usually the lower concentrations supported a good growth. After varying intervals of time, higher concentrations not showing growth were inoculated from the next lower concentration showing a pellicle; a sterile flask of the higher concentration in question was also inoculated —data reported are based on the inoculation into this previously ster- ile flask. The concentrations employed for the various amendments are calcu- lated in three ways in table 1: as per cent final concentration, as molarity, TABLE 1—SALT TOLERANCE Total NaCl concentra- Z. Chevalieri M.1 2. Chevalieri v. Andersonii H. 177 tion 2% 0.34 M pellicle in 5 days, sediment only, 10 days. 0.61 M 5% 0.85 M pellicle in 10 days. sediment only, 25 days, 1.53 M 8% no growth in 15 days; pellicle in | no growth in 25 days; no growth in 1.36 M 1 day when inoculated from 5%. 5 days when inoculated from 5%. 2.45 M 10% * | no growth in 15 days; pellicle in 1.7M 3 days when inoculated from 8%. 3.06 M 15% no growth in 20 days; faint sediment 2.55 M in 5 days when inoculated from 4.59 M 10%; no pellicle. 20% no growth in 25 days; no growth in 3.4 M 10 days when inoculated from sedi- 6.12 M ment of 15%. and as molarity < 1.8 to represent the effective molarity for the develop- ment of osmotic pressure by a binary electrolyte. It is indicated in table 1 that the strain of Z. Chevalieri used, while able during growth to be- NICKERSON: ZYGOSACCHAROMYCES 473 TABLE 2—SUGAR TOLERANCE AS SHOWN BY PELLICLE FORMATION Total concentration of glucose Organism 2% 6% 10% 14% 18% 22% 30% 42% 0.11M| 0.33M]| 0.55M| 0.77M| 0.99M/ 1.22M| 1.65M/ 2.33M Z. Chevalieri v. 2 2 2 2 2 3 3 7 Andersonti days days days days days days days days come adapted to increasing concentrations of solute, has an upper limit to this adaptation in the region of 4M solute concentration. Thus, while Zygosaccharomyces Guilliermondi is probably just a salt-adapted form of Z. Chevalieri, there is a difference in their salt tolerances of about 3M effective solute concentration. Under the conditions of the experi- ment and with the strain employed, Z. Chevalieri var. Andersonii ex- hibited no ability to form a pellicle on even 2% NaCl.. By 10 days it had formed good sediment, but no trace of pellicle. In the alcohol series (table 3), each of the three organisms formed a TABLE 3—ALCOHOL TOLERANCE Total alcohol Z. Chevalieri var. Z, Chevaliert M.1 4. farinosa ATCC 2252 concen- Andersonii H.177 tration 2% 0.43 M pellicle in 6 days. pellicle in 5 days. pellicle in 6 days. 5% no growth, 14 days; pel- | pellicle in 8 days. pellicle in 6 days. 1.09 M licle on inoc. from 2 % in 1 day. 10% no growth, 16 days; pel- | no growth, 10 days; pel- 2.17 M licle in 5 days on inoc. | licle in 2 days on inoc. | pellicle in 6 days. from 5%. from 5%. 15% no growth, 18 days; pel- | no growth, 12 days; pel- | no growth, 10 days; pel- 3.26 M licle islands in 5 dayson | licle in 6 days on inoc. | licle in 2 days on inoc. inoc. from 10%. from 10%. from 10%. 20% no growth, 18 days; no | no growth, 16 days; | no growth, 15 days; 4.34 M growth 5 days on inoc. | faint sediment only on | pellicle islands in 5 days from 15%. inoc. from 15%. on inoc. from 15%. well-developed pellicle that began to ascend the walls of the flask or tube shortly after being formed. Castelli used this pellicle phenomenon as a characteristic serving to differentiate Z. chiantigiana from Z. Cheva- lieri but it is evident that such an ability is common to most, if not all, members of the genus. TABLE 4—A COMPARISON OF MEMBERS OF Zygopichia Z. Chevalieri Z. Chevalieri Z. Chevalieri Zygosacch. Z. farinosa Z. chiantig- v. Andersonii v. Barkeri Guilliermondii iana Pellicle in liquid + + + + + + media Fermentation - - - - +d,1,m - Conjugation heterogamic heterogamic physiol. heterogamic iso- and heterogamic heterogamic heterogamic Spore shape hat-shaped or spherical and spherical and hat-shaped or spherical or oval spherical or spherical smooth smooth spherical flattened Spore size + 2.5 up +2.0u 44.3 p +2.0 4 2x3uyn + 284 Cell size in wort 34x48.5 uw 4x6.5y 4-5 x 5-7.8 uw 3-5 x 6-9 uw 2-5 x 4-16 2.4-3.6 x 7.2-9.6 pu 2-3 days old Mycelium on agar + _ — +. = + Gelatin liquefac- +21 +21 —20 —60 —60 tion in days Nitrite test _ - * + - - Ethanol utilization; + + * > + + pellicle formation Max. temp. for bud- 42-43 39-40 32.5 34 * 37 ding in wort in °C. Remarks: cells often cells elongate in grows in presence cells soon become 1,5-2.2 x 12-22 p old culture of 25% NaCl elongate central oil globule in spores * No data PLY PROL ‘T “IOA ‘VIMOTUVA NICKERSON: ZYGOSACCHAROMYCES 475 ZYGOSACCHAROMYCES vs. ZYGOPICHIA Following Stelling-Dekker and Klécker, the distinctions between Zygo- saccharomyces and Zygopichia, genera having gametic conjugation pre- ceding the formation of spores, are as follows: | Zygopichia: in wort a dry, dull pellicle within a short time (24-72 hours) after inoculation; spores angular, hemispherical, hat-shaped, or spherical. Zygosaccharomyces: no pellicle in wort, or at most a soft, slimy pellicle when fermentation is nearly complete, which is a considerable time (2-3 weeks or more) after inoculation; renniform, hat-shaped, or spherical spores. ZYGOPICHIA Klécker, Die Garungsorganismen, 3 Auflage, Berlin-Wien. 1924. The genus was established by transferring Zygosaccharomyces Cheva- liert Guilliermond to it as the type. It bears the same relation to Pichia that Zygosaccharomyces bears to Saccharomyces. In addition to the generic characters brought out in the above comparison, there are the following: fermentation, none or only very weak, nitrite tests negative, nitrate assimilation poor, cells of varying shape with a tendency towards pseudomycelial formation. KEY TO THE SPECIES OF ZYGOPICHIA * 1. On wort, a thin, smooth pellicle; no. fermentation 2. Spores hat-shaped; pseudomycelium formed on wort agar 1. Z. Chevalieri 2. Spores smooth only; no pseudomycelium formed 3. Spores small, about 2.0 4 diameter; heterogamic conjugation. 2. Z. Chevalieri vy. Andersonii 3. Spores large, greater than 4.0 ». diameter; gametes morphologically similar 3. Z. Chevalieri v. Barkeri 1. On wort, a thick, dry, wrinkled pellicle; fermentation of glucose, fructose, and mannose only 2. FObaee tis 4. Z. farinosa 1. Zygopichia Chevalieri (Guilliermond) Klécker, Die Giérungsorganismen, p. 289- 290, Berlin-Wien. 1924. Zygosaccharomyces Chevalieri Guilliermond, Ann. Sci. Natur. II Botan. 19: 1, 1914; C. R. Soc. Biol. Paris 1: 442. 1911. Pellicle formed promptly in wort, no fermentation of any sugar, wort gelatin or malt extract gelatin liquefied within 21 days. Cells in wort 3-5x4-8.5 p, pseudomycelium frequently formed. Asci arise after heterogamic conjugation or parthenogenetically; spores round, hemi- spherical or angular, approximately 2.5 w in diameter. Nitrite negative; good growth and pellicle formation with ethanol as the sole carbon source. Maximum temperature for budding, in wort, 42-43° C. Isolation: by Guilliermond from Biliwine (made from tubers of Osbeckia grandiflora) in West Africa. Cultures examined: Guilliermond Lab. de Parasit. no. 617, through Harvard Col- lection no. 175; E. M. Mrak no. 1, orginally from Tanner’s Collection. ® See also table 4 for a comparison of the species. 476 Fartowia, Vou. 1, 1944 Klécker’s disposition of this species was approved by Stelling-Dekker, and nothing found in this investigation warrants further discussion here. A study of adaptation to various chemical environments in the growth of this species will be reported later in this paper. 2. Zygopichia Chevalieri var. Andersonii Nickerson nom. nov. Zygosaccharomyces bisporus Anderson, Journ. Infect. Dis. 21: 341, 1917, nec Zygosaccharomyces bisporus Naganishi, Bot. Mag. Tokyo 31: 107-116, 1917. Pellicle formed promptly in wort, no fermentation of any sugar, gelatin liquefied within 21 days. Cells on wort agar average 4x6.5 p; no pseudomycelium formed on agar. Asci arise after heterogamic conju- gation or parthenogenetically; spores round and smooth, approximately 2.0 » diameter; very little variation from this average. Nitrite negative; good growth and pellicle formation with ethanol as the sole carbon source. Maximum temperature for budding, in synthetic medium and in wort, 39-40° C. Isolation: by H. W. Anderson from human feces. Culture examined: Guilliermond Lab. de Parasit. no. 569, through Harvard Co)- lection no, 177. An interesting coincidence occurred in 1917 when two men in widely separated parts of the world independently gave the same name to yeasts isolated by them. H. W. Anderson, in Urbana, Illinois, described one Z. bisporus, and H. Naganishi, in Tokyo, described the other. The first monographic treatment (Stelling-Dekker) after publication of these names accepted Naganishi’s species and did not mention Anderson’s. It is quite clear that Anderson and Naganishi did not describe the same yeast, for the latter’s species complies with Stelling-Dekker’s emendation of the’ genus Zygosaccharomyces, while the former’s, as here shown, does not. Since it is not possible to determine which specific name has priority, the writer has been guided by Stelling-Dekker’s acceptance of Naganishi’s name.* The culture of Z. Chevalieri var. Andersonii examined is believed to be a transfer of Anderson’s original isolate. It is identical in all respects with Anderson’s description. While this organism is closely similar to Z. Chevalieri, it is sufficiently different to warrant the varietal standing proposed. Similarities between the two forms are in regard to pellicle, absence of fermentation, gelatin liquefaction, mode of origin of asci, and approximate cell and spore sizes. Points of difference are that the pro- posed variety has smooth spores only, no pseudomycelial formation, and a maximum temperature for budding of 39-40° C. (this latter point a very small one, however). 3. Zygopichia Chevalieri var. Barkeri Nickerson var. nov. Yeast G, Pearce and Barker, J. Agric. Sci. Cambridge 3: 55, 1908. (From description given by Pearce and Barker.) In mediis maltatis membrana cito formatur; nullum saccharum fermentatur. Cellulae in mediis maltatis 4-5 x “International Rules, Art. 57. NICKERSON: ZYGOSACCHAROMYCES 477 5-7.8 wu; in culturis altis multae cellulae non nihil longiores. Asci post conjuga- tionem per cellulas formae similis pariunt; parthenogenesis quoque evenit. Sporum globosum leveque longitudo media lineae mediae 4.3 2; spora post conjugationem formata solum in una duae cellulae reperiuntur. Calor maximus gemmandi 32.5° C. (Description as given by Pearce and Barker.) Pellicle formed prompt- ly in wort; no fermentation of any sugar. Cells in wort 4-5 x 5-7.8 yp; elongated cells in old cultures. Asci arise after conjugation of morpho- logically similar cells, or parthenogenetically. Spores round and smooth, average 4.3 » in diameter; when formed after conjugation, are found in only one of the two gametes. Maximum temperature for budding 32.5° C. Isolation: by Pearce and Barker from bottled English cider. Culture examined: known only from the literature. This organism was considered by Klécker as belonging in Zygopichia, but he did nothing about its name. It is closely similar to Z. Chevalieri in pellicle formation, absence of fermentation, and approximate cell‘ size in young cultures, with elongate cells in old cultures. Sufficient differ- ences for the proposed varietal standing are found in possession of “physiological heterogamy” only, formation of only smooth spores, spore size, and the low maximum temperature for budding (32.5° C.). 4. Zygopichia farinosa (Lindner) Klocker, loc. cit. Saccharomyces farinosus Lindner, Woch. f. Brau. 11: 152, 1894. Pichia farinosa (Lindner) Hansen, Centralbl. f. Bakt. 12: 529-538, 1904. Zygosaccharomyces farinosus (Lindner) Papadakis, C. R. Soc. Biol. Paris. 86: 447, 1922. Prompt formation in wort of a thick, dry, wrinkled pellicle; ferments glucose, fructose, and mannose weakly. Gelatin not liquefied; nitrite test negative. Growth and pellicle formation with ethanol as sole car- bon source. Cells in wort 2-5 x 4-16 p, single or in budding chains; no pseudomycelium. Spores round to oval, and smooth, 2x3 y; formed after isogamic or heterogamic conjugation or parthenogenetically. Spores have a refractile granule in center. Isolated: by Lindner in Danzig from beer; by Saito from soy bean sauce. Culture examined: ATCC no. 2252; originally from Tanner Collection. This organism has had a rather checkered career. Discovered by Lindner, who considered it a Saccharomyces, it was shown by Hansen in 1904 to belong to Pichia on the basis of pellicle formation. This yeast was considered by Guilliermond as late as 1920 to have spores produced without conjugation, yet his student, Papadakis, in 1922 showed that a gametic conjugation was clearly involved, preceding spore formation. Klécker, in 1924, on establishing the genus, removed this species from Zygosaccharomyces along with Z: Chevalieri. Since Stelling-Dekker has reaffirmed this species, and nothing has turned up in the course of this investigation to raise any question, nothing more need be said. 478 Fartowl1A, VoL. 1, 1944 DOUBTFUL SPECIES 1. Zygosaccharomyces Guilliermondi Dufrénoy, Bull. Soc. Myc. Fr. 43: 272, 1927. (Description as translated from Dufrénoy.) Prompt pellicle forma- tion in wort; no fermentation; gelatin not liquefied within 20 days. Asci originate after heterogamic conjugation or parthenogenetically; spores hat-shaped or smooth, averaging 2.0 » diameter. Young cells in wort 3-5 x 6-9 u; pseudomycelium formed on agar. Will grow in media con- taining 25% NaCl. Maximum temperature for budding is 34° C. Isolation: by Dufrénoy in Corsica from the brine of the normal citron fermentation. Culture examined: known only from the literature. Dufrénoy used Lindner’s relatively inaccurate “little” method for study- ing fermentation of this yeast, and observed a slight fermentation of fructose, but none of glucose. Since it is highly unusual, if indeed pos- sible, for an organism to ferment any sugar and yet not ferment glucose, it is here considered that this yeast actually has no fermentative ability. This form is closely similar to Z. Chevalieri in regard to pellicle forma- tion, origin of asci, size and shape of spores, pseudomycelial formation, and lack of fermentation. It is doubtful that sufficient differences exist between the two to warrant varietal standing. It is a question whether this species should not be regarded merely as a salt adapted form of Z. Chevalieri, in view of the results of experiments on adaptation re- ported early in this paper. The time elapsed (20 days) in the judging of gelatin liquefaction as negative is somewhat shorter than one usually considers sufficient when working with yeasts. Preliminary findings which will be published at a later date indicate that high osmotic pressures have an inhibitory effect on proteolytic exoenzymes involved in gelatin liquefaction by two of the yeasts discussed here. This finding may pos- sibly be of interest in view of the results obtained by Johnson et al (1942) on the effect of barometric pressure on the luciferase mechanism of phosphorescent bacteria. In any event gelatin liquefaction is an un- satisfactory character on which to differentiate a species because of our lack of knowledge concerning its distribution among yeasts and the in- fluence environment may play upon it. In spite of this close similarity between Zygosaccharomyces Guilliermondi and Zygopichia Chevalieri, the author does not choose to reduce the former to synonymy with the latter at the moment. Dr. Dufrénoy, in a personal communication, in- forms me that his culture is possibly still in existence in the collection of Prof. Guilliermond at the Sorbonne. Until such time as the culture can be examined or is known definitely to be lost, decision may be reserved. 2. Zygopichia chiantigiana Castelli, Archiv f. Mikrobiol. 9: 449-468, 1938. (Description as translated from Castelli.) In malt after 24 hours at 25° C., cells globose or elongate, often slightly curved in the form of a sausage; cells usually contain a refractile globule; single or in groups of 5-6 cells. Average dimensions 3.8-4.5 x 7-9.5 ». After 5 days cells elongate, frequently forming pseudomycelia. On malt agar, young cells NICKERSON: ZYGOSACCHAROMYCES 479 3.8-5x 7-9 3; many cells possess extruded tubes after 3 days. By 7 days heterogamic conjugation abundant with some asci formed par- thenogenetically; usually 2 spores per ascus, spores globose or liek flattened, usually containing a central refractile globule; average size 2.6-3 p. On liquid media, as wine must or malt, a pellicle is rapidly formed, smooth and gray-white at first, it soon becomes strongly wrinkled and hazelnut in color. There is no fermentation of any sugar. Growth with ethanol is positive; scanty assimilation of nitrates; gelatin is not liquefied in 60 days; maximum temperature for budding Si: GC. Isolated: by Castelli from wine musts from the classic Chianti region and hills near Florence. Culture examined: known only from the literature. Castelli in his description of this species pointed out the close sim- ilarity to Z. Chevalieri, but was of the opinion that his organism differed sufficiently to warrant a new specific name. However, the characters used by Castelli to separate this yeast do not seem sufficiently strong to the writer to warrant such a separation. Cell size is a quite variable quan- tity with yeasts, and it is well known that the composition of the medium plays an important réle in this variation. He is in error, in the writer’s ‘opinion, in saying that Z. Chevalieri forms only a weak pellicle (velo sottile) on liquid media. The writer has repeatedly obtained films with authentic cultures of Z. Chevalieri that could be described by the words used by Castelli for the pellicle made by his yeast. Gelatin liquefaction is a rather uncertain character for differentiating yeasts, as previously pointed out. Rather than reduce this organism to synonymy with Zygo- pichia Chevalieri, the author prefers to follow the course chosen pre- viously, and await the time when it may be possible actually to examine this yeast. 3. Zygopichia Chevalieri var. fermentati Saito. | Such a species is listed in the 1940 catalogue of cultures issued by the Centraalbureau voor Schimmelcultures at Baarn. The author has been unable to find any reference in the literature to the name. It is listed here solely for the sake of completeness. EXCLUDED SPECIES The following species were placed by Klécker in Zygopichia. Stelling- Dekker reclaimed them for Zygosaccharomyces, for, though they may produce a pellicle on liquid media, it is only after considerable time and under special conditions (as with high salt concentrations). Zygosaccharomyces japonicus (Saito) Guilliermond, Rev. Gen.. Bot. 20: 32, 1908. Zygosaccharomyces japonicus var. soya (Saito) Stelling-Dekker, J. c. 1931. Zygosaccharomyces major var. salsus (Takahashi and Yukawa) Stelling-Dekker, I. c. 1931, A study of cultures of these organisms, obtained from the American Type Culture Collection, or from the Northern Regional Research Lab- 480 FarLowiA, VoL. 1, 1944 oratory of the Dept. of Agriculture, showed that a pellicle is not formed promptly in ordinary sugar-containing liquid media. Stelling-Dekker’s exclusion of these species from Zygopichia seems entirely justified. DISCUSSION At present the only character on which the genera Pichia and Hansenula are separated is that of nitrate utilization. As Mrak et al (1942) have pointed out, this is hardly a satisfactory condition, and they have re- vived, in part, Baltatu’s (1939) suggestion of combining these genera as subgenera under some single generic heading. Baltatu lumped Pichia and Hansenula together as subgenera of Myco- derma but his change has not been unequivocally received. To the writer it seems a mistake to place sporogenous yeasts in subgenera under an asporogenous genus. It would be a much better practice to combine the sporogenous genera and to allow the imperfect or asporogenous genera, in this case Mycoderma, to stand until their perfect stages have been clearly demonstrated.> Using grape must, Baltatu claimed to have induced sporulation in fourteen species of Mycoderma. While it is highly possible that the future might see the discovery of spores in other indi- viduals of this imperfect group, it is preferable to transfer at such time the species involved to the perfect group rather than to align them in their entirety at the present time. Since the name Pichia has priority over Hansenula in point of time of establishment and would take precedence in the event of a merging of these genera, the writer feels justified in adding to the literature the combinations proposed in this paper. While data on nitrate utilization are not available on two of the forms discussed, the writer is of the opinion that the close similarity between the proposed variety Barkeri and the parent species is sufficient justification for its inclusion here. To the author, it seems more satisfactory to test for the presence of nitrite than to report on the extent of growth with nitrate as the sole source of nitrogen. Excellent tests for nitrite, a reduction product of nitrate, exist (c. f. Manual of Methods, 1942). By using one such (sulfanilic acid and a-naphthylamine), it was found that all cultures of Zygopichia examined were completely negative; 3 species of Pichia examined were also negative, while 4 species of Hansenula and one of Zygohansenula gave the brilliant reddish color of a positive test for nitrite, signifying nitrate reduction. The usual difficulty in testing for nitrate utilization is that in eliminating sources of nitrogen other than nitrate from the culture medium, one invariably eliminates all sources of growth- nutrilites. Since the test is not to see if the yeast can grow in the ab- sence of “bios,” and since it is rather troublesome and expensive for routine work to make up a solution containing the known growth nutrilites as pure chemicals, the author feels it is preferable to permit yeast growth * See International Rules, Art. 57. NICKERSON: ZYGOSACCHAROMYCES 481 by adding a trace of yeast extract (0.10 g/liter) to a medium consisting otherwise of salts, glucose, and KNO3. Yeasts negative for nitrite make a meager growth, while yeasts that reduce nitrate flourish. Since the results of the nitrite test are clear-cut, it obviates the necessity of de- scribing the amount of growth in words. In cases where the nitrite test was negative, it was found, using the zinc dust test, that nitrate still ex- isted as such in the medium, thus precluding the rather obscure possi- bility that nitrate had been reduced beyond the nitrite stage. WHEATON COLLEGE Norton, Mass, BIBLIOGRAPHY Baars, J. K. Over sulfaatreductie door bacterien. Thesis, Delft. 1930. Baltatu, G. Mycoderma als echte Saccharomyceten. Zentralbl. f. Bakt. II, 101: 196-225. 1939. Guilliermond, A. The Yeasts. New York, 1920. (Translated by F. W. Tanner.) Johnson, F. H., D. Brown & D. Marsland. A basic mechanism in the biological effects of temperature, pressure and narcotics. Science 95: 200-203. 1942. Klécker, A. Die Garungsorganismen. Berlin-Wien. 1924. Manual of Methods for pure culture study of bacteria. Geneva, N. Y., 1942. Leonian, L. H. & V. G. Lilly. Induced autotrophism in yeast. Journ. Bact. 45: 329-339. 1943. Mrak, E. M., H. J. Phaff, R. H. Vaughn & H. N. Hansen. Yeasts occurring in souring figs. Journ. Bact. 44: 441-450. 1942. Nickerson, W. J. & K. V. Thimann. The chemical control of conjugation in Zygo- saccharomyces II. Am. Journ. Bot. 30: 94-101. 1943. Stelling-Dekker, N. M. Die Sporogenen Hefen. Amsterdam. 1931. 1(3): 483-488 FARLOWIA January, 1944 A NEW GENUS OF THE SCLEROTINIACEAE H. H: WHetzex * My conservative critics will doubtless raise an eyebrow at this presenta- tion of another monotypic genus in that group of stromatic inoperculate Discomycetes for which I have erected the family Sclerotiniaceae.” When I began my studies on these forms, now some 30 years ago, mycol- ogists treating any of these species commonly tossed them more or less indiscriminately into one of three or four genera, most frequently into Sclerotinia or Ciboria, genera erected by Fuckel back in 1869. My in- ‘vestigations on these forms indicate that there are at least 15 or 20 well marked generic groups in this family. Those which appear to me to have acceptable generic status are: Sclerotinia’ Fuckel 1869; Ciboria® Fuckel 1869; Rutstroemia Karsten 1871 (in part) emend. Rehm 1893 ;4 Stromatinia Boudier 1907;5 Phaeociboria von Héhnel 1918;® Lambertella von Héhnel 1918 (see also Whetzel 1943); Monilinia Honey 1928; Septotinia Whetzel 1937; Ovulinia Weiss 1940; and Martinia Whetzel 1942. It will be evident to my incredulous reader that I still have a substantial number of genera up my sleeve which I hope to spring upon the mycological public shortly. Of the ten above listed genera, four are as yet monotypic, but as I discovered in the case of Lambertella, they are likely to lose that status as more intensive searches in the field uncover undescribed species of the Sclerotiniaceae. With this orientation of my critical mycological fellows, I shall now proceed to add another monotypic genus to this growing family. A COPROPHILOUS SPECIES While on a collecting trip with some of my students at Malloryville, New York, on June 22, 1942, I picked up a small dung-ball, about half an inch long by a quarter of an inch in diameter among leaf mold on the hillside about the pitcher plant bog. The dung-ball looked like that of a partridge. It bore several tiny brown apothecial cups borne on the tips of long hair-like stipes. Ascospore shootings on potato dextrose agar gave a striking, black, indeterminate stroma (Fig. 2-3) from which there presently developed numerous long-stipitate apothecia like those on the original substrate. 1 Acknowledgments: I am grateful to Drs. David H. Linder and W. Lawrence White for critical suggestions in the preparation of this paper and to Dr. Rolf Singer who has kindly written the Latin descriptions. * See Whetzel, 1943, p. 18, footnote 2. ® Both these genera need to be revised and emended with Sclerotinia sclerotiorum (DBy) Fckl. and Ciboria caucus (Rebent.) Fckl. to be taken as type species respectively. “See White, 1941, p. 163, 169. 5 To be restricted to forms similar to the type species, S. rapulum Boud. ®*See Whetzel and White, 1940, p. 614. : 483 484 FarLowlia, Vor. 1, 1944 The dung-ball, meanwhile in a moist chamber, failed to produce more apothecia and was discarded. The striking characters of the apothecia which developed so abundantly from the stromata produced on the agar substratum lead to a careful study of the species which proves to be quite unlike any other of the fifteen or more genera of the Sclerotiniaceae known to me. I am, therefore, describing it as a new species and plac- ing it in a new genus. Coprotinia gen. nov. Stromate indistincto, nigro, 1-2 mm. crasso; strato corticali 1— vel pluri-stratoso, ex hyphis dense intertextis, atris consistente, medullam includente. Structura stro- matis structurae sclerotiorum Botrytis cinereae simillima. Carpophoris spermatigeris haud visis. Statu conidiali nulla. Apotheciis gregariis, longis, stipitibus gracilibus institutis, plus minusve fuscidulis; disco exiguo prae stipitis longitudine, margine fortiter recurvato in maturis, stipite criniformi, apicibus hypharum glanduliformibus ornato; ascis minutissimis, clavatis, basin versus gradatim attenuatis, apice incrassatis, poro iodi ope leviter caerulescente; ascosporis bi-seriatim dispositis, in apicem asci con- gregatis, minutis, elongate ellipsoideis, uni-cellulatis, hyalinis; paraphysibus cylin- draceis et tenuiter tunicatis. Stroma (on potato dextrose agar) indeterminate, black, 1-2 mm. thick; rind (external and submerged) one to several layers of densely inter- woven, slender, dark brown to black hyphae, enclosing a medulla; medulla of rather closely interwoven, slender, thin-walled hyphae embedded in a rubbery transparent matrix. Stromatic structure very like that of the sclerotia of Botrytis of the cinerea type. Spermatial fruit-bodies not seen. Conidial stage wanting. Apothecia gregarious, long, slender-stipitate, some shade of brown; disc small in comparison with length of stipe, the margin strongly recurved at maturity, stipe hair-like, the surface adorned with scattered glandular hyphal tips; asci very small, clavate, tapering gradually to the base, apex thickened, pore faintly J++; ascospores biseriate, crowded into the swollen apical part of the ascus, minute, long ellipsoid, one-celled, hyaline; paraphyses cylindric and thin-walled. Coprotinia minutula sp. nov. Stromate diffuso, indistincto, nigro, 1-2 mm. crasso; strato corticali ex hyphis dense intertextis, atro-fuscis, gracilibus consistente et uno vel nonnullis stratis com- posito, extus asperulato; medulla ex hyphis intertextis, ramosissimis, anastomosantibus, gracilibus, septatis, hyalinis, 3-3.5 u crassis efformata. Carpophoris spermatigeris haud observatis. Statu conidiali nullo. Apotheciis numerosis e superficie tota stromatis nusquam non enatis, longissime stipitatis; receptaculo 1-2 mm. in diametro, atrocastaneo ad primam maturitatem, vetustis expallentibus usque ad colorem pallide fulvidum, tenui, margine prima juventute fortiter recurvato; stipite gracillimo, criniformi, usque ad 20 mm. longo et % mm. crasso, sursum pallido, atro deorsum, apicibus hypharum glanduliformibus ornato; ascis brevibus, crassius- culis, clavatis, saepe conspicue inflatis sub ipso apice, 31-47 x 3.64.8 uw, apice ro- tundato, incrassato, poro iodi ope leviter caerulescente; ascosporis 8, minutis, bi- seriatim dispositis et in tertiam superiorem inflatam congregatis, elongate ellipsoideis, 4.8-6.0 x 1.5-2 y, hyalinis; paraphysibus cylindraceis, tenuiter-tunicatis nec septatis nisi ad basin, 3-3.5 x 33-45 uw. Stroma (on potato dextrose agar) diffuse, indeterminate, black, 1-2 mm. thick (Fig. 2-3) ; rind (external and submerged) composed of densely Joe i ea ee aaa Fig. 1-3. Coprotinia minutula: 1, Apothecia produced in culture, X 2; 2, Stroma formed on potato dextrose agar, photographed from upper surface, X 1; 3, Same as 2 but photographed with Petri dish inverted, x 1. 486 Fartowla, Vor. 1, 1944 interwoven dark brown, slender hyphae, one to several layers thick, the external portion here and there double with narrow strip of medulla be- tween the two layers; the surface rough and where not imbedded in the agar covered with a felty layer of loose brown hyphae; medulla of rather densely interwoven, much branched and anastomosed, slender, thin-walled, 5 Fig. 4-6. Coprotinia minutula: 4, Diagram of section through disc showing its general form at complete maturity, < 80 approx.; 5, Longitudinal section from lower part of stipe showing structure of stipe and protruding hyphal tips, x 1290; 6, Asci, paraphyses, and ascospores, « 1290. septate, hyaline hyphae, 3-5.5 » in diam., embedded in a transparent rubbery matrix. Spermatial fruit-bodies not observed. Conidial stage wanting. A pothecia arising in large numbers from all over the surface of the stroma (Fig. 1), very long stipitate; receptacle 1-2 mm. in diam., dark chestnut brown when first mature, bleaching out to “pale ochraceous tawny” (R) with age, thin, the margin very soon strongly recurved giving the appearance of tiny toadstools; stipe very slender, hair-like, up to 20 mm. long by 1% mm. thick, pale above, dark below, adorned with short glandular hyphal tips; asci short, stout, clavate, often markedly WHETZEL: NEw GENUS OF SCLEROTINIACEAE 487 inflated or swollen just below the tip, 31-47 x 3.6-4.8 yu, apex rounded, thickened, pore faintly J+-; ascospores 8 per ascus, minute, biseriate and crowded into the swollen upper third of the ascus, narrow ellipsoid, 4.86.0 x 1.5-2.0 » hyaline; paraphyses cylindric, thin-walled and aseptate except at the base, 3-3.5 x 33-40 yp. Hasirat: Collected only once on a small dung-ball of some unkown animal at Malloryville, N. Y., June 22, 1942. Known chiefly from stromata and apothecia developed in cultures derived from ascospore discharge on potato dextrose agar and on sterilized wheat in laboratory at Ithaca, N. Y. Type Specimen: C. U. Pl. Path. 31585. Stromata developed on potato dextrose agar and on sterilized wheat kernels. — This little discomycete presents several distinctive characters. Copro- philous species in the Sclerotiniaceae are rare. The only other species known to me to occur on dung is the form recorded by me” but collected by R. F. Cain on rabbit dung near Lac St. Joseph, Quebec, on August 25, 1938 (C. U. Pl. Path. 29646). That form I have referred to Mar- tinia panamaensis Whetzel. The species here described is, however, quite different since the apothecia develop from a diffuse stroma rather than from tiny distinct hemisphaerical sclerotia, and the spores are hyaline instead of colored. In certain of its aspects, Coprotinia minutula, reminds one at once of certain species of Lambertella. It cannot, however, be referred to that genus, since its stroma is of a more highly developed type, being in the structure of its rind and that of its medulla much more like the sclerotium of a Botrytis of the cinerea type. This, together with its colorless spores, sets it off not only from Lambertella but also from Martinia, although its apothecia, asci and ascospores are strikingly like those of the latter genus. It is perhaps most closely related to Rutstroemia, but the slender ' hair-like stipes of the apothecia would seem to deny reference of it to that genus. One of its most peculiar characters is the gland-like hyphal tips which adorn the stipe throughout its length and over the underside of the receptacle. These are of variable length, usually of 1 to 5 short cells, the apical cell being swollen and filled with an olivaceous-green substance of an oily character (Fig. 5). DEPARTMENT OF PLANT PATHOLOGY ~ CornELL UNIVERSITY IrHaca, NEw York 7™See Whetzel, 1942, p. 589. 488 FartowlA, Vou. 1, 1944 LITERATURE CITED Boudier, E. Histoire et classification des Discomycetes d’Europe, p. 1-221. (Paris) 1907, Fuckel, L. Symbolae mycologicae, p. 1-459. (Wiesbaden) 1869, Hohnel, F. von. Humarina tetrica Quel. In Rehm: Ascomycetes Exs. Fasc. 56 and 57. Ann Myc. 16: 219-220. 1918. Honey, E. E. The monilioid species of Sclerotinia. Mycologia 20: 127-157. 1928. Weiss, Freeman. Ovulinia, a new generic segregate from Sclerotinia. Phytopath. 30: 236-244. 1940. Whetzel, H. H. Septotinia, a new genus of the Ciborioideae. Mycologia 29: 128-146. 1937. . A new genus and new species of brown-spored inoperculate Discomycetes from Panama. Mycologia 34: 584-591. 1942. . A monograph of Lambertella, a genus of brown-spored inoperculate Dis- comycetes. Lloydia 6: 18-52. 1943. and White, W. L. Mbollisia tetrica, Peziza Sejournei and the genera Phaeo- ciboria and Pycnopeziza. Mycologia 32: 609-620. 1940. White, W. L. A monograph of the genus Rutstroemia (Discomycetes). Lloydia 4: 153-240. 1941. 1(3) : 489-490 FARLOWIA January, 1944 BOOK REVIEWS Tue BoretaceaE or NortH Carouina. By William Chambers Coker and Alma Holland Beers. Chapel Hill, University of North Carolina Press, 1943. viii, 96 pp. pl. .1-65, frontispiece. $7.00. The authors include three genera in this treatment: Boletus, Boletinus, and Strobilo- myces, all understood in a conservative sense, i.e., Boletus, by far the largest, with seventy species indicated and described for North Carolina, Boletinus with five species, and Strobilomyces with one species. The book is abundantly illustrated with many good and some excellent photographs, six plates in colors, five plates of spore draw- ings and one text illustration, showing the anatomy of several species. Some of the colored pictures are quite natural, and thus very helpful to the beginner. The micro- scopical pictures, however, are much too schematic and therefore of comparatively little help. Cystidia (with one exception), basidia, tramal hyphae, the covering layer of pileus and stipe, as well as chemical reactions are neither illustrated nor mentioned in the descriptions. The exact color of the spore print, a frequently neg- lected but very valuable character, is correctly indicated in nearly all descriptions. Two keys, both mainly based on macroscopical characters, are useful for everyone who tries to determine Boletes, although in a few instances they may lead the less experienced into trouble because they are not dichotomous, and therefore difficult to use. Those who are not primarily interested in taxonomy may regret the lack of com- mon names in the book, as well as the lack of indications concerning the practical interest of the species described. The authors mention only casually that “this is one of the safest groups ... for the beginner to experiment with, as only a few species are now thought to be poisonous.” The warning to keep hands off the whole Miniato-olivaceus group and all of the Luridi can be heartily subscribed to. Many a lay reader, however, may desire more information on the value of the single species for gastronomic purposes, while foresters and ecologists would appreciate more spe- cific data on the important problem of mycorrhizal relationship between forest trees and Boletaceae. The taxonomist, on the other hand, will find it difficult to follow the authors in many of their solutions of questions of nomenclature as well as in the general ar- rangement of the species. He will also regret the little space that has been devoted to synonymy, and the absence of Latin diagnoses for the new species and varieties proposed (Boletus felleus var. minor var. nov., B. satanas var. americanus var. nov., B. viridiflavus spec. noy., B. sub-fraternus spec. nov., B. parvulus spec. nov., B. amer- icanus var. reticulipes var. nov., B. subaureus var. siccipes var. nov.). The interpre- tation of some species of Boletus does not seem to be in accord with the original concept of their authors, as, for example, B. luridus Schaeffer which is evidently different from the plant described under this name in Europe, as the authors them- selves seem to feel. Notwithstanding certain shortcomings mentioned above, the book contains much valuable information for a wide variety of people interested in the study of Higher Fungi, and should be warmly welcomed not only in North Carolina, but also in other Eastern States, and especially in the South where the Boletaceae-flora, though rich and interesting, has never been monographed before. —R. S. and D. lak, bes PLANTS AND Vitamins. By W. H. Schopfer [translation by N. L. Noecker] Waltham, Mass. Chronica Botanica. 1943. ix, 293 pp. illus. $4.75. It may appear unusual to review in a taxonomic journal a book which deals primar- ily with a physiological subject, but after reading the book on plant vitamins, one can not help but be impressed by the part that taxonomy has had in the development of this field of research, and conversely, one is equally struck by the service that a knowledge of vitamins may render to taxonomy by making possible the more normal 489 490 FarLowla, Vou. 1, 1944 growth of organisms that have hitherto been grown on incomplete nutrient substrata. Anyone who has tried to determine fungi from cultures can readily realize what it would mean to have a medium on which these organisms would grow and behave as they do in nature. Thus, for those who deal with organisms in culture, whether or not they have sufficient chemical knowledge to understand the chemistry of vitamins, there may be found ample reason for reading this book and garnering facts that will be of service in their investigations since the author deals not only with the réle of vitamins in the nutrition of the higher and lower plants, but also introduces facts and suggestions on the part these substances play in symbiotism and parasitism. But lest it be thought that only taxonomists may find food for thought in this excellently presented and up-to-date summary of our knowledge of plant vitamins, it should be emphasized that throughout the work the author, while dealing with the occurrence, nature, and function of vitamins and kindred substances, has throughout presented many suggestions which should inspire further research in the allied fields of physi- ology, biochemistry, and genetics. Biologists are greatly indebted to the author for this valuable compilation of facts that have been presented in a careful yet stimu- lating manner, and to the publisher and translator who have made this work available in English at a time when biological discoveries may mean so much. — D. H. L. = * ‘ i " a Mgr 3 = z s tae , : x ‘ a a om ae Z : BD i Nm . : : pe ro = Pe oe ee, ct Se se ae i me Aes ee S eheaaitiony <= = - ; . -~ te . : i a x ¢ ; a na mea) : .: . a j , , ahane . # : = : : 7 P : be : . ; 5 m 2 i. ee Shea = f piel . 7 : ‘ ioe. © ¢ ‘ Z i OP as 77 rs [= ¥ nl Pe ie silt ada Wl ~ Ls na MANUSCRIPT Contributions from anyone will be considered, preference being given to papers dealing with phases of the taxonomy of the Fungi, Algae, -Musci, Hepaticae, and Lichens. Contributors should strive for a clear concise style of writing. A limit of 100 printed pages is set for any one article, but it is hoped that a paper of such length will be so arranged as to be divisible in two approximately equal parts. Writers are responsible for the accuracy of references to literature cited and to text illustrations. Papers containing lists of citations and descriptions of species should maintain a uniform and orderly arrangement of the parts of such citations, etc. Manuscripts must be typewritten, double spaced throughout, with a 144 inch margin on the left, on white paper, preferably 814x 11 inches. Because of the cost of reproducing photographs, carefully made line drawings in black ink are preferred, For the same reason, as many figures as are consistent with clarity should be grouped together to make up a plate. Numerous small figures in the text are to be avoided. Photographs, when necessary, should be printed on glossy, contrast paper and planned for reproduction as full page plates 4%4 x7 inches, the standard size for all plates. Avoid combining line drawings with photographs. To comply with the International Rules, Latin soe rvons must be soppued to validate new species and genera, PRICE OF REPRINTS Reprints should be ordered from the editor when proof is returned, 4pp. 8pp. I6pp. 32 pp. 48pp. 64pp. 50 copies ..... pge aves os $2.48 $3.62 $5.87 $10.81 $16.27 $19.95 WD copies 2 S98. GAG 990) A ge Bah tor ae PO Mapies ec es 450 631 960 1938 2703 34.04 Additional per C. .......... 128: 144 271 § 483 7.65 . 8.97 _ Covers for reprints, printed with title of article, author, source; cost for first fifty — $2.63. additional —1% cents each. PUBLICATIONS FOR SALE CLINTON, G. P. North American Ustilagineae. 1904. (Proc. Bost. Soc. Nat. Hist.) Unbound. .............. $ 2.00 FARLOW, W. G. Icones Farlowianae. Illustrations of the Larger Fungi of Eastern North America. 103 colored plates by J. Bridgham and L. C. C. Krieger. Folio. 1929. Cloth. JOURNAL OF MYCOLOGY. vol. 5, nos. 2, 3, 1889; vol. 6, 1890-91; vol. 7, 1891-94. Unbound. The lot. ........ PERSOON, C. H. Index Botanicus to his Mycologia Euro- paea, prepared by Dr. and Mrs. D. P. Rogers and Dr. E. V. Recor, dt. 29d Paper. ok cs ca ee SEYMOUR, A. B. Host Index of the Fungi of North America. Poy 08 Cloth THAXTER, R. Contribution towards a Monograph of the Laboulbeniaceae. (Mem. Amer. Acad. Arts & Sci.) Un- bound. | ioe a UR ee beter Us etre Weigies a MO ee THAXTER, R. Entomophthoreae of the United States. 1888. (Mem. Bost. Soc. Nat. Hist.) Unbound. .............. WHITE, W. L. & G. D. DARKER.. Supplement to the Host Index of the Fungi of North America. (In preparation.) Postage extra Address the Librarian, Farlow Reference Library, 20 Divinity Ave., Cambridge 38, Mass. 50.00 7.90 29 7.50 6.00 7.00 6.00 6.00 7.00 3.00 Vol. 1 JULY, 1944 No. 4 FARLOWIA _ A JOURNAL oF CrypToGaMic BoTANy EptroriaL Boarp DAVID H. LINDER, Editor E. V. SEELER, Jr., Mg. Ed. J. R. BartHOoLOMEW R. Sincer E. B. Bartram F. VerRDOoRN N. F. Conant W. H. Weston, Jr. H. F. Harris R. M. WrEetpen PUBLISHED BY FARLOW LIBRARY AND HERBARIUM OF HARVARD UNIVERSITY a 20 Divinity Avre., CAMBRIDGE, Mass. FARLOWIA Four numbers compose one volume of approximately five hundred pages. i Subseription price $5.00 per volume. Single numbers $1.50 each. Subscriptions and remittances should be addressed to FARLOWIA, 20 Divinity Ave., Cambridge, M ass., U. 8. A, Numbers lost i in the mails will be replaced only if reported within a reasonable period. CONTENTS OF VOL. 1, NO. 4 Marine Puyrocrocrapuy. By Valerie May.. Ss Gace eo ae GALAXAURA, A Dir.opiontic FLoRIDEAN GENUS WITHIN THE Oeine NEMALIO- nates. By Nils Svedelius........... Ce erat Oe ae a ieG ee es ees 495 MorruHotocic AND Puysiotocic InvesticaTions In Diatoms. By Sten Wiedling. . Pincge eee ceed cl eres ey ee ee yikes 501 NEw AND NotEwortTHy Pururerinse Mosses. By Edwin B. Bartram ii vee OU: On Two New Sprores or Mncaceros wits Norns on M. ARACHNOIDEUS, M. Denticutatus, M. GIGANTEUS, AND M. Granvis. By L. P. Khanna...... 515 SruprEs oN Po.ysticrus Crrcinatus AND Its RELATION To THE Burt-Rot or Spruce. By Roger Gosselin: .........ececececneees Bo Peat a eee .. 525 Norss on Wisconsin Parasitic Funai. IV. By H.C. Greene............ .. 569 A Discussion or Taxonomic Crrreria IN THE a ete gs By Alma J. WGOn §. Ooo eee eRe, hn, Cocoa Pade a a yk eee ahs CV eee 583 SruDIESs IN THE GENUS HELOTIUM, IV. By W. Lawrence White. . Viiercs ORE WPEURG: Ce re races ees pias aen Haha Peoioa ae Ms cee ee Inprex To VotuME1........ ak, acs Bae ea co veers viet ete OM Vol. 1, No. 3 was issued on January 15, 1944. CORRECTION Add 9 INDEX TO AUTHORS AND TITLES, Farlowia 1(4): 625, July, 1944. Ernst-Schwarzenbach, Marthe. The sexual dimorphism of the tropical mosses of the genus Macromitrium, 195-198. Aor Wha - FARLOWIA A JOURNAL OF CRYPTOGAMIC BOTANY wk | JuLy, 1944 No. 4 : MARINE PHYTOGEOGRAPHY VALERIE May From a comparative study of marine floras we may at some date be able to demonstrate the evolution of sea plants in both time and space. At present it is impossible to complete this comparative study since the algal exploration of many parts of the world is incomplete. Further, algal taxonomy lags far behind the corresponding study of. land plants. Thus any theory of phytogeography based on our present knowledge of marine plants rests on an insecure foundation. Variation in algal communities, within small areas, can be explained satisfactorily in terms of changes in exposure to dessication or heavy seas, changes in mineral and gaseous supplies, in intensity of light and heat, depth of water, and the nature of the sea floor. On this there are superimposed seasonal changes in the flora and interaction or competition between organisms. _ When a comparison is made of more widely separated districts, how- ever, such explanations are inadequate. The original conception of algal distribution was that each species had a centre of growth from which it was disseminated following the lines of the coast, each species being influenced by local factors such as depth of water. This idea ac- counted for local richness in certain groups of the flora and also the similarity between the algal floras bordering an ocean. Exceptions were quickly noted, however, many plants (e.g. Codium tomentosum) being - reported as cosmopolitan in distribution. Later work, as explained be- low, has suggested that this supposed cosmopolitan distribution may, in at least some cases, be due to taxonomic errors. By 1849 we find Harvey coming to the conclusion that temperature zones were highly important. Thus, in number of species, the “browns” (Melanophyceae) predominated in the tropics, the “reds” (Rhodophy- ceae) reached a maximum in the temperate zones, and the “greens” (Chlorophyceae) were polar or universal. Zoning by temperature may well be a misnomer for zoning by light intensity, as has been emphasized by Tilden. Besides this zoning by latitude, a certain amount of data indicates zoning by longitude; for instance in the Codiaceae, A. & E. 5. Gepp (1911) state that five large genera are widely distributed in both hemispheres, 491 . 492 FarLowla, VoL. 1, 1944 seven genera are in the Eastern Hemisphere only, one in the West Indies region and one in the Mediterranean Sea. Only very few species occur in both hemispheres. This grouping would seem due to some barrier (past or present) preventing east-west migration. Discontinuity of distribution has been attributed to the presence, or past existence, of barriers such as land masses, ocean deeps, or sandbank deserts. Likewise, wide dispersal has been explained as due to distri- bution by birds, ships, etc., and by ocean currents (speed as well as di- rection being important because the time of life of motile spores is limited). Svedelius (1924) suggested that the present distribution of algae may be due to geological changes in land and sea allowing migra- tion by way of water connections from the Indian Ocean to the Mediter- ranean Sea via Suez, and from the Pacific to the Atlantic Ocean via Panama. Our present knowledge is too meagre to test this theory. Fossil evidence is unsatisfactory because algal forms are usually soft, and hence their remains are scarce except for the specialized calcareous seaweeds of many so-called “coral” reefs. Lucas (1927) reported the discovery of Bytholotrephis gracilis James & Hall, from Victoria, Aus- tralia, “apparently identical” with forms found in the Lower Ordovician beds of North America. Another fossil species was like one found in England. Lucas assumed that Bytholotrephis was an “elemental or gen- eralised type,” dominant in England, North America, Australia, etc. This idea of an elemental or generalized type represents a theory that at one time uniform conditions and marine floras existed throughout the world. Later more temperate conditions prevailed north and south of the tropics and the floras of these zones, subjected to similar environ-— ments, changed along similar lines. Finally, it was suggested that, in the frigid zones, we would have identical representatives which were absent from temperate or torrid zones. This argument may hold for plankton (which are not considered in this article) but Gain (1912) has shown that great differences exist between the marine floras of the Arctic and Antarctic. For instance, he noted that not one species of the large groups of Fucaceae and Laminariaceae was common to both zones, although all species of these groups belong to cool regions. May (1940) applied Jaccard’s Coefficient of Correlation to the marine floras (Chlorophyceae and Melanophyceae only) of Australia, New Zea- land, Britain, and North America; also of Australia-Cape of Good Hope. From figures published by Gain (reference given above) the coefficient percentages for the Arctic and Antarctic floras are: Chlorophyceae 10.5 Melanophyceae 6.9 Rhodophyceae 7.8 All these coefficients are surprisingly alike and show that the degree of uniformity of the floras of the places examined is fairly constant. The _ correlation tends to be higher among the Chlorophyceae than the Melano- phyceae. The Chlorophyceae include more so-called “cosmopolitan” species. May: Marine PHYTOGEOGRAPHY .493 It seems the wide distributions reported for so-called “cosmopolitan” plants may very well be due to faulty systematic work. Setchell (1914) examined Scinaia furcellata, previously reported as “cosmopolitan.” He found that this species was in reality limited in distribution. Plants re- ported under this specific name 4s occurring in other districts were either different species or even different genera. Thus the “cosmopolitan” distribution resolved itself into there being a group of more or less closely related species, each occupying its own limited region. Ectocarpus confervoides, on the other hand, appears to be truly cos- mopolitan, but different phases of its life history predominate in different places. The reason for this is not understood, although the appearance of many algae is known to change with habitat, i. e. the form assumed depends on the environment. Weber van Bosse (1904) reported that many characters used in classi- fying plants “were not implicitly to be relied upon in all species.” These characters included manner of ramification, diameter of conceptacles, and the number or manner in which they appear on the joints, characters which are still widely used. Perhaps the present unsatisfactory con- dition of algal systematics could best be overcome by more care being paid to the criteria used in classification. Certainly we need more knowledge of the specific limits of the plants we deal with before a phytogeographic comparison between floras can be satisfactory. To conclude, phytogeography of marine plants is an almost untouched field. Without more work of systematic nature and further collections being made, this study is unable to advance beyond vague. hypotheses. All we can say with certainty is that latitude and the presence or absence of barriers (either geological or present-day) between suitable habitats appear to control the distribution of marine vegetation. - NATIONAL HERBARIUM SYDNEY, AUSTRALIA BIBLIOGRAPHY Gain, L. La Flore Algologique des Régions Antarctiques et Subantarctiques. Deuxiéme Expédition Antarctique Francaise (1908-1910) Commandée par le Dr. Jean Charcot. 218 pp. 8 pl. Masson, Paris. 1912. Gepp, A. & E. S. Gepp. The Codiaceae of the Siboga Expedition. Monographie LXII of: Uitkomsten op Zoologisch, Botanisch, Oceanographisch en Geologisch Gehied. 150 pp. 22 pl. Leiden. 1911. Lueas, A. H. S. On Additional Occurrence of Bytholotrephis in Victoria. Mem. Nat. Mus. Melbourne No. 7: 157-158. pl. 14. 1927. May, V. A Comparison between Marine Floras. Contrib. N. 5S. Wales Nat. Herb. 1 (2): 94-98. 1940. Setchell, W. A. The Scinaia Assemblage. Univ. Calif. Publ. Bot. @ (5): 79-152. pl. 10-16. 1914. Svedelius, N. On the Discontinuous Geographical Distribution of Some Tropical and Subtropical Marine Algae. Ark. f. Bot. 19 (3): 70 pp. 1924. Weber van Bosse, A. & M. Foslie. The Corallinaceae of the Siboga-Expedition. Monographie LXI of: Uitkomsten op Zoologisch, Botanisch, Oceanographisch en Geologisch Gehied. 110 pp. 16 pl. Leiden. 1904. 1(4): 495-499 FARLOWIA July, 1944 GALAXAURA, A DIPLOBIONTIC FLORIDEAN GENUS WITHIN THE ORDER NEMALIONALES Nits SVEDELIUS One of the most remarkable discoveries in the sphere of floridean re- searches was the observation of M. A. Howe (1917, 1918) that in the genus Galaxaura there is a structural dimorphism of sexual and tetra- sporic plants. This was previously quite unknown in this algal group and the discovery was the more remarkable since this genus belongs to the order Nemalionales among the Florideae, which order comprehends especially haplobiontic types, i. e., such types in which the reduction division occurs immediately after the fertilization of the female nucleus and in which there is therefore no tetraspore formation whatsoever. This structural dimorphism in respect to the anatomy was naturally quite un- known to F. R. Kjellman when, in the year 1900, he published his com- prehensive treatise of the genus Galaxaura and to a great extent based the diagnoses of the sections in particular on anatomical characters. These sections of Kjellman contain either purely sexual or purely tetrasporic plants. After Howe’s discovery, this classification was quite untenable and it is now an important task for algological research to unite Kjell- man’s different species which are, as we now know, only different gen- erations of one and the same species. The consequence of all this is that a close investigation of the development of Galaxaura is highly neces- sary, the more so as the divergent anatomy of the sexual and the tetra- sporic plants also must be made clear from the developmental point of view. I have delivered a contribution to the solution of these problems in a paper “Zytologisch-entwicklungsgeschichtliche Studien tiber Galaxaura, eine diplobiontische Nemalionales-Gattung” (1942) and I will here give a short account of its main results. This seems to be all the more re- quired as Galaxaura in many other respects too proved to be a new and rather divergent type of organization amongst the Florideae. By my investigations, I believe I have established that the sexual and the-tetrasporic plants are also distinguished by a certain difference re- garding their ramification and succession of shoots. The sexual plants have a more lively ramification caused by proliferations from new grow- ing points, whereas the tetrasporic plants have poorer, mainly dichotomous ramification and a’ more continuous apical growth. This seems to depend : upon the fact that the fertile shoots of the sexual plants are commonly almost totally thrown off after fertilization, whereas the tetrasporic shoots are not deformed to such a high degree by the formation of tetraspores, but can survive and continue their growth. The dimorphism regarding the anatomy that Howe first observed is restricted to the assimilating layer of the cortex, and depends on a very early differentiation, just below the growing point, which appears in such a way that the cortex of the tetrasporic plants produce still another cell- layer besides those belonging to the sexual plants. The assimilating cells 495 496 Fartowia, VoL. 1, 1944 of the tetrasporic plants are supported by special stalk-cells and the cells lying between these stalk cells develop in another way. Either they become stretched laterally with the result that the assimilating cells are isolated from each other so that small cavities are formed between them, or else they develop into big, rounded, colorless cells between the stalk- cells. Earlier statements about the cystocarp of Galaxaura are very scarce and, moreover, quite false. Thus the statement that the cystocarp of Galaxaura has an involucre or pericarp formed from the lowest cell of the carpogonial branch is incorrect and derived from a false observation. In this, Galaxaura differs from both Scinaia and Gloiophloea, Pseudo- scinaia and Actinotrichia. On the other hand, Galaxaura in this respect agrees with Chaetangium (Martin 1939). The side branchlets of the three-celled carpogonial branch develop very early and become large, and rich in plasmatic content, but with very weakly staining nuclei. Further, all the cells of side branchlets get fer- tilized diploid nuclei. After fertilization, the diploid nucleus either enters directly into the hypogynous cell (G. corymbifera) or first divides once in the carpogone itself, after which the daughter nuclei migrate into the different inferior cells (G. Diesingiana). In the emigration of the fer- tilized nucleus from the carpogone, Galaxaura corresponds with Scinaia and Chaetangium, but differs from these genera in an important respect: no immediate reduction division occurs in Galaxaura, which thus is a diplobiontic genus within the Nemalionales having special diploid tetra- sporic plants with the reduction division at tetraspore formation. This I have plainly observed in G. Diesingiana and G. tenera. When the diploid nuclei have entered the hypogynous cell, a lively nuclear division arises, and these diploid nuclei enter all the cells of the branchlets of the carpogonial branch except the empty carpogonium itself. Open pore connections now arise between the hypogynous cell and the cells of the side branchlets. The gonimoblast filaments soon begin to develop and elongate in different directions, namely: 1) straight upwards, i.e., through the basal part of the carpogonium; 2) laterally from the side branchlets of the hypogynous cell; 3) from the first or lowest cell of the carpogonial branch. It ought to be pointed out that there is a differentiation between such gonimoblast filaments as grow straight up- wards and begin the carpospore formation immediately, and those that later on grow out laterally, giving rise to secondary gonimoblast centra. Finally, there are also formed sterile nutritive filaments which grow downward and seek out the bigger leading cells in the center of the alga, which cells become twined round by these nutritive filaments. All gonimo- blasts, the primary as well as the secondary, have, however, a common ostiole through the calcified cortical layer. The unity of the cystocarp is maintained in spite of several distinct gonimoblast centers. Regarding the formation of the carpospores, it is to be observed that, after they have been dispersed, regeneration generally takes place so that SVEDELIUS: GALAXAURA ' 497 new sporangia are formed underneath the emptied ones. On the other hand, no paraphyses are formed, just as in Scinaia, Statements about paraphyses in Galaxaura in the literature are false, as are the statements about a special pericarp formed by filaments from lower cells of the carpogonial branch. ‘ Thus the formation of the cystocarp in Galaxaura has been found to be a type which is quite new among the Rhodophyceae, and characterized by the fact that the whole carpogonial branch receives diploid nuclei, and that the whole carpogonial branch also forms starting-points for the gonimoblast. The carpogonial branch can be comprehended as a com- pound auxiliary system. The whole gonimoblast can be divided into primary and secondary gonimoblasts whereby the primary is the one which is formed in connection with the central fusion-cell, and the sec- ondary the one originating from the centers which have proceeded from the horizontally extending gonimoblast filaments. : Galaxaura is also remarkable for the fact that its spermatangia are formed in deep cavities, a very exceptional case among the Rhodophyceae. An investigation of the development has proved, however, that the very earliest stages correspond rather well with those of Scinaia, but in Scinaia the spermatangial branches protrude and push between the epidermal cells to become superficial. In Galaxaura the development proceeds in another direction. The branching is especially lateral, and the sur- rounding cortex. cells stretch around the spermatangial branchlets with the result that they very early become depressed in a deep cavity, the walls of which are covered by the spermatangial branchlets. However different the spermatangial branches in Scinaia and Galaxaura may still seem in fully developed stages, when the development has been clarified, they can without difficulty be classed under a uniform scheme or type of organization. The genus Galaxaura belongs among the very few genera amongst the Nemalionales which are characterized by the formation of tetraspores. As it has now been proved that Galaxaura is a real diplobiont, there is of course no reason to suppose anything else than that the tetraspore forma- tion is also combined with a reduction division. I have also been able to prove this reduction division in two species belonging to two different sections, namely G. Diesingiana and G. tenera. It proceeds normally and the nucleus of the tetraspore mother-cell passes through prophase with spireme and diakinesis with about ten bivalents (gemini). The position of the tetrasporangia varies in different species and on this a division of the genus Galaxaura into sections may be founded better than on the older one which was. based mainly upon anatomical char- acters. Preliminarily, I divide Galaxaura into three sections, namely: Sect. 1. Rhodura (Kjellm.) Sved. emend., characterized by quite sessile sporangia formed terminally or laterally upon rather long assimilation filaments. The sporangia never regenerate. 498 Fartowia, Vou. 1, 1944 Sect. 2. Acrosporangiatae Sved., characterized by terminal sporangia formed upon assimilating cells from a special small stalk-cell. After the dispersion of the tetra- spores, the stalk-cell forms a new tetrasporangium in which the nucleus passes through a new reduction division and a new tetrad is formed. This process is repeated many (5-6) times, and after each regeneration the basal part of the regenerated sporangium always remains as a collar at the base of the new sporangium. Some- times a new and purely vegetative cell is interpolated between the sporangia. Sect. 3. Pleurosporangiatae Sved., characterized by the constantly lateral sporangia originating from the assimilating cells by means of a special stalk cell. Regenera- tion occurs here also but, in contrast to the section Acrosporangiatae, only once. The repeated regenerations combined with repeated reduction divisions is very remarkable and with no equivalent in any other floridean genus. An investigation of the closely related chaetangiaceous genera Scinaia, Chaetangium, and Galaxaura has shown that the first two genera are haplobiontic and lack tetraspore generation, whereas Galaxaura is diplo- biontic with special tetrasporic plants. According to statements in the literature, Gloiophloea and Pseudoscinaia are very probably haplobionts, whereas, as I have now shown, Galaxaura is diplobiontic with special tetrasporic plants. Actinotrichia is also diplobiontic. Thus it is evident, just as I have already pointed out in my Scinaia paper of 1915, that we must be very careful to use haplo- and diplobiontic organization as a sys- tematic character for characterizing the larger groups. Closely related genera behave differently in this respect and in the genus Chantransia even different species behave differently. Thus it is impossible to make. haplobiontic organization into a main character for the order Nemalionales even if the majority of its genera are haplobionts. It seems to me that this character must not be accorded higher taxonomic value than, for example, homospory and heterospory for the taxonomy of the Pterido- phyta. We may say that in the order Nemalionales we find amongst the Rhodophyceae the transition types between haplobiontic and diplobiontic organization, that is, the organization which later becomes the predom- inant one of the higher Red Algae. Regarding the special taxonomy of the family Chaetangiaceae I here give a revised classification in which the two subfamilies Scinaiae and Chaetangieae are maintained with partly altered diagnoses: Subfamily I. Scinaieae (Trevis.) Sved. emend. The lowest cell of the carpogonial branch forms a sterile pericarp around the gonimoblast, i.e. the cystocarp is provided with a wall. The spermatangia are generally superficial. A. Haplobionts: Scinaia, Gloiophloea, Pseudoscinaia. B. Diplobiont: Actinotrichia. Subfamily II. Chaetangieae (Kuetz.) Sved. emend. The sterile pericarp about the gonimoblast lacking; gonimoblast developing from nearly all cells of the carpogonial branch, the cystocarp is thus without wall. Spermatangia are immersed in cavities. A. Haplobiont: Chaetangium. B. Diplobiont: Galaxaura. It appears from this system that diplobiontic organization must have SVEDELIUS: GALAXAURA 499 arisen at different occasions during the evolution of the Chaetangiaceae. This seems to me to be of great interest and it is also of fundamental im- portance for the real understanding of the evolution of the plants from haploid to the prevailing diploid organization. Unrversity oF UppsaLa UppsaLa, SWEDEN LITERATURE CITED © Howe, M. A. (1917). A note on the structural dimorphism of sexual and tetra- sporic plants of Galaxaura obtusata. Bull. Torrey Bot. Club 43: 621-624. (1918). Further notes on the structural dimorphism of sexual and tetra- sporic plants in the genus Galaxaura. Mem. Brooklyn Bot. Gard. 1: 191-197. pl. 3, 4. Kjellman, F. R. (1900). Om floridé-slagtet Galaxaura, dess organografi och syste- matik. K. Svenska Vet. Akad. Handl. 33 (1): 109 p. pl. 1-20. Martin, M. T. (1939). The structure and reproduction of Chaetangium saccatum (Lamour.) J. Ag. II. Female plants. Jour. Linn. Soc. London (Bot.) 52: 115-144, pl. 1. Setchell, W. A. (1914). The Scinaia assemblage. Univ. Calif. Publ. Bot. 6 (5): 79-152. pl. 10-16. Svedelius, N. (1915). Zytologisch-entwicklungsgeschichtliche Studien iiber Scinaia furcellata. Nova Acta Reg. Soc.-Scient. Upsaliensis. Ser. IV. 4 (4): 1-55. - (1939). Uber den Bau und die Entwicklung der Spermatangiengruben bei der Florideengattung Galaxaura. Bot. Notiser 1939: 591-606. - (1942). On the development of the cystocarp in the genus Galaxaura and the auxiliary cells in the order of Nemalionales. Blumea. Suppl. II (Dr. A. A. Weber van Bosse Jubilee Volume) p. 72-90. : - (1942). Zytologisch-entwicklungsgeschichtliche Studien tiber Galaxaura, eine diplobiontische Nemalionales-Gattung. Nova Acta Reg. Soc. Scient. Up- saliensis. Ser. IV. 13 (4): 1-154. r Weber van Bosse, A. (1913-28). Liste des Algues du Siboga. Siboga-Expeditie Monog. 59 (a—d): 1-533. pl. 1-16. * Wa. aa ee ee Go) ls fee”. eee ee ee Wg? gee ge eee nl eT Sr gre ae Cae Pas eel Sap. ake RL a ee Prin oe ey | Ba oN “ oy recy een Ee Pe ae ee AOS SSR eT. aetna cee aay ne ete Ae yee Air Shel Mie ae : A kt ie he 1(4) : 501-502 FARLOWIA July, 1944 MORPHOLOGIC AND PHYSIOLOGIC INVESTIGATIONS IN DIATOMS STEN WIEDLING Having earlier studied culture methods for planktonic fresh-water algae, especially diatoms, the author in 1934 started an experimental investiga- tion of the Nitzschiae with the intention of finding out the conditions of auxospore formation in these diatoms. Both fresh-water forms and those from brackish and salt water were cultured (Botaniska Notiser 1941: 37. 1941). The list of species according to the determinations of Dr. Astrid Cleve-Euler, the grand old lady of Scandinavian diatomology, com- prises the following forms: Nitzschia ovalis Arnott fa., N. subtilis var. paleacea Grun., N. palea var. debilis (Kg.) Grun., N. palea var. tropica? Grun., N. Riuvemeiona Hilse, N. Kiitzingiana var. exilis Grun., and N. communis Rabenhorst. In all, about twenty-five strains were kept and some of them are still being cultured. Three standard substrates were used, all of them based on agar and inorganic salts; ferric citrate, however, was added as an iron source since Carsten Olsen in 1929 demonstrated its superiority to inor- ganic iron salts in plant cultures. For the fresh-water forms, distilled or tap water was added and for the other forms surface water from the Sound was found suitable (cf. Harvey, J., Marine Biol. Assoc. 23: 499, 1939). The results of this investigation on the reproduction of the Nitzschiae were partly not according to expectation (Naturwiss. 31: 115. 1943). Several strains as those of N. subtilis var. paleacea Grun., N. Kiitzingiana Hilse, and N. palea var. debilis (Kg.) Grun. did not show any decrease in the apical length, which instead was constant. This experience is an- other evidence against the universal significance of the MacDonald-Pfitzer rule for diatom variation (J. D. MacDonald, Ann. Mag. Nat. Hist. 3: 1. 1869; E. Pfitzer, Sitzungsber. Niederrhein. Ges. Natur-Heilkde, Bonn 1869: 71; Bot. Zeitung 27: 779. 1869). Other diatoms with constant apical length were earlier found by Allen and Nelson (N. closterium W. Sm. fa. minutissima, Journ. Mar. Biol. Assoc. 8: 421. 1910) and Geitler (Eunotia pectinalis var. minor (Kiitz.) Rabenh., Arch. Protistenk. 78: 1. 1932). The forms with a constant apical length may represent a certain stage in the phylogenetic development of diatoms. Evidently our views on diatom variability and reproduction must be thoroughly revised. Another interesting item was the formation of cell aggregates (“col- onies”) of a typical appearance (“forma multiplex’) in the otherwise solitary N. Kiitzingiana var. exilis Grun. (Botaniska Notiser 1940: 403. 1940). This form also has a specific tendency of passing to a skeleton- free phase (Botaniska Notiser 1941: 33. 1941). The skeleton disappeared after the normal form had been cultivated for several years, during which time the decreasing average apical length had been followed down to a 501 502 FarLtowia, Voi. 1, 1944 certain minimum size which had been reached without auxospore forma- tion. The cause of this might be that the strains in question have, during cultivation, been obtained in clones which consequently lacked sexual partners, in this case necessary for the formation of auxospores. This is in contrast to the skeleton-free forms described by Richter, Bachrach, Geitler, and Stosch which were probably produced through environmental influence on the forms growing vegetatively. The skeleton-free types, although not a part of the normal develop- ment, are of a great interest since they may show resemblance to the phylo- genetic ancestors of diatoms, i.e. unicellular brown-colored algae without a silica skeleton. The common origin of the Bacillariophyta and Phaeo- phyta is thus likely. The morphologic results have as yet been presented only as preliminary communications but the author hopes to be able to issue a more complete and definite report of these parts of his investigations. An interesting feature in the physiologic investigation was the demon- stration of antagonism between the sulphanilamides and p-aminobenzoic acid (Naturwiss. 29: 455. 1941; Science 94: 389. 1941; Botaniska Notiser 1941: 375. 1941). This is the first time that such an antagonism was found to exist in other organisms than bacteria. According to Fildes’ theory (Lancet 238: 955, 1940), p-aminobenzoic acid is thus also an essential metabolite for these autotrophic algae. In the experiments on the antagonism between sulphanilamides and p-aminobenzoic acid, it ap- peared that sulphanilamide inhibited growth more than sulphapyridine, which was in its turn more growth inhibiting than sulphathiazole. In the presence of p-aminobenzoic acid, however, conditions were the reverse. ‘From this it was interpreted that sulphapyridine and sulphathiazole also influenced other biochemical processes than those connected with the p-aminobenzoic acid (cf. Nature 150: 290. 1942). This opinion has found support in observations of other experiments with other organisms (West, Proc. Soc. Exp. Biol. Med. 46: 369. 1941; Dorfman & Koser, Journ. Infect. Dis. 71: 241. 1942; Teply et al, Journ. Pharmacol. 77: 207. 1943). Astra CENTRAL LABORATORIES SODERTALJE, SWEDEN oT, 5: eine Se 1(4) : 503-513 FARLOWIA July, 1944. NEW AND NOTEWORTHY PHILIPPINE MOSSES _Epwin B. BARTRAM Since the “Mosses of the Philippines” was published in 1939, several important and quite extensive collections have added a number of sig- nificant species to the local moss flora which are here placed on record. In the following list of twenty-six species, seven are apparently new to science, twelve are species previously unrecorded from the Islands, and self-explanatory remarks accompany the remaining seven. The supplementary collections from which these records are derived total well over 400 numbers. Only the more noteworthy collections are listed here but the remainder is important from the standpoint of local distribution. The collectors to whom we are indebted and the principal sources of the material are as follows: Luzon and Mindoro — Mr. José V. Santos in 1938; Mindanao, Palawan and Mindoro — Mr. Liborio E. Ebalo, 1939-1940; Mindanao — Mr. A. Lynn Zwickey, Expedition of the Arnold Arboretum, Harvard University, and the Botanical Garden of the University of Michigan, 1938. I am further obligated to Mr. Santos for his labors in packeting and labelling the specimens accumulated by the University of Michigan so that they would be available for study. The collections from Mindanao, Mindoro and Palawan are especially valuable as much field work remains to be done in these areas, especially Mindanao, and it may be long years before these prolific regions are again open to- further exploration. With these additions the Moss Flora of the Philippines is expanded to 931 species representing 184 genera. It is highly probable that the total will exceed 600 species when the possibilities of the Archipelago are more fully developed. The types of the new species are in the writer’s herbarium and in the herbarium of the University of Michigan. A complete set of the series collected by Mr. Zwickey in Mindanao including the duplicate types will also be found in the Farlow Herbarium, Harvard University. FISSIDENTACEAE Fissidens (Crispidium) Zwickeyi Bartr. sp. nov. Fic. 1-4, Dioicus. Caulis 5 mm. altus. Folia circa 10 juga, sicca falcato-flexuosa, oblongo- - ovata, obtusa, breviter apiculata, ad 1.6 mm. longa, 0.5 mm. lata, elimbata, lamina dorsalis ad basin costae rotundata, margines ubique crenulati, cellulae hexagonae, chlorophyllosae, laevissimae, diam. 10 yp, costa percurrens vel infra apicem soluta. Fructus terminalis, seta 2-3 mm. alta, superne arcuata, theca inclinata, deoperculata 0.8 mm. longa, sicca sub ore contracta. Mindanao: Lanao Prov., vicinity of Olangu near Momungan, alt. 400— 500 m., in thin layer of clay on moist rock, 25 Nov. 1938, A. L. Zwickey, B38. Near F, sylvaticus Griff. but I think clearly distinguished from this species by the more broadly pointed leaves, the essentially smooth lamina cells and the inclined or horizontal capsules. 503 504. FarRLowliA, VoL. 1, 1944 Fissidens lagunensis Bartr. nom nov. Fissidens diversiretis Bartr. Philipp. Journ. Sci. 68 (1-4): 21. 1939. I find that the name Fissidens diversiretis was used twice prior to its publication in the “Mosses of the Philippines.” Brotherus describes I. diversiretis from Yunnan Prov., China! followed by F. diversiretis Dixon from Fiji? in 1930. Since F. diversiretis Broth. has priority, I propose the above new name for the Philippine plants. Fissidens Wichurae Broth. & Fleisch. Hedw. 38: (127). 1899. Mindanao: Lanao Prov., vicinity of Dansalan, alt. 700-800 m., on clay hummock, forest floor, Nov. 23, 1938, A. L. Zwickey, 794. New to the Philippine Islands. Distribution: Java. DICRANACEAE Angstroemia orientalis Mitt. Luzon: Mountain Prov., Mt. Data, 26 March 1938, Josée V. Santos, 937a, in part. Previously known in the Philippines from only a single locality on Mt. Pulog. LEUCOBRYACEAE Leucophanes Massarti Ren. & Card. Rev. Bryol. 23: 99. 1896. Mindanao: Lanao Prov. Ulango Mountain, on bark of tree, 26 Feb. to 30 March 1941, Liborio E. Ebalo, 1176. _New to the Philippine Islands. Distribution: Java. This collection, which matches almost exactly Fleischer’s Musc. Frond. Arch. Ind. no. 205 from Java, is distinguished by the longer leaves (7-9 mm.) in contrast to the short leaves (3-6 mm.) of L. octoblephari- oides Brid. CALYMPERACEAE Syrrhopodon hispido-ciliatus Bartr. nom. nov. Syrrhopodon perarmatus Bartr. Farlowia 1 (1): 42. 1943 non Syrrhopodon perar- matus Broth. Journ. Proc. Roy. Soc. N. S. Wales 49: 133. 1915. Mindanao: Zamboanga Prov., Muralong Mountain near Kabasalan, on dead log, 27 Nov. 1940, Liborio FE. Ebalo, 737. New to the Philippine Islands. Distribution: New Guinea. Mr. Dixon has kindly called to my attention that the name S. perarmatus is preoccupied by a New Hebrides moss as indicated above, hence the change. It is an interesting coincidence that this species should turn up in Mindanao so soon after its discovery in Papua, New Guinea. It will be *Symbolae Sinicae, Part IV, p. 11, 1929. * Proc. Linn. Soc. N. S. Wales 55 (3): 270. 1930. BARTRAM: PHILIPPINE MossEs 505 readily separated from S. philippinensis Bartr. by the narrower, rigidly erect-spreading leaves acute at the apex and the more densely hispid costa. Calymperes porrectum Mitt. Linn. Soc. Journ. (Bot.) 10: 172. 1869. Calymperes salakense Besch. Ann. Sci. Nat. ser. 8. 1: 302. 1895. Mindanao: Lanao Prov., vicinity of Dansalan, alt. 700-800 m., on trunk of tree, Oct. 6, 1938, A. Lynn Zwickey, 268. Distribution: Malakka, Java, New Guinea, Samoa.’ A recent study of some Samoan collections indicates that C. porrectum Mitt. and C. salakense Besch. are identical. As Mitten’s name has unques- tioned priority I am reducing C. salakense to synonymy. When the “Mosses of the Philippines” was published I had seen no material of this species from the local area. The Mindanao collection is a robust form showing the upper leaf margins coarsely and distantly toothed, the teniolae well in at the leaf shoulders and the cancellinae slightly rounded above, but these variations are of slight importance. In all the material of this species that I have seen, the abnormal leaves are mixed with the normal ones and reduced above to a bare costa which is gemmiferous at the tip, suggestive of pseudopodia. Calymperes strictifolium (Mitt.) Roth, Hedwigia 51: 127. 1911. Syrrhopodon strictifolius Mitt. in Seemann FI. Vit. p. 388. 1873. Syrrhopodon tuberculosus Dix & Ther. Journ. Linn. Soc. (Bot.) 43: 303. 1916. Mindanao: Lanao Prov., vicinity of Davis Falls, alt. 200-300 m., moist crevices and beneath “overhangs” of rocks, 29 Oct. 1938, A. Lynn Lwickey, 552. | Distribution: Borneo, Fiji, Samoa. Comparison with a part of the type collection of Syrrhopodon stricti- jolius Mitt. fails to reveal any practical distinctions between this species and Calymperes tuberculosum (Dix. & Ther.) Broth. so there is no al- ternative but to reduce the latter name to synonymy. Calymperes (Eucalymperes) Ebaloi Bartr. sp. nov. Fic. 5-10. Caespites densi, sordide virides, haud nitidi. Caules brevissimi, 4-8 mm. alti, dense foliosi. Folia sicca et humida erecta, leniter flexuosa, haud contorta, ad 16 mm. longa, e basi breve ovata longe et anguste ligulata, limbata, sensim late acuta, ‘vix 0.25 mm. lata, lamina inferne sensim contracta, longe supra basin angustissima vel nulla; marginibus ubique et remote denticulatis, ad basin dense spinoso-serratis; costa valida, percurrente, dorso ad apicem solum denticulata; cellulae superiores laevissimae, subquadratae, diam. 10-12 y.; cancellina bene definita, supra acutata, teniola ad basin indistincta, in lamina validiuscula, fere ad apicem attingens, intro- marginata. Seta tenuis 12 mm. longa, laevis; theca cylindrica, deoperculata 1.5 mm. longa. Caetera ignota. Mindanao: Zamboanga Prov., Lilimbrog Mountain near Kabasalan, on tree trunk, 24 Nov. 1940, Liborio Ela Ebalo, 664. Widely distinct from C. serratum A. Br. in the petiole-like base, the intramarginal leaf border and the straight or only slightly flexuose leaves. I have an undescribed species from Fiji with similar leaves in which the oy hoe ig spall aii CR al ac 306 FarLowia, VOL. i 1944, lamina is gradually contracted below to a stretch of bare costa, then wid- ened abruptly again at the shoulders of the leaf base, but in the Fiji plants the leaves are shorter without an intramarginal border, the lamina cells papillose and the leaf shoulders not spinose-serrate. MNIACEAE Orthomniopsis japonica Broth. Ofv. Finska Vet.-Soc. Forh. 49 (10): 1. 1905-06. Luzon: Bontoc Subprov., June-July 1911, Father M. Vanoverbergh, 1282. Distribution: Assam, New Guinea, Japan. Through the kindness of Dr. Quisumbing I have seen part of the above collection which is unquestionably this species. For some reason it was not included in the material sent from the Bureau of Science as a basis for the study of the local moss flora. BARTRAMIACEAE Philonotis evanidinervis Fleisch. Laubmoosfl. Java 2: 412. 1902-04. Mindanao: Lanao Prov., vicinity of Dansalan, alt. 700-800 m., on rock, edge of stream, secondary forest toward Remain, 28 October 1938, A. Lynn Zwickey, 539. Ss New to the Philippine Islands. Distribution: Java. An interesting extension of this localized species previously known only from several localities in West Java. RHACOPILACEAE Rhacopilum magnirete Bartr. sp. nov. Fic. 11-14, Sat robustum, caespites late extensi, lutescentes, haud nitidi. Caules intertexti, fere ad apicem dense rufo-tomentosi, irregulariter ramosi. Folia late ovata, acuta, breviter cuspidata, 2 mm. longa, 1 mm. lata. sicca fortiter inflexa, humida late patentia; margines plani, supra medium irregulariter denticulati; costa tenuis, superne valde angustiora, sensim in cuspidem attenuata, saepe infra apicem dissoluta; cellulae superiores majusculae, pellucidae, laeves, hexagonae, 22-25 y. longae, parietibus firmis, basin versus paullo majores, internae longiores. Amphigastria multo minora, ovato-lanceolata, sensim acuminata, costa longe excurrente. Caetera ignota. Mindanao: Lanao Prov., Saguiran Mountain, on tree trunk, 26 Feb. to 30 March 1941, Liborio E. Ebalo, 1124. This species is possibly nearest R. Schmidi (C. M.) Jaeg. but the densely tomentose stems, more coarsely toothed leaves, the slender costa ending in the shorter point or even disappearing below the apex, together with the larger pellucid and distinct upper leaf cells are distinctive fea- tures. From R. spectabile Reinw. & Hsch. it will be separated by the less robust habit, larger lamina cells, less coarsely toothed leaf margins and the absence of the area of lax juxtacostal cells in the leaf base. THUIDIACEAE Herpetineurum toccoae (Sull. & Lesq.) Card. Beihefte Bot. Centralb. 19 (2): 127. 1905. Anomodon toccoae Sull. & Lesq. Musc. Bor, Am. (Ed. 1) no. 240, 1856. a BARTRAM: PHILIPPINE MosseEs Serre | OE: Luzon: Malinao, Atimanan, Tayabas Prov., on ground, 25 Oct. 1927, José V. Santos, 862. New to Philippine Islands. Distribution: Africa, Asia, Ceylon, Sumatra, Java, Celebes, New Cale- donia, Formosa, South America, Guatemala, Mexico, Southern United States, It is not surprising to find this almost cosmopolitan species in the Philippines. Although not common anywhere it seems to occur spe- radically in temperate and tropical regions throughout the world. PTEROBRYACEAE Endotrichella nematosa Bartr. sp. nov. FIG. 15-17. Robusta,. flavescens, nitida. Caules secundarii ad 5 cm. alti, complanati, densi- folii, circa 8 mm. lati, superne saepe attenuati et prope apicem propagula numerosa aurantiaca filiformia articulata instructi. Folia 5 mm. longa, 1.5 mm. lata, oblongo- lanceolata, ecostata, plicata, parum auriculata, in acumen cuspidatum raptim an- gustata; margines infra recurvi, superne distanter fortiter serrati, cellulae superiores anguste rhomboideae, perincrassatae, basilares lineares, infimae fuscae, fortiter porosae, alares oblongae, laxae, numerosae. Perichaetium parvum; theca exserta, leniter inclinata, deoperculata breviter oblonga, 1.8 mm. longa, 1 mm. lata; seta circa 2 mm. longa. Caetera ignota. Panay: Mt. Bulilao, Capiz Prov., June 1919, A. Martelino & G. Edano, 35812a. Caroline Islands: Ponape Island, Kuporujo, grows on top of the mountain on epiphytic orchid, 13 March 1936, Masahiko Takamatsu, Herb. B. P. Bishop Museum, 66la (TYPE). An outstanding species in the caudate tips of the secondary stems which are abundantly felted with erect, filiform, orange-brown, articulated brood filaments as in some of the species of Cyathophorella. Garovaglia Zwickeyi Bart. sp. nov. FIG. 18-20. Rigida, aureo-viridis, nitidula. Caules circa 4 cm. alti, sat gracili. Folia dense conferta, erecta, arcte appressa, late orbiculato-ovata, haud undulata, fortiter plicata, breviter cuspidata, 2.5 mm. longa, 2 mm. lata; margines plani, leniter denticulati; costae nullae; cellulae breviter rhomboideae, parietibus firmis, alares rectangulares, numerosae. Perichaetium Jonge exsertum, folia perichaetialia abrupte breviter cuspidata, superne erosa, theca immersa. Caetera ignota. Mindanao: Lanao Prov., vicinity of Olangu near Momungan, alt. 400— 500 m., on trunk of tree, 10 November 1938, A. Lynn Zwickey, 700. A very distinct species in the short, broad, closely appressed leaves giving the rather slender, subterete stems a catenulate appearance due to the reflexed upper leaf margins. It may be compared in a broad way with G. rigida Dix. of Siam but is widely different in the much shorter and relatively broader erect appressed leaves with the margins weakly denticulate. 3 NECKERACEAE : Neckeropsis nitidula (Mitt.) Fleisch. Laubmoosfl. Java 3: 882. 1906-08. Luzon: Tayabas Prov., Danlagan, Guinayagan, on tree branch, 19 Oct. 1936, José V. Santos, 482. vi fe neh 508 Fartowia, VoL. 1, 1944 New to Philippine Islands. Distribution: China, Japan, Formosa, Tonkin. Notwithstanding my remarks on page 245 of the “Mosses of the Philip- pines” I am happy to definitely record the presence of this species in the local area. Pinnatella ambigua (Bryol. Jav.) Fleisch. Mindoro: Pandan Mountain, Bulalacao, on stone, 6-11 Nov. 1939, Liborio E. Ebalo, 239. Distribution: Bhota, Burma, Sumatra, Java. Although the plates in the “Bryologia Javanica” representing P. ambigua and P. Kuhliana respectively show clearly the distinctive difference in the shape of the stem leaves, unfortunately I failed to grasp this distinc- tion when studying the Philippine collections and as a result confused the two species. P. ambigua is at once distinguished by the stem leaves grad- ually narrowed from an ovate base to a relatively slender acuminate point in bold contrast to the oblong-ovate abruptly short-pointed leaves of P. Kuhliana. P. ambigua is apparently rare in the Philippines. Fleischer cites a col- lection from Luzon in the Laubmoosfl. Java, p. 919 but the only speci- men I have seen is the one cited above. Pinnatella Kuhliana (Lac.) Fleisch. Hedwigia 45: 80. 1906. New to the Philippine Islands. Distribution: Sumatra, Java, Ceram, Celebes, New Guinea, Fiji, Samoa, Tahiti. As explained above the description, figures and collections given under the heading of P. ambigua in the “Mosses of the Philippines” should all be referred to Pinnatella Kuhliana. This species does not seem to have been recorded from the Philippines but a careful check of all the avail- able specimens shows that it is widely distributed throughout the archi- pelago and decidedly more frequent than P. ambigua. Pinnatella mucronata (Lac.) Fleisch. Hedwigia 45: 50. 1906. Palawan: Tagbarungis Mt., vicinity of Puerto Princesa, on tree trunk, 14 February 1940, Liborio E. Ebalo, 489. New to the Philippine Islands. Distribution: Sumatra, Java, Borneo, Ceram, Celebes, New Guinea, Samoa. HOOKERIACEAE Daltonia armata Bartr. sp. nov. FIG. 21-24, Pallide viridis, nitidula. Caulis ad 1 cm, altus. Folia 3.5 mm. longa, 1 mm. lata, oblongo-ovata, longe piliformiter aristata, erecto-patentia, sicca leniter contorta, ubique angustissime limbata, integerrima; costa tenuissima, vix dimidiam partem folii attingens; cellulae superiores elongate hexagonae, 40-60 x 10-16 u, parietibus tenuiusculis, basilares lineares. Seta 3 mm. longa, fere ad basin argute tuberculosa; theca erecta, deoperculata 1.2 mm. longa; calyptra fimbriata. BarTRAM: PHILIPPINE MossEs : 509 Mindanao: Lanao Prov., vicinity of Dansalan, alt. 700-800 m., on culm of climbing bamboo, Sacred Mountain, 3 Nov. 1938, A. Lynn Zwickey, 638. The leaves of D. armata are similar in shape to those of D. aristifolia Ren. & Card. of Java but the resemblance ends here. In D. armata the upper lamina cells are oval-hexagonal up to 60 » long and the setae are densely armed throughout with high tubercular-like spines up to 25 » high. PLAGIOTHECIACEAE Plagiothecium neckeroideum Br. & Schp. var. javense Fleisch. Laubmoosfl. Java 4: 1168. 1920. Luzon: Mountain Prov., Mt. Pawai, Benguet, on ground, 27 March 1938, | José V. Santos, 1035. Distribution: Sikkim, Java. In this collection the setae are 3-4 cm. long, thus agreeing enh the description of Fleischer’s var. javense. In the absence of any other well marked differences it seems probable that the variety is hardly more than a robust. form of the species. SEMATOPHYLLACEAE Heterophyllum Santosii Bartr. sp. nov. FIG. 25-28. Autoicum, pallide aureum, nitidum, Caulis prostratus, elongatus, regulariter pin- natim ramosus, ad 9 cm. longus, ramis vix 1 cm. longis, leniter flexuosis, apice fal- cato. Folia caulina falcata, 2.5-3 mm. longa, e basi late oblonga raptim lineari- lanceolata, superne spinoso-serrata; marginibus inferne integris, anguste revolutis; costa bina, brevissima; cellulae superiores lineares, parietibus, hyalinis, firmis, basilares longiores, angustiores, infimae laxiores, fuscae, valde porosae, alares num- erosae, oblongae, hyalinae vel fuscae, haud vesiculosae. Folia ramea minora, breviora, angustiora, sensim acuminata, argute serrata, apicem versus saepe rugoso-undulata. Seta 37 mm. longa, rubella, superne tenuissima; theca suberecta, deoperculata 2.5 mm. longa. Caetera ignota. Luzon: Mountain Prov., Mt. Data, Benguet, on tree trunk, 26 March | 1938. José V. Santos, 957. A splendid species, possibly allied to H. brachycarpum (Mitt.) Fleisch. of Japan but widely different in the elongated stems regularly and closely pinnate forming narrow fronds when moist. The unusually long seta shown by the only available sporophyte is also a distinctive feature. The genus adds a new and interesting element to the local flora from a region which has been carefully worked over by many previous collectors, so Mr. Santos may well be proud of his discovery. Glossadelphus similans (Bryol. Jav.) Fleisch. E. & P. Nat. Pflanzenfam, Musci p. 1093. 1908. Luzon: Mountain Prov., Mt. Pawai, Benguet, on dead log, 27 March 1938, José V. Santos, 1027. New to Philippine Islands. Distribution: Java. 510 FarLowlA, VoL. 1, 1944 Glossadelphus hermaphroditus Fleisch. Laubmoosfl. Java 4: 1359. 1920. Mindanao: Lanao Prov., vicinity of Guassi-Dansalan Road, alt. 700- 800 m., on boulder in stream bed near Togaya, 2 Nov. 1938, A. Lynn Zwickey, 618. New to Philippine Islands. Distribution: Java. Pseudohypnella verrucosa (Doz. & Molk.) Fleisch. Laubmoosfl. Java 4: 1368. 1921, Mindanao: Lanao Prov., Mt. Makaturing, alt. 800-1000 m., on rotting wood, 22 Oct. 1938, A. Lynn Zwickey, 489. New to Philippine Islands. Distribution: Ceylon, Java, Banka. POLYTRICHACEAE Pogonatum macrophylloides Broth. Mitt. Inst. allgem. Bot. 7 (2): 140. 1928. Luzon: Mountain Prov., Mt. Pawai, Benguet, on ground, 27 March 1938, José V. Santos, 1026. New to Philippine Islands. Distribution: Borneo. BusHKILL, Pike Co. PENNSYLVANIA ah 5. Te oe: ee ee ae, 2a. “4 ze > y pial 4 ¥ 4 be tae eel ae ae eee is td 4 Sera e ‘ ‘ x 512, FarLtowlA, Vo. 1, 1944 EXPLANATION OF PLATE Fig. 1-4. Fissidens Zwickeyi Bartr. 1. Plants « 1. 2. Leaf x 20, 3. Apex of leaf « 200. 4. Capsule x 7. Fig. 5-10. Calymperes Ebaloi Bartr. 2: Plant x 1, 6. Leaf x 4. 7. Apex of leaf x 50. 8. Leaf base x 20. 9, Upper cells and margin x 200. 10. Part of cross section from upper half of leaf x 200. Fig. 11-14. Rhacopilum magnirete Bartr. ll. Part of plant x 1. 12. Leaf x 15. 13. Apex of leaf x 40. 14. Upper leaf cells and margin x 200. Fig. 15-17. Endotrichella nematosa Bartr. 15. Plant x 1. 16. Part of propagulum x 200. 17, Leaf < 7. ‘ Fig. 18-20. Garovaglia Zwickeyi Bartr. 18. Plant x 1. 19. Leaf « 16. 20. Upper leaf cells and margin x 200. Fig. 21-24. Daltonia armata Bartr. 2): Plant < 7, 22. Capsule x 9, 23, Leaf x 10. 24. Upper leaf cells and margin x 200. Fig. 25-28. Heterophyllum Santosii Bartr. 25. Plant x \%. 26. Stem leaf « 13. 27. Branch leaf « 13. : 28. Basal angle of leaf x 80. Figures 1-28 ak icin ac ia 8 Ui tik i ag i cay % Ie ga gm gta! se = pat ‘ a Gen er a ee Fa } 4 i 2 an? ‘ i" ae a am ee = x " . 7 : ; ; , , Taga? ta . F eo yee ‘i f i j oer dean ne gsi cil : 4 7 : : : fla | 7 i : i a A a et as ee : et ata ©: E a ee ee het 1(4) : 515-523 FARLOWIA July, 1944 ON TWO NEW SPECIES OF MEGACEROS WITH NOTES ON M. ARACHNOIDEUS, M. DENTICULATUS, M. GIGANTEUS, AND M. GRANDIS L. P. Kanna For the material which forms the basis of this note, the writer is in- debted to Mrs. E. A. Hodgson, Wairoa, P. O. Hawkes Bay, New Zealand, to whom he wishes to express his thanks. Thanks are also due to Dr. Eduardo Quisumbing, Curator, Philippine National Herbarium, Manila, -and to Dr. R. E. Holttum, Director, Botanical Gardens, Singapore, for _ the loan of specimens from their herbaria. Megaceros Hodgsoniae sp. nov. ; Fics. 1-6. Planta monoica. Sectio transversa 5-6 cellis excelsitato, solida. Cellae super- ficiales 30-70 X 25-45 y.. Involucre geminatum, 5-6 mm. longitudine, 0.5-1.5 mm. latitudine, cylindricum, attenuatum. Capsula 8-15 mm. longitudine, 0.25-0.35 mm. latitudine. Sporae pallidae, verrucolosae, 30-40 Une Monoecious. In patches. Thallus up to 25 mm. long, nearly flat, smooth, divided into broad obovate lobes, more or less rounded at the apex with margin nearly entire; transverse section 5-6 cells high in the middle, without lacunae; surface cells 30-70 x 25-45 p. Involucre fre- quently geminate, 5-6 mm. long and 0.5-1.5 mm. broad, cylindrical, slightly narrowing at truncate apex. Capsule 8-15 mm. long and 0.25—0.35 mm. broad, wall cells thickened, light brown. Spores 30-40 p, pale, verrucose. On Popa Rock in creek in deep shade, Kiwi, April, 1935. From the comparative chart of the species of Megaceros Campbell 1907 appended, the following species agree with the form described in having the verrucose type of spore surface: M. arachnoideus Stephani 1916, M. denticulatus (Lehman 1857) Stephani 1916, M. fuegiensis Stephani 1923, M. martinicensis Stephani 1916. The present form differs from M. martinicensis by the shorter capsule and the larger thallus, from M. arachnoideus, M. denticulatus and M. fuegiensis by the shorter capsule and the structure of the thallus. It is therefore necessary to create a new species for which the name M. Hodgsoniae is suggested. Megaceros Zotovii sp. nov. Fics. 7-10. Planta monoica. Sectio transversa 10-12 cellis excelsitato, solida; cellae super- ficiales 30-45 x 15-25 y. Involucre solitarium, cylindricum, attenuatum, 12-16 mm. longitudine, 1.5-2 mm. latitudine. Capsula 40-60 mm. longa, 0.5 mm. lata. Sporae pallidae, verruculosae, 30-35 wu. Monoecious. In patches,:light green. Thallus up to 50 mm. in di- ameter, fan-shaped, with obovate lobes, rounded at the apex with margin finely crenate, transverse section 10-12 cells high in the middle, without lacunae; dorsal epidermal cells 30-45 x 15-25 p. Involucre 12-16 mm. long and 1.5-2 mm. broad, cylindrical, narrowing towards the nearly truncate apex. Capsule 40-60 mm. long and 0.5 mm. broad. Spores 30-35 y, pale, somewhat irregular in shape, verrucose. Elators un- branched, 0.2-0.3 mm. long. Epidermal cells of the capsule somewhat 515 eee te ee Sey ER ee 516 FarLtowl1a, Vou. 1, 1944 thickened. Cehatorawa Saddle, Tarama Mts., New Zealand, 26 Nov. 1933, V. D. Zotov. : The above described species agrees with the following in having ver- rucose spores: M. arachnoideus Stephani 1916, M. denticulatus (Lehmann 1857) Stephani 1916, M. fuegiensis Stephani 1923, M. Hodgsoniae Khanna and M. martinicensis Stephani 1916. The present form differs from M. denticulatus and M. Hodgsoniae by its longer involucre and capsule, from M. arachnoideus, M. fuegiensis and M. martinicensis by the bigger in- volucre and other characters of the thallus. Megaceros giganteus (Lehmann et Lindenberg) Stephani Fics. 11-15. Monoecious. In light green patches. Thallus up to 50 mm. in di- ameter, divided into obovate lobes, depressed in the center, margin slightly ascending-irregular; transverse section 8-10 cells high in the middle, without lacunae; surface cells 8-20 x 20-30 ». Involucre 8-15 mm. long and 1.0-1.5 mm. broad, cylindrical, slightly narrowed towards the apex. the mouth lobulate. Capsule 40-60 mm. long and 0.4—0.8 mm. broad, epidermal cells of the capsule, long, narrow, highly thickened, and some cells with brownish contents. Spores 30-35 p, pale, asperulate. Elators long and slender, sometimes 0.5 mm. in length, unbranched. Antheridia singly on prominent stalks. Water course, north of Field Flat, 8 Oct. 1933, V. D. Zotov. The above described species agrees with the following in having the same type of spore surface: M. caledonicus Stephani 1916, M. celebensis Stephani 1916, M. crassus Stephani 1916, M. endiviaefolius (Montagne 1856) Stephani 1916, M. giganteus (Lehmann et Lindenberg 1834) Stephani 1916, M. Jamesonii (Taylor 1848) Stephani 1916, M. lacerus (Nees 1844.) Stephani 1916, M. longispirus (Carrington & Pearson 1887) Stephani 1916, M. monandrus Stephani 1916, M. Novae-guineae Stephani 1916, M. Novae-zelandiae Stephani 1916, M. pallens (Stephani 1892) Stephani 1916, M. parvisporus Stephani 1916, M. solidus Stephani 19106. The present species differs from M. caledonicus, M. celebensis, M. crassus, M. lacerus, M. monandrus, M. Novae-guineae, M. pallens, and M. solidus by the larger thallus and involucre; from M. endiviaefolius by its shorter capsule; from M. Jamesonii by its smaller spore and other vegetative characters; from M. lacerus by the shape and the other char- acters of the thallus. M. giganteus, M. Novae-zelandiae and M. parvi- sporus and the above described species differ from each other in minor characters which are not sufficiently distinct to be of specific value. The writer therefore is of the opinion that these may be considered as a single species — M. giganteus (Lehman et Lindenberg 1834) Stephani 1916. _ The writer has been unable to consult Stephani’s material of these forms and is therefore unable to state whether differences exist which would supplement the already existing description. Judging from the char- acters so far given, however, he is of the opinion that there is only one species, and the other two listed are environment forms without spe- cific validity. Ly ee ee ee a Pe, eee er es ae eee ae fs ee ete a Se eee ee ee ee re bee ee eh a ee eee. oe ee ie F ERROR Scat Rete Ten ap eee AAR, Bip Ce hg tM ee Rr PU See eget QOS a ing MENS cask hy Oy aera OPS meg eR ae MRT cage ales ae ne ae Ss oe Perr nt tp 8 ob ij ns am * : ; : ¥ te KHANNA: MEGACEROS Syke Megaceros grandis (Angstrom) Stephani FIcs. 16-20. Monoecious. On rotten log, mixed with mosses. In light green patches. Thallus 12-20 mm. in diameter, fan-shaped or nearly so; slightly de- pressed in the center, the margin ascending; divided into numerous nar- row linear lobes, dorsal surface rough with several fine lamellae; surface cells 35-55 x 20-40 yp, transverse section 5 cells high in the middle, with- out lacunae. Involucre 4.5-6.5 mm. long and 0.8-1.0 mm. broad, cylin- drical, narrowing to the almost truncate apex. Capsule 12-40 mm. long - and 0.3-0.5 mm. broad, epidermal cells long, thick-walled. Spores 25-35 p, pale green, granular; elators unbranched, 0.18—-0.2 mm. long. Antheridia borne singly in the antheridial cavity. Stream side—on a log in a bush about 3000 ft. above sea level; on Tauhara, New Zealand. From the comparative chart of Megaceros Campbell, 1907, the following species agree with that described above in having the same type of spore surface: M. grandis (Angstrom 1873) Stephani 1916, M. vescoanus Stephani 1916, M. vincentianus (Lehmann et Lindenberg 1834) Ste- phani 1916. The present species can be separated from M. vincentianus by its shorter capsule and other structures of the thallus; M. grandis, M. vescoanus and the above differ from each other in minor characters which are not sufficiently distinct to be of specific value. The writer, therefore, is of the opinion that these may be considered as a single species — M. grandis (Angstrom 1873) Stephani 1916. Megaceros denticulatus (Lehmann) Stephani Fics. 21-25. Monoecious. In overlapping patches. Thallus variable, up to 55 mm. long and 40 mm. broad, with irregular branches, rounded at the apex, with margin irregular; from the ventral surface marginal buds are given off; the upper superficial cells 25-30 x 10-20 ». Involucre 5-7 mm. long and 0.8 mm. broad, narrowing towards the apex, the mouth lobulate. Capsule 16-20 mm. long and 0.5 mm. broad, brownish. Spores 35-40 p, pale yellow, verrucose. Capsule wall cells highly thickened. From the description this plant appears to be M. denticulatus (Leh- mann 1857) Stephani 1916— though the present form differs by the larger involucre and the smaller spores — but these differences within a narrow limit are not sufficiently distinct to be of specific value. Megaceros arachnoideus Stephani FI¢s. 26-29. Monoecious. In patches. Thallus 15 mm. in diameter, divided into broad obovate lobes, with margin lacerate; dorsal surface rough with several leaf-like lamellae; transverse section 10-14 cells high in the middle, without lacunae; upper superficial cells 35-45x 20-45 p. — In- volucre 6-10 mm. long and 0.6-1.2 mm. broad, narrowing towards the apex, mouth lobulate. Capsule 25-35 mm. long and 0.3—0.4 mm. broad, brownish. Spores 25-40 », somewhat irregular in shape, greenish yellow, verrucose. Elators long and slender, sometimes 0.2 mm. in length. On damp earth by creek by [Mols St.?] New Zealand, 26 Dec. 1932, kK. A. Hodgson. 518 FarLtowl14, VoL. 1, 1944 This form agrees with M. arachnoideus Stephani 1916 but shows minor differences. University OF RANGOON InpIA REFERENCES Campbell, D. H. Studies on some Javanese Anthocerotaceae, I. Ann, Bot. 21: 467-486. pl. 44-46. 1907. Stephani, F. Species Hepaticarum 5: 945-957. 1916. —_————. Species Hepaticarum 6: 424. 1923. alatifrons Steph. amoenus Steph, aneuraeformis Steph. arachnoideus Steph. caledonicus Steph. callistictus Steph. carnosus Steph, celebensis Steph. columbianus Steph. crassus Steph. cristisporus Steph. denticulatus (Lehm.) Steph. endiviaefolius (Mont.) Steph. flagellaris (Mitt.) Steph. flavens (Spruce) Suegiensis Steph. giganteus (Lehm. & Lindenb.) Steph. gracilis (Reichardt) Steph. grandis (Angstr.) Steph. guatemalensis Steph. jamaicensis Steph. Jamesonii (Tayl.) Steph. lacerus (Nees) Steph. laciniatus (Schw.) Steph. leptohymenius (Tayl.) Steph. longispirus (Carr. & Pears.) Steph. martinicensis Steph. mexicanus Steph. minarum (Nees) Steph. monandrus Steph. monospirus Steph. muriculatus Steph. Novae-guineae Steph. Novae-zelandiae Steph. Nymanii Steph. pallens (Steph.) Steph. parvisporus Steph. e salakensis Campb. schizophyllus (Gottsche) Steph. solidus Steph. Stahlii Steph. tjibodensis Campb. tosanus Steph. . ’ Vescoanus Steph. vincentianus (Lehm. & Lin- denb.) Steph. Wiemanii Steph. KHANNA: MEGACEROS APPENDIX SPECIES OF MEGACEROS Size of Invo- Cap- plant lucre_ sule mm. mm. mm. 20x3 7 30 20x3 7 30 40x 5 5 30 40x25 7 30 20x3 5 40 20 x3 3 15 30 5 50 20 x 12 8 30 15x5 12 40 20 6 40 40 15 70 50 2 20 60 x 7 7 90 40x 5 a 40 30x 5 8 60 30x 8 5 40 45 x7 10 40 20 5 40 30x10 10 30 40x 5 9 60 50x10 10 40 40x10 10 50 40 7 40 40 5 40 50x10 10 40 50 x 20 5 50 10x 4 4 40 15x4 4 40 20 5 40 30 5 20 35 7 25 30 5 45 20x3 5 40 40 5 30 30 4 35 25x5 5 40 30x 5 10 40 30 6 40 15 5 20 20x 4 6 20 30 15 80 50x15 10 90 30 i 20 40 x7 7 40 40 x 6 5 50 20x10 15 50 Spore 27 uw, small papillae in center 27 pn, umbonate 36 », umbonate 36 uw, yellow, verrucose 27 uw, Minutely asper 22 uw, muricate - 27 uw, yellow, muricate 27 uw, minutely asper 36 uw, umbonate 36 uw, asper 27 uw, yellow, papillate 45 yu, yellow, verrucose 36 yu, asper 18 nz, muricate 27 w, muricate 36 yw, verrucose 27 uw, asper 36 u, papillate 27 uw, granulate 27 uw, papillate 27 u, papillate 54 yw, asper 36 yu, asper — u, muricate 36 yu, papillate 36 », minutely asper ‘27 uw, dark, verrucose 27 yp, papillate— 27 w, yellow, papillate 27 », minutely asper 27 yw, papillate 27 uw, Muricate 27 yw, asper 27 w, asper 36 yu, papillate 27 uw, asper 18 uw, asper F 28 uw, minutely echinate 27 uw, tuberculate ! 36 uw, asper 27 wu, yellow, densely papillate 27 wu, papillate 27 uw, muriculate 18 yn, granular 27 w, granular 27 wu, umbonate 219 Distribution Brazil Guadeloupe Brazil New Zealand New Caledonia Brazil Australia Celebes Colombia Tasmania Costa Rica New Zealand Chile Fiji Peru Tierra del Fuego New Zealand Australia Tahiti Guatemala Jamaica Peru Africa Mexico New Zealand Tasmania Martinique Mexico Brazil Asia , New Caledonia New Caledonia New Guinea New Zealand New Guinea New Zealand Samoa Java Cuba Guadeloupe Java Java, Sumatra Japan Tahiti Guadeloupe, St. Vincent Brazil 520 FarLtow1a, Vou. 1, 1944. EXPLANATION OF FIGURES 1-15 In figures 1 and 11, the scale equals 1 millimeter; in all other figures the scale equals 20 yu. Megaceros Hodgsoniae. Figs. 1-6. 1. Young plant showing sporophyte. 2. Dorsal epidermal cells. 3-4, Spores. 5. A portion of an elator. 6. Capsule wall cells. Megaceros Zotovii. Figs. 7-10. 7. Dorsal epidermal cells. 8. Different views of spores. 9. Part of elator. 10. Capsule wall cells. Megaceros giganteus. Figs. 11-15. 11. Thallus showing sporophyte. 12. Dorsal epidermal cells. 13. Spore. 14, Elator. 15. Capsule wall cells. Se eg ee ae Se ee Na “Scale 521 KHANNA: MEGACEROS ) Figures 1-1 eae se Le ne ea eer ee Cae ee ee ee ee ee x ee 7, = * ih sap ee we Of a. See OG wll) Se Te, a 522 Far.towlia, VoL. 1, 1944 EXPLANATION OF FIGURES 16-29 In figures 16, 21, and 26, the scale equals 1 millimeter; in all other figures the scale equals 20 u. Megaceros grandis. Figs. 16-20. 16. A young plant. 17. Dorsal epidermal cells of the thallus. 18. Spore. 19. A portion of an elator. 20. Epidermal cells of the capsule wall. Megaceros denticulatus. Figs. 21-25. 21. A portion of the lobe of the thallus showing the irregular margin. 22, Epidermal cells of the thallus. 23. Capsule cell walls. 24. Spore. 25. Portion of an elator. Megaceros arachnoideus. Figs. 26-29. 26. Young plant. 27. Dorsal epidermal cells of the thallus. 28. Spores. 29. A portion of an elator. KHANNA: MEGACEROS Ficures 16-29 See ee ee gee ES eS SS he OE, se eM RE a EN ee ee ke ee ae ee re a erate ea mee oe : ere ' Ss ene 1(4): 525-568 FARLOWIA July, 1944 STUDIES ON POLYSTICTUS CIRCINATUS AND ITS RELATION TO BUTT-ROT OF SPRUCE Rocer GossE.Lin! During the writer’s study of the butt-rots of spruces in the Province of (Quebec, it was found that the most important fungi causing butt-rots are Porta subacida, Polyporus balsameus, Polyporus Schweinitzii, and Poly- porus circinatus. This last was chosen for study because of the irregular occurrence of the species and of its importance locally in causing ex- cessive windfall. It soon became evident that what had been determined as P. circinatus actually included two distinct fungi. One was characterized by its bright- er color and by the fact that it occurred chiefly on the ground; the other was characterized by the darker color of the sporophores and by the fact that it occurred mostly on the trunks of the trees. These differences be- tween the two forms which were so obvious in the field, led to the inves- tigations of the taxonomy. It was found that the two organisms could be separated microscopically by the character of the setae present in the hymenium and this led to their determination as Polyporus tomentosus Fr. and Polyporus circinatus Fr. From a review of the literature it appears that there has been consid- erable confusion as to the identity of these two forms. This confusion has arisen since Fries described the two species, P. tomentosus in 1821 and P. circinatus in 1848, and it came from the fact that he described P. tomen- tosus as having a homogeneous context, yet according to specimens iden- tified by Fries, both P. tomentosus and P. circinatus have a duplex context. The only difference, excepting the color and the shape, are the straight setae for the former and the predominantly curved setae for the latter. Peck (1878) described a new species from New York, which he called Polyporus dualis. His species was sessile or short, lateral-stemmed and of rather dark color. Later, Ellis and Everhart (1889) examined speci- mens from Peck and found the curved setae characteristic of P. circinatus Fr. Cooke (1886) and Saccardo (1888) have given P. dualis Pk. as a synonym of Polystictus circinatus (Fr.) Cke. So it is fairly safe to say that Peck’s type specimen is P. circinatus Fr. In 1882, Karsten transferred Polyporus tomentosus to Polystictus. Later (1889) he segregated the new genus Onnia from Polystictus, placing em- phasis upon the setal character of the hymenium. He distinguished O. tomentosus from QO. circinatus by the homogeneous context of the former. Ellis and Everhart (1889) examined Karsten’s specimen later and found straight setae for the plant he designated as Onnia tomentosus. Although Ellis and Everhart’s illustration (1889) did not make this point 1 Thesis submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at Harvard University. Contribution from the Laboratories of Cryptogamic Botany and the Farlow Her- barium, Harvard University, no. 225. 529 526 FartowlA, Vou. 1, 1944 clear, Karsten’s specimens show that there is a duplex context, hence his Onnia tomentosus is the same as P. tomentosus Fr. Little is known about his O. circinatus. While Karsten was creating the genus Onnia, Ellis and Everhart (1889) created the genus Mucronoporus based on Polyporus circinatus. They described straight setae for M. tomentosus and curved ones for M. cir- cinatus. Subsequent mycologists have considered these species synony- mous with the two Friesian ones. | In 1900, Patouillard made a new genus Xanthochrous based on P. cir- cinatus. He described the setae as being straight. Apparently his X. circinatus is P. tomentosus Fr. Murrill (1904) refers to Fries’ authentic specimens, stating that “the type of Polyporus tomentosus at Upsala corresponds in all respects with the plants found in America, having the same kind of spines and a dual context.” Since Nannfeldt has reported in a personal letter to Haddow (1941) that there are no type specimens of either of Fries’ species at Upsala, it is difficult to understand to which specimen Murrill refers. It is worth observing, however, that he attributed to his Friesian specimen of P. tomentosus a double context. Murrill (1916) comprises under the single species Coltricia tomentosa both P. tomentosus and P. circinatus of Fries. There apparently was much confusion in Lloyd’s mind in regard to the status of these two species. In 1908, he described P. circinatus Fr. as having straight setae, as in P. tomentosus Fr. Later in 1912, he stated that P. tomentosus Fr. was unknown from the United States and ques- tioned whether P. circinatus Fr. was really distinct from P. tomentosus Fr., and finally suggested that “there are three (stipitate) forms of Poly- porus tomentosus Fr.: first (typical), with thin context, straight setae; second (circinatus, American), thick context, straight setae; third (cir- cinatus, European and American), curved setae.” Lloyd (1908), on account of the resemblance of P. dualis Pk. to Fries’ figure of P. triqueter, suspected it to be P. triqueter of Europe. Bresadola, (1920) following Lloyd’s opinion, considered P. dualis Pk. to be a synonym of P. triqueter Fr. Fries described P. triqueter in 1838, while P. circinatus was described only in 1848. He did not mention any rela- tionship between the two species, hence P. triqueter Fr., if different from P. dualis Pk. as seems likely, cannot be used as a synonym for P. circinatus and there is little need to consider it in this discussion. | Neuman (1914) reported P. tomentosus from Wisconsin but gave no description of the microscopic characters. He considered P. dualis as a synonym of P. tomentosus and for that reason the exactitude of his iden- tification may be questioned. Lloyd (1920) described a new species, Polyporus peakensis Lloyd, from a single specimen collected by Hedgcock at Pikes Peak in Colorado. Later, Shope (1931) stated it was conspecific with P. circinatus Fr. GossELIN: STUDIES ON POLYSTICTUS CIRCINATUS 527 Sattory and Maire (1922) believed that P. circinatus is only a variety of P. tomentosus, a point of view shared by Haddow (1941). Konrad and Maublanc (1926) under Fomes (Xanthochrous) circinatus furnished a picture of curved setae and described it as straight or curved. We previously mentioned the same thing for the wood-loving forms we found in the field. It should be P. circinatus Fr. Bourdot and Galzin (1927) made the same observations on what they called P. circinatus ae Shope (1931), Overholts (1933) and Lowe (1934) followed Lloyd’s opinion and do not recognize P. tomentosus in the American Flora. Furthermore, Overholts states that all American forms should be referred under the name P. circinatus. All three described the typical form as having straight setae because they have been led astray by the statement of Fries that the species was characterized by the homogeneous context of the fruiting bodies. Concerning P. dualis Pk., Shope considered it the same as P. circinatus, while Overholts gave a varietal rank because of the curved setae which in the typical form are straight. Lowe considered it a good species. Jérstad and Juul (1938) worked out the taxonomy of the species with Norwegian material. They recognized two varieties to P. tomentosus Pr.: P. tomentosus var. americanus and P. tomentosus var. circinatus. They made the variety americanus because of Lloyd’s statement (1912) that P. tomentosus Fr. does not exist in the American flora, but only a related form. The variety circinatus was suggested for P. circinatus Fr. because they said there is no definite line of demarcation between Fries’ species. They make no mention of Sartory and Maire who made the same va- riety in 1922. Haddow, in 1941, based all his discussion on specimens of the Curtis Herbarium identified and labelled by Fries himself. He recognized the duplex context for both P. tomentosus and P. circinatus; the former had straight setae and the latter curved ones. Haddow did not agree that the curved setae character was sufficient to make a new species as did Fries, thus he called it P. tomentosus var. circinatus. As a resumé, the following table shows the real position of each specimen. Author Polyporus tomentosus Fr. Polyporus circinatus Fr. - Fries (1851) Polystictus tomentosus Peck (1878) P. dualis Cooke (1886) Polystictus tomentosus Polystictus circinatus Saccardo (1888) Polystictus tomentosus Polystictus circinatus Karsten (1889) . Onnia tomentosus Onnia circinatus? Ellis & Everhart (1889) Mucronoporus tomentosus Mucronoporus circinatus Hennings (1898) Polystictus tomentosus Polystictus circinatus Patouillard (1900) Xanthochrous circinatus Murrill (1904) Coltricia tomentosa Coltricia tomentosa ~ Lloyd (1908) (1912) Polyporus tomentosus Polyporus dualis Polyporus circinatus Ames (1913) ; Polystictus tomentosus Polystictus circinatus Neuman (1914) Polyporus tomentosus Polyporus dualis 528 FarLowia, Voi. 1, 1944 Author Polyporus tomentosus Fr. Polyporus circinatus Fr. Bresadola (1920) Polyporus dualis Polyporus triqueter Sartory & Maire (1922) Polyporus tomentosus P. tomentosus var. circinatus Konrad & Maublanc (1926) Fomes (Xanthochrous) circinatus Bourdot & Galzin (1927) Polyporus circinatus Killerman (1928) Polystictus tomentosus Polystictus circinatus var. triqueter Shope (1931) Polyporus circinatus Polyporus dualis Overholts (1933) Polyporus circinatus Polyporus circinatus var. dualis Lowe (1934)- Polyporus circinatus Polyporus dualis Jérstad & Juul (1938) Polyporus tomentosus P. tomentosus var. circinatus P. tomentosus var. americanus Haddow (1941) Polyporus tomentosus P. tomentosus var. circinatus It is clear, from what has been written above, that the tangled taxonomy of the two types would not have resulted had it not been for Fries’ early statement that Polyporus tomentosus possessed a homogeneous context, whereas a study of his specimens by the writer and others, has shown that the context of the sporophore is duplex in both types, the first of which should be recognized as P. tomentosus (with straight setae) and the second P. circinatus (with curved or straight setae in the same fructi- fication). Assuming for the time being that the two species belong in the genus Polyporus, it appears to the writer that Fries was correct in making these separations, since, as it has been stated above, the two species are obviously different when seen together in the field (Plate I, fig. 1, 2). It is for this reason that the writer considers it desirable to follow Fries in recognizing these species as distinct rather than to follow Sartory and Maire, and Haddow in reducing P. circinatus to varietal rank under P. tomentosus. The generic position of these two species would be greatly simplified if they were kept in the genus Polyporus as has been done by many, al- though Murrill (1916) and Donk (1933) and others have tended to divide the genus into smaller units and probably rightly so. On the basis of priority, Fries’ genus Polystictus (1851) must be accepted for this and related species, since P. tomentosus was specifically mentioned as an ex- ample of the genus and therefore can be taken to represent the type of the genus. For this reason, it seems best to consider that Polyporus tomen- tosus and Polyporus circinatus should be known as Polystictus tomentosus (Fr.) Fr. and Polystictus circinatus (Fr.) Cke. DISTRIBUTION Since they have first been described by Fries, P. tomentosus and P. circinatus have been found abundantly in Asia, Europe and America. GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS 529 Because frequently they have been misnamed, the writer has prepared from the literature a list of localities and has tried to indicate the dis- tribution under P. tomentosus Fr. and P. circinatus Fr. Where there is any question of the accuracy of the determination, the author name is ° followed by a (?). Nova Scotia New Brunswick Locality Polystictus tomentosus Polystictus circinatus (Fr.) Fr. (Fr.) Cke. Africa Lloyd(?) (1920) Austria Lloyd (1908) China Teng (1932) Canada Prince Ed. Is. Ell. & Evh. (1889) Ontario Lloyd (1908), Faull (1922) Manitoba & Bisby (1938) Saskatchewan McKay(?) (1904, 1908) Hay (1903, 1905) Wehmeyer (1940) Hay (1903, 1905) Germany All. & Schn. (1886), Hen- Hennings (1898) Murrill nings (1898), Murrill (1904) (1904) Japan Shirai(?) (1927 Sweden Fries (1863), Lloyd (1908, Fries (1863), Lloyd (1908), United States California Colorado Dist. of Columbia Florida Idaho (northern) 1912), Murrill (1904, 1908) Maurrill (1908) Lloyd (1911, 1915) Kauffman (1921), Lloyd (1908, 1916, 1920), Murrill (1904), Seaver & Shope (1936), Shope (1931) Lloyd (1922) Lloyd (1920) Boyce (1938), Hubert (1929, Haskell & Wood(?) (1930), 1931), Martin (1929) Hubert (1931) Murrill (1915) Towa Wolf (1931) Maine Lloyd (1923), Murrill (1904) White(?) (1902), Ricker (?) (1902) Maryland Lloyd(?) (1908) Massachusetts Lloyd (1907, 1908, 1911, 1912, 1913, 1914 1915, 1921), Murrill (1904) Michigan Kauffman (1911), Lloyd (1912), Longyear (1904), Overholts (1916), Povah (1935) Minnesota Hubert (1924, 1931), Lloyd Haskell & Wood (1930) (1920, 1922) Martin (?) (1925) Montana Hubert (1931) New Hampshire Murrill (1904) New Jersey Murrill (1904) Ellis(?) (1890), Ell. & Everh.(?) (1889) New York Burnham & Latham (1914, Peck (1878, 1894) 1924), Lloyd (1913, 1915, 1916, 1920, 1923), Murrill (1904), Peck (1869, 1880, 1893, 1899) , Saccardo (1888) ‘eteahaeil oe he rele iL ae 530 Fartowia, Vou. 1, 1944 Locality Polystictus tomentosus Polystictus circinatus (Fr.) Fr. (Fr.) Cke. North Carolina Curtis (1867), Lloyd (1914) Atkinson & Schrenk(?) (1893) Pennsylvania Overholts (1933) Rhode Island Bennett (1888) Tennessee (east) . Hesler (1929) Vermont Lloyd (1908, 1911, 1915), Murrill (1904) Washington Hubert (1931), Lloyd (1916) West Virginia Lloyd (1908, 1920), Mur- rill (1904) _ Wisconsin Dodge (1914), Neuman (1914), Overholts (1916) Wyoming Seaver & Shope (1935, 1936) To complete this list, the writer has plotted on a map the North Amer- ican localities of the specimens of P. circinatus (Fr.) Cke. kept in the Farlow Herbarium. It seems that this species reaches its southern limit at the 36th parallel of latitude (Plate I, fig. 3). It is probably limited by the temperature and if so it could be expected farther south in the mountains. History AS A PARASITE Many writers have discussed the taxonomy of the two Friesian species since 1878, and still more mycologists have collected them in the forests, but very few, either in the Eastern or Western Hemisphere, have worked out the parasitism of these species. In the Eastern Hemisphere, Jgrstad and Juul (1938), after a discussion of the taxonomy of the species, questioned whether P. tomentosus Fr. has any parasitic tendencies and mentioned the fact that in Sweden it has been found on a root of spruce which did not present any rot. However, the ability of P. circinatus to cause rot is not doubtful and they gave the general characters of the rot as it occurs in Sweden and mentioned the similarity that this. particular one has with that caused by Fomes pini. The first in North America to report this species as a parasite was Dr. Faull (1922) who found that, in Ontario, it was causing a root- and butt-rot in spruce, hemlock and eastern white pine. Giving the descrip- tion and habits of the sporophores, he identified them as being P. tomen- tosus F'r., but his herbarium specimens for that region show that P. cir- cinatus Fr. was also abundant. He mentioned the fact that the infection was very much localized and further added that in the Otter District of Ontario a high percentage of white pine is affected. Hubert (1931) reported that P. circinatus attacked roots and butt sec- tions of western white pines in northern Idaho, but considered it to be a wound fungus and commonly associated with fire scars. He made a comparative study of P. circinatus and Trametes Pini and clearly de- scribed the similarities and differences between the two rots based on the macroscopic and microscopic characters of the two different decays, and the differences in the cultural characters of the two pathogens. GOSSELIN: STUDIES ON PoLysTICTUS CIRCINATUS 531 Christensen (1940) published a brief note on P. circinatus, giving the . cultural characteristics and a description of the decay and of the sporo- phores and mentioning the importance of this disease in northern Min- nesota as the cause of windthrow and premature death of the trees. From this brief resumé of the literature it is apparent that although we have a certain body of facts in regard to the type of decay, the micro- scopical and macroscopical characters of the decayed wood, and of the cultural characters of the fungus, we have very little knowledge concern- ig the parasitic nature of the fungus, of geographical or other factors influencing infection, and little information about the biology of Polystic- tus circinatus. For these reasons and since the fungus seems to be of local importance because of its effect on the spruce forests of the Province of Quebec, the writer decided to make more intensive studies of the fungus in an endeavor to discover further facts bearing on the biology of the fungus and its relation to the host. At the same time, it was hoped to explain the relation of other butt-rot fungi to their hosts. STUDIES ON THE HOST-PARASITE RELATIONS FIELD OBSERVATIONS In undertaking this study of Polystictus circinatus, an intensive survey was made to determine the occurrence of this speciés in the Province of Quebec, especially along the north shore of the St. Lawrence River and in the Gaspé Peninsula as well as in the Laurentide National Park area. On the Gaspé Peninsula, and more especially along the northern side, the fungus appears to be of rather frequent occurrence, whereas on the south side it is somewhat scarcer. On the northern side of the Gaspé Peninsula in the vicinity of Mt. Albert, infection? was locally very high, (100%), in the York River area infection amounted to 95%, and at Lake Ste. Anne approximately 50%. On the north shore of the St. Lawrence River, near Lake Walker, the infection in the valley was about 20% and on the mountain slope up to 80-90%. On the other hand, in the Lauren- tide National Park, no evidence of infection could be discovered, even though that region is intermediate between Ontario, where Faull (1922) states that the fungus is very abundant, and the North Shore of the St. Lawrence. The high degree of rotting of spruces by Polystictus circinatus indicates that this fungus may be of considerable importance in the Province of Quebec, and the gap in the distribution of the species represented by the complete or nearly complete absence of the fungus in the Laurentide Na- tional Park indicates that there may possibly be some ecological factor involved which not only would furnish information on the distribution of P. circinatus but might also help to explain host-parasite relations in the heavily rotted areas. * As shown by the obvious symptoms of butt-rot at stump section. 552 FartowiA, VoL. 1, 1944 For the understanding of the behavior of the disease, it would be of interest to know if this disease occurs on trees weakened by age, crowding, competition with other species, by storm or insect damages. Christensen (1940), according to observations made in Minnesota, states that P. circinatus can infect trees as young as fifteen to twenty-five years old. From the writer’s observations it can be stated that trees of forty years age presented visible decay at the stump section. Since they must have been infected earlier, it is probable that the age of infection given by Christensen is near the truth. According to the same observations, Christensen stated that “The fungus does not merely accompany stagnation and senescence but, if present at all, is likely to be one of the primary causes of such conditions.” The writer agrees with Christensen that it sometimes happens that P. circinatus will kill the tree, but he does not think that it will always bring a premature senescence of the trees. From the data obtained from the Gaspé and North Shore regions, it is shown that the average age for the infected trees in Gaspé is 90 years, while it is 130 years for the North Shore region. The average age for sound trees is respectively 85 and 114 years. Thus, it seems significant. that, on the whole, infection by P. circinatus appears to delay senescence. The effect of crowding does not seem to have much influence on the occurrence of the disease. The disease was found where tree cover was very dense or loose, fruiting bodies having been found in places where the canopy was so dense that herbaceous plants could not grow, as well as in places where the canopy was sufficiently open to permit the growth of these covering plants. Hence, crowding is not a determining factor for the disease. Infection occurred either in pure stands of spruces or in mixed stands of spruce and fir. But in these mixed stands, infection has rarely been found when the percentage of firs was higher than 60%. This might be explained by the fact that the fir grows where soil is richer and where infection of spruce is relatively infrequent. Incidentally, P. circinatus was not found in stands of mixed soft and hardwoods, although P. tomen- tosus occurred in such stands. Again this may be explained by the fact that infection is less frequent in richer soils. Competition between host species does not seem to favor any tendency for infection. It can be stated, rather, that infection is most commonly found in pure stands where conditions for the growth of the trees are less favorable. In the Gaspé region and on the north shore of the St. Lawrence River, observations were made on the occurrence of the butt-rot; there was no correlation between spruce sawfly epidemic, which was very severe, and the rotting of spruce by P. circinatus. At Mount Albert and at Lake Ste. Anne the black spruce was rotted, whether damaged or undamaged by the spruce sawfly, and the disease was found independently on both of them. Furthermore, in places like Petite Riviére-a-Marte and York River where there was up to 95% and 100% infection by the disease, the damage by the spruce sawfly was nonexistent. Except for the spruce sawfly epidemic GOSSELIN: STUDIES ON POLysTICTUS CIRCINATUS 533 we have not seen, during our work, any sort of insect attack. It is there- fore fairly evident that the rot caused by P. circinatus is not related to, or a result of, insect attacks. Storms which break branches or make scars on the trunks of trees do not appear to have any influence on infection by P. circinatus, since infec- tion takes place through underground parts. Nevertheless, a large num- ber of forest soils in Quebec being thin, the rooting of trees is superficial and hence, under the action of the wind the trees oscillate and some small roots may break off and infection by P. circinatus may take place. Yet careful studies did not furnish evidence of increased infection. In addi- tion, it may be remarked that in no case did entrance of the fungus into the wood take place through other parts than those underground, or which were at least in contact with the duff. Hubert (1931) states that this disease is often associated with fire scars but we believe that those scars had to be in contact with the ground. In places where infection was up to 85% and 100%, we have not seen a single case in which infection oc- curred at other places than those mentioned above. However, from inocu- lation experiments carried out in Duchesnay, it has been found that if the mycelium comes in contact with a scar on the trunk, it will invade the wood cells. Thus, it can be suspected that under natural conditions, the spores of the fungus must germinate in the humus. In our North Shore experiments we have organized some kind of rudi- mentary meteorological stations in order to determine if temperatures, maximum and minimum, the quantity of rainfall, the relative humidity of the air, and the evaporation rate would be reflected in the number of in- fected and non-infected stands. The different measurements obtained at the two places had no significance, and none of the meteorological factors could be correlated with infection or non-infection of the stand. As a rule we can state that the fungus invades very young trees and is seen in old ones, but the occurrence of this disease is not correlated with any crowding, competition or damage of any sort, or with local meteoro- logical factors. EFFECTS OF TEMPERATURE AND HUMIDITY ON SPOROPHORE PRODUCTION Bearing in mind that the practical aspect of the subject is the control of the disease, field observations were made to determine the factors bring- ing about the fructification of the fungus, since the control of a disease is sometimes attempted by the removal of the fruiting bodies. During the summer of 1941, which the writer spent in Shelter Bay on the North Shore of the St. Lawrence River, it was only on the 15th of August that he found sporophores of Polystictus circinatus. As they are produced at about the same time of the year, he has tried to correlate the maximum and minimum temperature and the relative humidity of the air with the appearance of sporophores. For this purpose the following fig- ures of temperature and humidity have been brought together in table I. 534 FartowisA, Vou. 1, 1944 Table I Givine THE Maximum AND MINIMUM TEMPERATURE AND THE RELATIVE HUMIDITY or THE Arr IN SHELTER Bay, FROM JULY THE 15TH TO AUGuUsT THE 21sT 1941 Temperature Temperature Date Max. Min. Humidity Date Max. Min. Humidity 15 July 72.0 51.0 86 4 August 57.0 49.0 65 A sez 55.0 48.0 55 ele 59.0 47.0 50 Arg ae 64.0 53.0 88 te 58.0 45.0 58 1 Blea 64.0 54.0 58 pony 67.0 40.0 57 19 46.0 70 eat 54.0 40.0 79 De 72.0 48.0 74 ae 52:0 39.5 82 ps ay 82.0 50.0 82 ke 65.3 45.0 67 A ahh, 93.0 61.0 69 i Pe A 38.6 53 pe Ua 92.0 64.0 51 12 : 68.0 47.5 74 ~ | ia 61.0 54.0 79 1s el 49.0 38.0 86 ae 69.0 49.0 60 ist 58.5 35.5 93 a ea 70.0 47.0 37 year 66.5 32.0 ref 7) aed 72.0 47.0 33 ll bee ed 64.5 51.5 94 29 «“ 71.0 53.0 47 i EF eh ia 62.3 43.3 82 Ft ae 61.0 51.0 43 bees f 60.5 36.5 74 Eth 58.0 48.0 94 iL earl 70.0 B1.5 65 1 August 61.2 45.0 68 5 A a 60.3 48.3 100 Batt 59.0 53.5 100 Bikwak ts 67.0 33.8 76 5 eS oe 48.0 84 It can be seen from the graphs of maximum and minimum temperatures (Graph 1) that the average of both has a tendency to be lower after the 15th of July. Also, it can be seen that the maximum temperature after the 30th of July averaged 60° F. and fell below 50° F. on the 15th of August. It seems that the 60° F. temperature is the critical one for initiating the fructification of P. circinatus. Since the figures given in Table I and in Graph 1 are for air temperature it should be pointed out that the temperature of the soil and of the tree would most likely be neither as low nor as high as the air temperature. Although those data have been taken only during one season and at one locality, they seem to indicate fairly well that when the average temperature of the day ap- proaches the minimum temperature for mycelial growth, the vegetative phase is checked and the reproductive stage is initiated a short time before fruiting bodies actually appear. Reference to Graph 2, presenting the percentage of relative humidity of the air against the time of the year make it evident that after the first day of August and up to the time fruiting bodies appeared, the percentage of relative humidity did not go below 50% and averaged 81.7%, while before that date, from the 15th of July up to August, the relative humidity was as low as 35% and averaged 64.3%. From the 15th of August up to GrapH 1. (Above) The relation of maximum and minimum temperatures to the appearance of sporophores. GrapH 2. (Below). The relation of atmospheric humidity to the appearance of sporophores. GOSSELIN: STUDIES ON PoLysTicTUS CIRCINATUS Sao TEMPERATURE Appearance of the first sporophore Degrees Fahrenheit 4 { 7? i duly he Anon, HUMIDITY Appearance of the first sporophore Percentage of relative humid lity of ihe air 1 t July * August GrapH 1, 2 536 Fartowia, Vou. 1, 1944 September, the best time for sporophore production, the relative humidity was never lower than 65%, with an average of 85.7%. We have remarked © that the sporophores come on trunks or roots always very close to the soil, a fact that substantiates the evidence tending to show that a relatively high humidity favors not only growth, but at a later date, when vegetative growth has been checked, also favors the production of fruiting bodies. Just how much the percentage of relative humidity becomes a limiting factor cannot be asserted from the data we have, but it is at least certain that an average of around 80% is favorable. As far as light is concerned, it can be said that the sporophores are more generally found in diffused light. However, in one case, one was found exposed to the afternoon sunlight. From laboratory studies it has been found that the related P. tomentosus grows better in diffused than in bright light or in darkness. Although not conclusive, these data furnish good indications that the appearance of the sporophores of P. circinatus is correlated principally with temperature and humidity and that the time of fruiting corresponds with the period when the low temperature checks vegetative growth and stimulates reproductive growth. This period also correlates with the time when the average relative humidity is on the increase and thus is furnished the water necessary to enable the fungus to complete the fruiting stage and to ripen the sporocarps. Exposure to light appears, if it is im- portant at all, to play only a very secondary réle in initiating the forma- tion of the fruiting body. However, it is possible that once the fructifica- tions are above the surface of the soil, light may have some influence on the formation of the hymenium and on the production of the spores. If this is so, since the fruiting bodies are produced mostly in the deep shade, it is evident that only very low intensities of light are needed to exert an influence on fructification. Errects on Host In addition to some ecological data concerning the production of sporo- phores, field observations have also given some information concerning the effects of this disease on its host as, for instance, the external charac- ters which diseased trees present and the kinds of breaking the rotted, fallen trees will present. The writer has observed that in early stages of infection the trees do not present externally any signs of disease, but as the infection reaches an advanced stage, the foliage of the trees turns pale yellow-green. Another external character which sometimes accompanies the advanced stage of decay is the exudation of resin at the base of the trees. Although these characters are not specific to the infection by P. circinatus, they help to spot the disease in regions where it is known to occur. It has already been stated that field observations indicate that the infection of the host takes place through the lateral roots, as is shown by the fact that when subjected to strong winds the trees blow over very readily and in a char- GOSSELIN: STUDIES ON PoLysTICTUS CIRCINATUS DOU acteristic manner. By the comparison of trees infected by P. circinatus and of those attacked by Poria subacida, this point can easily be brought out. In this latter species, it was found that penetration was through the central or tap root and from there the rot progressed into the stump leaving the lateral roots unharmed; as a result the trees broke off about one to two feet above ground level. In P. circinatus, on the other hand, when the lateral roots have been weakened by decay the trees blew over and at the same time the tap roots were pulled from the ground. Further studies substantiate the above evidence. For example, at Shelter Bay in the valley level where both fungi occurred, the trees were blown over in a manner characteristic of trees infected by the two species of fungi. In this connection it should be pointed out that in the valley the soil is sufficiently deep so that the spruces develop both tap and lateral roots. On the moun- tain slopes however, where the rock is covered only by a layer of humus, and as a result tap roots are not developed, Poria subacida was absent and only P. circinatus was evident. Under such conditions as are present in the mountain habitat, only lateral roots could be infected. Thus there is furnished the added evidence that P. circinatus is confined to the lateral roots, at least during the early stages of growth. GROWTH STUDIES During the course of the studies in connection with penetration of the host by the fungus it became evident that it was desirable to make a com- parative study of the growth of trees infected by P. circinatus*® and non- infected ones, and at the same time to endeavor to discover the relations between the habitat and the infection of the trees as illustrated by their rate of growth. A survey was also made to find out whether or not there was any correlation between the age of the tree and the incidence of butt- rot. For this purpose studies were made in five localities, namely: the valley and mountain slope and summit in the vicinity of Lake Walker on the North Shore of the St. Lawrence River, and in the vicinity of York River and Lake Ste. Anne on the Gaspé Peninsula where ecological condi- tions seemed to be similar. In these studies sections were made at D.B.H. of trees infected by P. circinatus and those not infected. In each type, the age of the tree was determined by a count of the annual rings and the diameter of the tree measured for each ten-year interval. The trees were segregated in age classes: (40-60) (70-90) (100-120) ... etc. For each year class and for each locality the diameter of the average tree was calculated and from that the area of D.B.H. section was obtained for each ten-year interval. * During the discussion on the growth rate of the trees, the terms “trees infected by P. circinatus” and “trees non-infected by P. circinatus” have been used having in mind the fact that the trees do or-do not present visible signs of decay. It cannot be stated that trees which had no decay caused by P. circinatus were not-in mycor- rhizal-association with this fungus, nor can it be said that no other mycorrhizal fungus is present. 2 538 FarLowiA, VoL. 1, 1944 We assumed that the surface of the section of a conifer at breast height increased in proportion to the increase in volume of the tree. The growth rate was obtained by subtracting the logarithm of the surface of the sec- tion of two subsequent decades. Appendix I gives the area at D.B.H. section for each ten years and for each classification, Appendix II gives the corresponding growth-rate. In Graph 5, the rate of growth of infected and non-infected trees for the 130-150 year class have been plotted for different sites. From this graph it can be seen that in the York River and the Valley areas where the soils are favorable, the rate of growth of non- infected trees is greater than that of infected trees. Contrarily, on the East Slope and Mountain Top sites where the soil is shallow and leached and hence less favorable, the rate of growth of the infected trees is greater than those that are not infected. The significance of these facts will be discussed shortly. At this point it may be objected that there are not valid grounds for drawing any conclusions since the trees of each class may have encountered different ecological conditions during the different years of their growth. To answer this objection, the average rate of - growth of trees for a given site and of infected and non-infected trees, together with the rate of growth of individual trees in each classification, was plotted by age (Graph 3) and by year (Graph 4). A comparison of these two graphs demonstrate that although there are minor differences, there is in general a very close agreement between the results derived by the two methods. Thus, whichever method is employed, the results are approximately the same and equally significant. For convenience and as a resumé of Appendix II, Table II has been prepared to show for each age class and for each region the percentage of number of decades in trees rotted by P. circinatus in which the rate of growth was higher than those not rotted by this particular fungus. Accompanying these percentages are symbols which indicate whether the area at D.B.H. section of the average tree rotted by P. circinatus is greater, equal, or smaller than that of the non-rotted trees. Table II Gaspé Peninsula Lake Walker Area Age class L. Ste. Anne York River Valley East Slope Mountain Top 40-60 50% [] 0% -- 67% ++ 70-90 100% + 86% + 75% ++ 29% [] 100-120 30% ++ 70% -- 4%+ 56% + 64% ++ 130-150 42% —— 42% — 54% + 42% ++ 160-180 388% —— 74% ++ 190-210 38% —— Average 62% 538% 38% 65%. - AT% Area of rotted trees definitely higher ++, somewhat higher +, indifferent [ ], somewhat smaller —, definitely smaller ——. GrapH 3. The average rate of growth (heavy lines) and the growth of individual trees (light lines) of the 130-150 year class plotted by age. Grapu 4. The average rate of growth (heavy lines) and the growth of individual trees (light lines) of the 130-150 year class plotted by year. ee ee Growth rate 0.9 fr 0.9 - aa 0.8 og f : \ O7- i O7- \ ——. Trees infected by P,circl \ . ——— Trees non-infected by P.circinatus Or ae ee Mme Pare 0g \ Nett eet eal a tN a Bee ono Trees infected by P, circinatus ‘ \ ==---+ Trees infected by P. circinatug ; 0.6 0.6 F 2 r) he 0.5 € 05 F = 2 o Ode ode O.3- 03 - O2- 02 SS OLE orb s oN No n n n 1 i n 1 1 10 1872 1882 1392 1902 1912 1922 1932 1942 : Date GraPH 3 GRAPH 4 SALYNIDUI) SALIILSATOG NO SAIGNLS :NITASSON 6€S 540 Fartowia, VoL. 1, 1944 AGE CLASS 130 = 150 GRAPH 5 ri York River 1 w % 0.45 + ‘ _ g J qq 0.25 + \ "———~ Trees non-infected by P.circinatus | \ o----* Trees infected by P.circinatus - | | t mae 1 1 ? . : aie SEES mote oe) P ce = ? R Mountain Top Aaa E \ a J t el rf £ 2 1 t 1 ie alae ee. ee es 10 30 60 70 90 110 130 160 ones “He'd ¥# wory soyou] ouenbe ne eee eS ee GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS 541 Reference to Appendix I and Table II shows that in the two localities: York River and Valley, the area at D.B.H. section of the average rotted tree by P. circinatus is lower than that of the non-rotted trees except in age classes 70-90 and 100-120, but in all other localities the rotted trees have larger area at D.B.H. section than the non-rotted ones. Thus, it can be concluded that either the infection of the tree by this fungus is beneficial to the host or else that this fungus invades rapidly growing trees. This last hypothesis has to be discarded because the data from York River and Valley regions (Graph 5) show that the reverse is true. Examination of growth rates, Appendix II and Table II more especially, show that in all cases there always has been a period during the life of the tree when the growth-rate of trees infected by P. circinatus was higher than the non- infected ones, except for the age class 40-60 of York River. The per- centages given in Table II show how long was this period. To explain why in York River and in the valley of the Lake Walker site, the area at D.B.H. section of trees infected by P. circinatus is smaller, it may be said that in these two regions the conditions for growth of the trees were not as poor as in the other regions, and that the growth of young non-rotted trees was at the beginning very good, while for some unknown reason it was some- what delayed in the trees rotted by P. circinatus. Thus. starting with a larger capital, the area at D.B.H. continued to be larger throughout the life of the tree even though the rate of growth was smaller during a few decades. On the other hand, in the other localities where the conditions of tree growth were poorer, the initial capital of the non-rotted trees was smaller or only slightly greater and was later exceeded by the faster growing but rotted trees. In general it can be said that when there is little sub-soil, as under the conditions existing in the mountain slope and top, the trees show a larger diameter when rotted than when not rotted; on the contrary, when there is a considerable depth of sub-soil such as exists in the valleys, then the reverse holds true, but in all cases, with one exception, rotted trees always present a. higher rate of growth, at least for a time, than do the non- rotted ones. Soi, CONSTITUTION Not having found any correlations between the occurrence of the disease and the state of crowding or competition between species, or the appear- ance of any insect epidemics, to explain the different relations between fungus and host in the various habitats, we have tried by soil analysis to correlate the occurrence of this disease with some salt deficiency. For this purpose we have used the method given by Hester, Blume and Shelton (1937) for rapid soil analysis. The results are shown in Appendix III which have been made to illustrate the effect of the concentration in the GrapH 5. The rate of growth and area at D.B.H. section of infected and non- . infected tree at four different sites. 542 FarLtowia, VoL. 1, 1944 soil of phosphorus, CaO, NH4, and potassium on the percentage of rotted trees in the stand. The conclusions are briefly summarized as follows: 1) — Phosphorus: From 0% to 30% infection, increasing concentra- tion of phosphorus is correlated with increasing infection of the stand, but for higher infection than 30% no correlation can be made with the con- centration of this element. 2) — Calcium: From 30% to 80% infection, increasing concentration of calcium seems to be correlated with increasing infection of the stand, but outside of these percentages of infection, nothing can be correlated. 3) — Ammonium: From 10% to 80% infection, increasing infection of the stand is fairly well correlated with increasing concentration of am- monium in the soil. 4) — Potassium: Up to 50% infection, decreasing potassium concentra- tion in the soil accompanies increasing infection of the stand. 5) — H-ion concentration: In all the cases seen, infection was never found in stands where the humus had a greater H-ion concentration than pH 3.5. It seems that the average H-ion concentration has a tendency to be lower in stands with a higher percentage of infection, although this does not seem very specific. From culture studies, we know that this fungus can grow in artificial media having a pH between 3.5 and 8.6. The medium with a pH of 6.4 supported the fastest growth, but since there were not any cultures intermediate between pH 4.5 and pH 6.4, it cannot be stated that pH 6.4 is the optimum for growth of the fungus. Never- theless, the optimum pH appears to be around pH 6.4. The pH of the humus of almost all infected stands are comprised between pH 3.58 and pH 4.40. Thus, we can state that generally the mycelium infects trees when conditions are not the optimum for the growth of the fungus, and accordingly the infection of trees could be interpreted as a reaction of the fungus against adverse conditions. Thus, in so far as the percentage of rotted trees in the stand is not too high, the increasing degree of infection of the stand can be correlated with increasing concentration in the soil of phosphorus and ammonium, and decreasing concentration of potassium and calcium. Now, instead of correlating singly the elements with the degree of in- fection of the stand, we will consider the elements by groups of two: phos- phorus and calcium, ammonium and potassium. 1) — Phosphorus and calcium: It can be seen that, when infection is at or below 30%, the phosphorus concentration increases and calcium concentration decreases. From 30% to 50% infection, the phosphorous concentration decreases while calcium concentration increases. This goes on up to 70% infection. From that degree of infection up to 100%, calcium concentration will act in the same manner as phosphorus con- centration. : 2) — Ammonium and potassium; Ammonium and potassium concen- _ trations always act in opposite ways, except between 0% and 10%, 50% eae ee eee ar ee pe EE De ee ER eS eg ae ee ee i ees ae ie ae GOSSELIN: STUDIES ON PoLysTICTUS CIRCINATUS 543 and 60% infection of the stand. So, it seems that as long as there are opposite (or mirror) correlations between phosphorus and calcium con- centrations in the soil, the stand will have only weak infection. But as soon as these two elements correlate in the same sense, the stand is sus- ceptible to a high degree of infection. In general, however, the correla- tion between ammonium and potassium does not seem so important. NEEDLE ANALYSIS According to experiments done at the Black Rock Forest, New York, the salt concentration in the leaves, or needles, is nearly proportional in pure culture to the concentration of these elements in the soil. So, to complete the information given by the soil analysis, we have analyzed needles of spruces from localities in which there were different degrees of rotting. The potassium, ammonium and phosphorus percentage of the dry weight increase in the needles with the increase of infection up to 30% (see Appendix IV). Even if in the soil the potassium concentra- tion is low, the percentage of dry weight of this element in the needles is relatively high. It should be remarked also that the data in the three cases are about similar. Although there are some indications from the soil analysis that increas- ing phosphorus and ammonium and decreasing potassium and calcium will favor rotting, those indications are valid only in the cases of low percentages of rotting. There must be some other factors which, in higher percentages, will govern the infection, but unfortunately we have not been ‘able to determine such factors. Even the needle analysis does not give much information for greater rotting than 30%. EVIDENCE OF SYMBIOTIC RELATIONSHIP If soil and needle analysis do not give much information as to the factors determining the occurrence of rot, they do furnish good evidence that this parasitic fungus forms a symbiotic association with its host before becoming parasitic. The evidence is furnished by the fact that, as has just been pointed out above, under unfavorable conditions existing on the moun- tain top, rotted trees show greater growth than the non-rotted trees, whereas in the valleys where the essential nutrient elements have accumulated, the reverse is true and indicates that the symbiotic relationship is only effective when there is some mineral deficiency. The evidence is also furnished by the fact that infection takes place by the lateral roots and that sporo- phores of P. circinatus have been found associated with tree rootlets. The results of needle analysis also tend to show that there is a symbiotic rela- tionship between the host and the fungus. As stated previously, the introduction of the fungus into the host is made by way of the underground parts of the tree. Thus, the only possi- ble way for the mycelium to penetrate the host will be either by under- ground scars, by direct penetration through the bark or symbiotic asso- ciation with the tree. We have dug out about a dozen rotted trees in Ste. ee 544, FarLowl1A, Vout. 1, 1944 Catherine and in Shelter Bay, and there are only a few cases where infec- tion took place through scars on the roots. In all the other cases we could not find any particular place where the mycelium would have pene- trated. Rather, we have seen the rot behaving exactly as a river. The main body of the river is the main lateral root and all or many of the secondary roots bring rot to the main lateral roots, and these meet in the trunk. Dr. Faull, in a personal communication, stated that he followed the rot up to the very tip of a root. The writer has made similar observa- tions in Shelter Bay. Hence the mycelium must have penetrated directly through the bark or by symbiotic association. We do not have any evi- dence that it came through the bark, while we do have some for the sym- biotic association hypothesis. At least twice, sporophores of P. circinatus were found growing out of nests of rootlets. Hatch (1937) reported that sporophores coming out of rootlet-nests are quite frequent in the usual mycorrhizal fungi, and therefore since P. circinatus has twice been found connected with rootlet- nests, the evidence for its mycorrhizal relationships is further strengthened. As has been previously stated, in the Gaspé and in the Lake Walker area the growth rate of rotted trees was higher than that of healthy ones. It is accepted that trees with mycorrhizae will have a larger rate of growth than those without, at least when the mycorrhizal relationship is efficient. On the other hand, we have seen that in the Valley (North Shore) of the St. Lawrence River and on York River (Gaspé), the diameter was smaller for rotted trees than for healthy ones. This has been correlated with the fact that in these two places the soil is richer in nutrient salts and better, apparently, for tree growth without the aid of P. circinatus as a partner, at least when it does not cause any rot.1_ We have seen also that the humus of these two places had less phosphorus than the others had. Rosendahl (1941) found that when the phosphorus concentration of soil was low, although mycorrhizal relationship was established, the intake of nitrogen and potassium was less than in similar seedlings grown with an adequate supply of phosphorus. Therefore if symbiotic relationship is to be of greatest benefit to the tree, then a suitable concentration of phos- phorus is required. TaBLe III THe CONCENTRATION OF THE Various MINERAL ELEMENTS IN THE Soi Is GIVEN ACCORDING TO THE DIFFERENT LOCALITIES STUDIED pH Kk NH, Fr CaO Locality Inf Non InfNon Inf Non Inf Non Inf Non Gaspé Peninsula Lake Ste. Anne 3.81 3.79 31 35 23 19 5.00 3.638 165 1381 York River 3.80 38.74 39 35 26 24 4.75 6.50 97 260 L. Walker area Valley 3.74 4.07 20 22 20s 22 2.60 12.50 80 40 Mountain Top 4.06 3.75 40 8 13 23 12.50 4.20 80 24 East Slope 3.95 3.95 5 23 15 32 12.50 10.25 40 48 ‘Refer to footnote page 537. Pst. ee ee GOSSELIN: STUDIES ON POLYSsTICTUS CIRCINATUS 545 As seen from Table III the writer’s observations substantiate this con- clusion since in the valley of the Lake Walker area and York River area in Gaspé, the concentration of phosphorus was relatively low and this was correlated with a low percentage of length of time beneficial to rotted trees. It can be seen also that at Lake Ste. Anne where the concentration of phosphorus was not optimum, the infection in age class 100-120 was beneficial for only 30% of the time (Table II). The growth rates have been discussed in greater detail in a previous paragraph. Another good character which seems to indicate the symbiotic relations of P, circinatus with the trees, is the salt content of the needles. Routien and Dawson (1943) suggest “that mycorrhizae increase the salt absorbing capacity of the roots . . . The increased salt uptake may then be reflected in a greater rate of growth.” Finn (1942) stated: “It is seen that nitrogen and potassium were taken up in larger quantities by the inoculated seed- lings than by the uninoculated ones. Phosphorus was absorbed in a slightly larger quantity by the inoculated seedlings than by the seedlings which were not inoculated.” Thus, it is admitted now that plants with mycorrhizae will generally absorb more nitrogen, potassium and phos- phorus than those without. In order to check the mycorrhizal nature of P. circinatus, we have chemically analyzed spruce needles to find the percentage of these three elements in the needles both in rotted and non-rotted trees. The needle analysis reveals that the phosphorus percentage in relation to the dry weight of the needles is lower for rotted trees than for healthy ones in the general average and in the data from York River, while it is higher at Lake Ste. Anne and at Chandler. Data from Finn (1942) reveal that the phosphorus intake by the mycorrhizal seedlings is higher than for the non-mycorrhizal ones, but the percentage to the dry weight is smaller even though the intake is larger. The difference, as shown by Finn, is 0.020% while the writer found a difference of only 0.011%. On York River this percentage is lower or only 0.007% while at Lake Ste. Anne it is higher or 0.049% and at Chandler 0.054%. This, however, has not definite significance because when there is a arenes amount of phosphorus in the needles, there is a correspondingly greater concentration of this element in the soil. Thus, it cannot be unqualifiedly concluded that the infection by P. circinatus was beneficial to the trees although there seems to be evidence tending to indicate that such is the case. As in the phosphorus relations, the percentage of nitrogen in the needles of rotted trees is higher, in the general average, than in the needles of healthy ones. Although the general average is higher for rotted trees than for healthy ones (Appendix VI), the percentage at Lake Ste. Anne and at York River are lower, respectively of 0.002% and 0.003%, al- though the soil analysis from these places (Appendix V) shows that soil under rotted stands had a higher concentration of ammonium salts than the healthy ones. Even if the nitrogen percentage of the needles is lower 546 FarLowl1A, VoL. 1, 1944 (0.002%) for rotted trees than for healthy ones, this difference is so small that it probably is of little significance. However it is interesting to note that nitrogen concentration in the needles is higher for rotted trees than for healthy ones only when the concentration of nitrogen is lower in the soil. For example, at Lake Ste. Anne and on York River, the rotted stands had higher concentration of ammonium salt than the non-rotted ones and the reverse was true for the percentage of nitrogen in the needles. Exactly the contrary is found at Chandler. Neverthe- less there seem to be indications that the infection by P. circinatus is some- times beneficial to the host as a result of increasing the intake of nitrogen. For the percentage of potassium in the needles, the analysis revealed that, in the general average, this percentage is lower for rotted trees than for non-rotted ones; however in the rotted trees of Lake Ste. Anne and of Chandler this percentage is much higher. This low percentage in rotted trees seems to be prevalent almost exclusively at York River. At that place we have found percentages as low as 0.062, 0.071, 0.085, and 0.094. In none of the other places were percentages found as low as these, and it was remarked that these low percentages came with heavily rotted trees. In Gaspé only 22% of the infected trees were rotted as high as D.B.H. section, but of the trees which have a low percentage of potassium 60% were rotted up to the D.B.H. section. In general the potassium percentage of the needles is low in all rotted trees of York River, 0.236% compared with 0.372% at Chandler and 0.418% at Lake Ste. Anne. This excessively low average might perhaps be explained by the fact that 88% of the most heavily rotted trees were from York River region. It can be seen from soil analysis that, at Lake Ste. Anne, al- though the concentration of potassium in the soil was smaller for infected stands than for healthy ones, the percentage of this element in the needles was greater. It is logical to suppose that the infection, at least at Lake Ste. Anne, has been beneficial to the trees for larger intake of potassium. Thus if the data from York River region are excepted, we can state that the infection of the trees with P. circinatus has increased the intake of mineral salts. This beneficial effect should not be interpreted too categorically, but rather as a probability. It should be remembered also that the needles analyzed might, in many instances, have come from trees in which the parasitic have exceeded the beneficial effects of the previous symbiotic association. This would have lowered the average percentage for the rotted trees and might explain why, in the York River region where the trees are more heavily rotted than in the other regions, the percentages of all elements are so low. The field observations, in addition to some facts concerning the be- havior of the disease caused by P. circinatus, has brought out facts which seem to indicate fairly well that this fungus penetrates into its host by symbiotic association. Each fact in itself is not very conclusive but they all substantiate the same conclusion. See ae eee ae ae a eee Or ye ee Ree Mee to ee GOSSELIN: STUDIES ON PoLystTicTUS CIRCINATUS 547 EFFECTS OF SEEDLING INOCULATION WITH P. CIRCINATUS Having in mind all the evidence from field observations that P. circinatus was penetrating into the host through symbiotic association, the writer has tried to establish the mycorrhizal relationship between spruce seedlings and the fungus under laboratory control. First, we have tried to culture seedlings under aseptic conditions with the Hatch culture chambers, but the experiment did not succeed. Later, loamy-clay soil was obtained from Saskatchewan where no trees are grow- ing, and consequently in which there are no mycorrhizae forming fungi. This was suggested by the experiments done by Hatch (1937). After having sterilized and stratified spruce seeds, they were put in the ice box for two weeks in order to obtain a better germination. The loamy-clay soil was placed in wooden boxes previously sterilized and the seeds were sown. After germination in July, 1942, the seedlings grew in a green- house until January, 1943, when they were placed under the snow to allow a dormant period. At the beginning of April, 1943, they were dug out of the snow and put in the cold room (33° F.) for fifteen days to make the transition between the temperatures of the snow cover and the green- house less abrupt. After two weeks in the greenhouse, when the buds were bursting, half of the seedlings were inoculated on May 1, 1943, with cultures of P. circinatus, while the other half was reserved to serve as a check, although maintained under the same growing conditions. After three months of growth, namely on August 31, the seedlings from both series were removed and studied to determine what difference ex- isted. It was immediately obvious (Plate II) that the inoculated seedlings had not only made a greater growth but that the needles were of a deeper - green color and that the primary roots had produced more secondary roots. To determine the relative growth of the inoculated seedlings and those not inoculated, they were measured from the primary node to the apex. The results of these measurements are presented in Graph 6 in which the actual figures are represented by crosses for the non-inoculated seedlings and circles for the inoculated ones. These figures are based on the meas- urements of a hundred seedlings for each series. It is immediately obvious that the non-infected seedlings show a fairly definite curve, whereas the inoculated ones follow essentially the same curve, although there is some deviation from the curve presented by the non-inoculated seedlings. This deviation, it seems, may be explained by the fact that the period of growth after inoculation was so short that complete mycorrhizal relationship had not yet become established. Despite this erratic behavior, however, it is quite obvious that even in so short a time as three months, the fungus did stimulate the growth of the spruce seedlings. A further examination of the seedlings shows that while the roots of the uninoculated ones are not infrequently longer than the inoculated ones, nevertheless the latter were characterized by the production of a greater number of secondary rootlets which were frequently branched. In the eee FES Se Ag pelt eo) dh tae le Ne en ois hs 548 FarLtowia, VoL. 1, 1944 uninoculated plants these secondary rootlets, comparatively speaking, were very rarely branched. Thus, it can be seen that the fungus in this in- stance has added to the efficiency of the root system for absorption. This becomes all the more evident when it is seen that, despite shorter roots with increased secondary rootlets, the secondary internode of the stem of the inoculated seedlings, on the basis of ten specimens taken at random, totaled to 142.5 mm. whereas in the non-inoculated seedlings that were measured in the same way, the total was 113 mm. Furthermore, the 28 265 24- Frequency - r q \y/ peat as lL 1 L l 1 i js a! _J ll 12 13 #14 «#215 16 «#17 «#618 «#619 @ Length of internode in millimeters GrapH 6. The relative rate of growth of inoculated (0) and non-inoculated (+) seedlings. inoculated seedlings bore 727 needles and the uninoculated 548 needles, or respectively 51 and 48.5 needles per cm. of internode. In addition to these obvious differences between the two series, there is the further difference that is furnished by the darker color of the needles of infected seedlings. The needles of non-inoculated seedlings in contrast to those inoculated were mostly yellowish green and in addition show the presence of anthocyanins, an indication, according to Mitchell (1934), that there is a phosphorus deficiency in the soil. Since the inoculated trees did not show this symptom of deficiency, it seems likely that the fungal symbiont helped to make available a larger phosphorus supply. Un- fortunately the information is not available to enable us to make any statement as to how the process takes place. GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS 949 The obvious differences observed in the size of inoculated and un- inoculated seedlings, and the greater number of needles per centimeter of internode for the inoculated than for the uninoculated seedlings, led us to examine the microscopical characters of rootlets of the inoculated ones. Rootlets which we considered as having mycorrhizal relationship on account of their larger size were killed and imbedded in paraffin. It can be seen from the sections that the mycorrhizal relationship is of the regular type. The Hartig net is two to three cells deep, without apparent intracellular invasion. The mantle is homogenous (Plate III, fig. 1) and somewhat parenchymatous in structure, about three to four cells deep, with a smooth margin from which arise very few hyphae which penetrate into the surrounding cells. Sections made of rootlets found in a rootlet nest (Plate III, fig. 2) under a sporophore of P. circinatus show essen- tially the same type of mycorrhizal relationship. The mantle is homo- venous, although thicker, parenchymatous in structure, with a smooth margin. The tannin layer is not very apparent, and locally seems to be two cells thick. The Hartig net is not so thick as the type we obtained by inoculation, but essentially the same type of mycorrhizal relationship is evident in the artificially and the naturally invaded rootlets. Thus, although the experiment was not done under aseptic conditions, it seems very probable that the endophyte here concerned is P. circinatus. When all characters are taken into consideration, it seems safe to con- clude that the fungus has a beneficial effect on its phanerogamic partner, at least in the early stages. Also these experimental results tend to sub- stantiate the conclusions already drawn from field observations. The observations that we have made so far, either on growth rates or on soil or needles analysis, seem to indicate that the infection of the trees by P. circinatus behaves as if the fungus was mycorrhizal. However, in our northern forests where the pH of the soil is relatively high and where nitrogen is chiefly supplied as ammonium salt, it would seem that the trees must be in mycorrhizal association to live under such conditions. We have seen that trees infected with P. circinatus behave, in comparison with trees not infected by this species, much in the same manner as trees that are in association with known mycorrhizal fungi. This relation be- tween the fungus and the tree exists until some unfavorable factor, as yet unknown, upsets the balance between the symbionts. When this happens, then it would seem that the fungus readily becomes parasitic, and the mycelium, instead of remaining intercellular, as in the Hartig net, pene- trates into the cells (Plate III, fig. 3) and travels backward along the rootlets, killing them as it proceeds. This would then explain the reason why the butt-rot seems to have its inception in the lateral roots rather than through wounds or through the tap root. In Polystictus circinatus, then, we have a butt-rot fungus that in the earlier stages of the develop- ment of the trees, instead of being a parasite, actually is beneficial until the host reaches a certain age, which might indicate senescence, the age being determined by local ecological conditions. FSAI CBee 990 FarLtowia, Vou. 1, 1944 SUMMARY The white pocket butt-rot of conifers caused by Polystictus circinatus (Fr.) Cke. has been studied because of its local importance in the Province of Quebec and because of its peculiar behavior. Field observations have been made to correlate the occurrence of this disease with some ecological factors. The disease was localized in such definite loci of infection that it was thought that it was tied up with con- ditions of temperature, inséct epidemics, mechanical damage, or with the chemical nature of the soil. However, none of those factors appeared to be responsible for the occurrence of the disease. i At the same time as the ecological factors were discarded as having no correlation with the disease, the field observations brought up evidence that this disease penetrated into its host by mycorrhizal association. Al- though none of the facts furnished any conclusive evidence, they all sub- stantiate the same conclusion, To bring more evidence, the writer has tried to establish mycorrhizal relationship between spruce seedlings and the fungus. The results indi- cate that the fungus has really formed a mycorrhizal association of the usual type. If P. circinatus penetrates into its host through symbiotic association, and there is presumptive evidence that such is the case, this fact will open an entire new field in forest pathology. It will associate the two concepts of symbiotism and parasitism which were previously separated by air-tight walls. Those who are specifically interested in symbiotic problems will have to consider the pathological aspect of the question and the forest pathologists will have to be better acquainted with symbiotism in order to deal correctly with certain of their problems. Foresters will no longer be able to consider the butt-rots only as a reducing factor for the volume of the tree but will have to determine under which conditions the damage from parasitism exceeds the beneficial effects of symbiotism on trees, if it is demonstrated later, as it is not at all illogical to suppose, that butt- rotting fungi penetrate into the host by mycorrhizal relationship. These fungi in the Province of Quebec are all quite abundant; before trying to eliminate them by short cycle rotations, it will be necessary to know what part they are playing in the biological complex of our forest soil. Future researches will have to determine under which conditions parasitism follows the symbiotic association. This field is new and wide open. It will certainly be of considerable interest to those who will have the advantage of working on such prob- lems, and it promises to be at the same time of practical interest to foresters. GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS 551 ACKNOWLEDGMENTS The writer wishes to thank Dr. David H. Linder, under whose direc- tion this work was done, for giving his time so spontaneously, for his imaginative suggestions on possible hypotheses, his objective criticism of experiments, and his correction of the manuscript. He also wishes to thank Professor Emeritus J. H. Faull under whom this study was initiated and who furnished the first knowledge on this disease; and to Dr. Gast of the Harvard Forest, who has added to the writer’s knowledge of mycor- rhizal problems, for constructive criticism of this study, and for informa- tion concerning methods for the chemical analysis of spruce needles. The writer wishes also to thank the officials of the Department of Lands and Forests of Quebec for having made possible this study by the financial help they provided for the academic studies and for the field investiga- tions; among these officials the writer wishes to thank especially M. A. Bédard, Deputy-Minister, M. F. Boutin, Chief of Forestry Service, and Dr. R. Pomerleau, Director of the Bureau of Forest Pathology, under whom the writer has been working during all this study. He wishes also to thank Professor Z. Rousseat of Laval Forestry School for his unfailing interest in this study and for the laboratory facilities he extended to the writer at the Forestry School. ~ __ The writer is indebted to Dr. Amiot for the help he so kindly gave during chemical analysis of needles, to M. Y. Garand, Soil Analyst in the Department of Colonization, for advice on soil analysis, and to M. G. Bolduc, Forest. Ranger, for his whole-hearted help during field and lab- oratory work. The writer is also indebted to Dr. W. H. Gileson, Superintendent of the - Experimental Farm, Indian Head, Saskatchewan, who so kindly furnished the prairie soil. HarvaArD UNIVERSITY Camepripce, Mass. 552 FartowlA, VoL. 1, 1944 Appendix I | AREA OF D.B.H. SECTION FOR EACH AGE CLASS Dara FROM YORK River, GASPE | Age Class: I (40-60) (70-90) (100-120) | (130-150) Number of trees: | (1) (6) (13) (19) (25) (24) (2) (4) A B A 5 A B A B {i 34 1.27 1.34 0.61 1.09 0.58 1.03 0.21 5.35 4.13 4.48 2.41 3.06 1.61 4.97 0.58 20.37 8.44 7.96 5.48 5.88 2.77 ROL aT I15 49.15 18.24 11.08 9.45 7.98 4.01 13-87 1.61 40.65 16.73 14.29 14.08 9.80 5.62 20.39 2.24 58.67 23.00 17-00. 17°91 11.65 7.34 26.10 2.77 21.40 22.20 13.87 9.10 0,30 3.90 22.20 26.36 16.05 11.08 4.80 4.84 26.36" 18.38 13.04 46.20 5.89 20.89 14.72 53.00 F288 25.20 17.91 31.50 8.61 19.62 Poe 9.81 77.00 11.08 Data From Lake STE. ANNE, GASPE Age Class: (40-60) (70-90) cea Number of trees: \ (7) (3) (23) iQ) (4) (3) A B A B A B 0.54 0.67 0.39 0.26 0.12 0.39 1.76 1.91 1.47 1.21 0.50 1.34 3.26 Peay § 2.96 2.96 1.03 2.77 5.09 5.88 5.22 | 5.62 1.99 5.35 7.18 7.64 7.64 8.44 2.96 8.44 7.80 10.15 12.43 4.72 12.04 12.84 16.27 6.88 16.27 13.87 18.63 9.28 21.14 19.87 12.24 27.80 14.08 32.80 19.62 42.10 44.30 A—tTrees not infected (see footnote page 537). B—Trees infected with P. circinatus. GOSSELIN: STUDIES ON PoLysticTus CIRCINATUS Appendix I (continued) DATA FROM THE VALLEY, LAKE WALKER AREA Age Class: (100-120) (130-150) (160-180) Number of trees: (44) (5) (45) (23) (7) (9) A B A B A B 0.20 0.32 0.18 0.16 0.23 0.04 0.67 La 0.72 0.58 0.98 0.11 1.40 2.68 1.54 1.34 2.24 0.26 2.50 5.09 2.68 2.32 4.36 0.42 3.90 7.80 4.12 3.68 Tale) Oat 5.48 10.71 5.15 5.36 1k.27- 7106 8.12 13.04 7.80 7.03 16.05 1.47 10.538 15.82 9.98 9.10 21.14 ° 1.99 12.84 20.12 12.64 11.46 26.65 2.59 16.73 20.88 14.93 13.66 32.15 3.386 22.78 17.44 15.82 Sholo yA ie 20.12 18.15 43.21 5.22 23.00 19.87 49.72 6.02 55.45 6.88 60.61 7.64 67.38 Data FROM THE East Store, Lake WALKER AREA Age Class: (40-60) (70-90) (100-120) (130-150) Number of trees: at) (10) (2) (11) (6) (5) (1) A Bas A B A B A B 0.42 0.77 0 bbs" 0.18 0.20 0.18 0.20 0.18 2.41 1.99 0.42 0.46 Oot 40). 72 0.62 0.62 3.06 4.12 ieQesiy lL 1.40 1.48: a Eas gee 3) el 5.09 8.12 1.68 1.91 2.0 412.56 2.24 1°76 7.18 TA | aie 8.26 4.12 2.16 2,16 4.24 4.97 4.60 6.30 4.24 4.48 5.62 7.68 5.48 8.44 5.48 6.59 6.88 10.34 eUGs Wy. 7 1 6.88 8.61 7.96 15.82 9.98 138.04 8.61 11.46 12.04 15.16 10.58 13.24 12.64 14.50 15.16 15.82 16.97 17.44 18.63 19.62 A—tTrees not infected. B—tTrees infected with P. circinatus. B1s73) (190-210) (2) (3) A B 0.10 0.01 0.29 0.04 0.54 0.06 1.15 0.13 1.99 0.26 3.36 0.39 4.36 0.62 6.02 0.82 7.18 1.09 9.10 1.40 11.65-. «1.76 18.87 ~ 2,93 16.97 2.59 20.87 3.06 23.81 3.46 26.92 3.79 30.25 4.24 33.78 4.36 37.83 4.60 (160-180) (1) (1) A B O48) O48 0.39 0.50 On77) IS 124% 2.24 1764. O79 2.24 8.35 287. wag 3.47 9.63 4.48 12.64 5.35 14.50 6.16 16.73 7.03 19.62 8.61 22.19 9.63 24.65 10.16 26.36 11.46 28.79 504 FarLtowi14, VoL. 1, 1944 Appendix I (concluded) Dara FROM THE Mountain Top, LAKE WALKER AREA Age Class: (70-90) Number of the trees: (21) (3) > ie CNORNH OO EOE Ean] y DarAWnAI DMO — =) or on —_ Age Class: (40-60) Number of trees: (1) (1) A B 0.6020 0.5104 0.5806 0.3110 0.1982 0.1956 0.1018 0.1016 0.1594 0.13880 GROWTH RATE BY AGE CLASSES (100-120) (31) (1) A B 0.16 0.138 0.58 0.39 1.21 0.50 Dae 2.87 3.26 4.48 4.84 7.03 7.03 10.16 9.09 14.29 1427 17.44 12.74 20.37 15.16 23.81 15.37 25.79 Appendix II Data FROM YoRK River, Gaspé (70-90) (13) A 0.5246 0.2498 0.1438 0.1104 0.0980 0.0774 0.0158 0.0748 (19) B 0.5864 0.3574 0.2366 0.1730 0.1044 0.0932 0.0748 (100-120) (25) A 0.4484 0.2842 0.0956 0.1260 0.0750 0.0756 0.0634 0.0590 0.0554 0.0818 A—Trees not infected (See footnote page 537). B—tTrees infected with P. circinatus. (24) B 0.4436 0.2354 0.1608 0.1460 0.1160 0.0934 0.0858 0.0706 0.0526 0.0852 0.0396 (130-150) (13) (1) A B 0.16 0.20 0.58 0.98 1.09 1.99 1.83 4.12 2.87 6.59 4 24 10.16 5.89 13,24 7.96 15.82 9.98 17.44 12.43 19.62 14.72 21.14 17.20 22.19 19.62 23.81 24.65 (130-150) (2) (4) A B 0.6826 0.4520 0.3388 0.2960 0.2072 0.1468 0.1670 0.1422 0.1072 0.0932 0.0646 0.1484 0.0602 0.0940 0.1238 0.0848 0.0592 0.0956 0.0648 0.0694 0.0634 0.0566 0.0342 0.0532 GOSSELIN: STUDIES ON PoLysTicTUS CIRCINATUS Appendix II (continued) DaTA FROM LAKE STE. ANNE, GASPE Age Class: (40-60) (70-90) Number of trees: (7) (3) (23) (9) A B ay B 0.5142 0.4556 0.5822 0.6716 0 0.2682 0.2718 0.3036 0.3884 0. 0.1938 0.2168 0.2464 0.2780 0. 0.1492 0.1134 0.1654 0.1770 0. 0.0358 0.1238 0.1682 0. 0.1014 0.1168 0. 0.03386 0.0588 0. 0.0180 0. 0 0 Data FROM THE VALLEY, LAKE WALKER AREA Age Class: (100-120) (180-150) (160-180) Number of trees: (44) (5) (45) (23) (7) (9) A B A B A B 0.5166 0.5576 0.6020 0.5706 0.6274 0.4682 0.3216 0.3672 0.3328 0.3628 0.3604 0.3522 0.2500 0.2794 0.2398 0.2392 0.2898 0.2128 0.19388 0.1850 0.1878 0.2002 0.2170 0.2014 0.1480 0.1378 0.1442 0.1636 0.1956 0.1878 0.1704 0.0854 0.1324 0.1176 0.1536 0.1544 0.1128 0.0840 0.1072 0.1118 0.1198 0.1310 0.0862 0.1044 0.1024 0.1004 0.1006 0.1140 0.1152 0.0162 0.0726 0.0762 0.0814 0.0888 0.0368 0.0672 0.0638 0.0626 0.1024 0.0622 0.0596 0.0658 0.0620 0.0580 0.0394 0.0610 0.0580 0.0474 0.0454 0.0386 0.0460 A—tTrees not infected. B—Trees infected with P. circinatus. A eh eRe OE PEI EC Rt ee ee ER pe Ta 555 (100-120) (4) (3) A B . 63874 0.5408 6168 0.38162 2854 0.2858 1726 0.1980 2024 0.1542 1640 0.13808 1298 0.1138 1200 0.1192 .0610 0.0714 . 1440 0.1084 0.0224 (190-210) yee) A B 0.4748 0.4860 0.2724 0.2182 0.3296 0.38204 0.2384 0.2818 0.2278 0.1742 0.1126 0.2096 0.1406 0.1158 0.0774 0.1262 0.1026 0.1100 0.1076 0.0978 0.0756 0.1044 0.0876 0.0632 0.0794 0.0730 0.0678 0.0542 0.0536 0.0386 0.0504 0.0490 0.0478 0.0118 0.0492 0.02382 996 FarLowiA, Vou. 1, 1944. Appendix II (concluded) Data FROM THE East SLoPE, LAKE WALKER AREA Age Class: (40-60) (70-90) (100-120) (130-150) (160-180) Number of trees: (9) (1) (10) (2) (11) (6) (5) (1) (1) (1) A B A B A B A B A B 0.7574 0.4152 0.4312 0.4082 0.5166 0.6020 0.4862 0.6666 0.4570 0.5682 0.1040 0.3166 0.3892 0.3992 0.3216 0.3128 0.3098 0.2652 0.2980 0.338 0.2214 0.2940 0.2130 0.2208 0.1856 0.2284 0.2444 0.1846 0.2428 0.2860 0.1492 0.2162 0.2454 0.1804 0.2182 0.1500 0.2544 0.1186 0.2290 0.1848 0.1694 0.1592 0.1842 0.1280 0.1516 0.1044 0.1500 0.1220 0.1894 0.0766 0.1268 0.1116 0.1678 0.1178 0.1460 0.0884 0.1292 0.1618 0.1084 0.0968 0.1160 0.0826 0.1092 0.0630 0.1846 0.0984 0.0854 0.0970 0.1244 0.1112 0.1180 0.0814 0.0654. 0.0874 0.0628 0.0774 0.0598 0.0794 0.0394 0.0614 0.0622 0.0790 0.0878 0.0572 0.0690 0.0490 0.0422 0.0878 0.0536 0.0405 0.0512 0.0488 0.0456 0.0234 0.0292 0.0522 0.0282 Data FROM THE Mountain Top, Lake WALKER AREA Age Class: (70-90) (100-120) (130-150) Number of the trees: (20) (3) (31) (1) (13) (1) A B A B A B 0.5726 0.4546 0.5706 0.4560 0.5706 0.6800 0.3422 0.4814 0.3194 0.1110 0.2736 0.3098 0.1978 0.1826 0.2826 0.7606 0.2260 0.3166 0.2128 0.1878 0.1476 0.19388 0.1940 0.2034 0.1608 0.1744 0.1718 0.1960 0.1702 0.1882 0.1442 0.0934 ~ 0.1620 0.1600 0.1424 0.1150 0.1390 0.1288 0.1118 0.1480 0.1310 0.0772 0.1892 0.0930 0.0864 0.0984 0.0422 0.0634 0.0676 0.0954 0.0512 0.0654 0.0678 0.07382 0.0326 0.0062 0.0346 0.0676 0.0210 0.0572 0.0306 0.0150 A—Trees not infected. B—tTrees infected with P. circinatus. GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS Appendix III 997 CONCENTRATION IN P.P.M. OF PHospHORUS, PorassiuM, CALCIUM AND AMMONIUM AND H-10N CONCENTRATION OF THE HuMUS OF STANDS OF VARIOUS DrGREES OF Rorrine Percentage of rotted trees in the stand Phosphorus Potassium Calcium Ammonium 5) a i 0% 5 aly .(3.82)* 35 (10) 154 (104) 33 (13) 3.83 (0. 5% 6.87 (4.06) 47 (5) 110 (78) 23 (2) 3.58 (0. 10% 6.25 (3.64) 39 (4) 300 (50) 19 (5) 4.22 (0 15% 5.0 40 165 (69) 20 3.70 20% 8.55 (3.33) 35 (5) 272 (78) 21 (4) 4.37 (0 30% 10.00 (3.54) 33 (6) 53 (38) 24 (5) 3.72 (0 50% 5.09 (8.45) 34 (14) 222 (99) 26 (2) 4.20 (0 60% 7.81 (3.98) 40 (8) 111 (67) 27 (10) 3.76 (0 70% 4.38 (0.63) 47 (5) 53 (19) 25 (5) 3.92 (0 80% 8.75 (8.57) 42 (24) 300 (41) 30 (7) 4,27 (0 90% 3.82 (0.82) 33 (12) 93 (67) 26 (8) 3.76 (0 95% 8.75 (4.61) 40 (10) 165 (85) 27 (3) 3.93 (0 100% 2.17 (2.05) 32 (13) 300 (50) 23 (7) 5.38 (1 * Standard deviation. Appendix IV PERCENTAGE OF THE Dry WerIGHT oF THE NEEDLES OF PHOSPHORUS, PoTassIUM AND NITROGEN Percentage of rotted trees in the stand om oo ooo Phosphorus .188% (0.055)* .146% (0.032) .143% (0.027) 157% (0.042) 194% (0.017) 184% (0.028) 166% (0.030) 148% 0.148% (0.029) 0. 0.153% (0.028) 182% (0.007) * Standard deviation. ooo ooo O'S ore Potassium .3839% (0.70) .277% (0.053) .278% (0.060) .404% (0.082) .545% (0.091) . 306% (0.034) .216% (0.070) .198% .232% (0.135) .264% (0.115) .294% (0.038) Nitrogen 29) 06) .5) 03) .09) 37) 25) 06) seis 16) 25) 02) 0.658% (0.062) 0.602% (0.051) 0.664% (0.042) 0.404% (0.082) 0.744% 0.631% (0.062) 0.620% (0.051) 0.626% 0.652% (0.093) 0.661% (0.061) 0.781% 558 Fartowia, Vou. 1, 1944 Appendix V Tue pH Anp THE ConcENTRATION OF K, NHy, P, CaO In THE SorL OF THE VARIOUS © Loca.ities STUDIED pH Kk NH, P CaO Locality Inf Non InfNonInfNon Inf Non Inf Non Gaspé Peninsula Gaspé Park 4.95 3.81 41 38 20 21 2.75 4.13 290 72 Lake Ste. Anne 3.81 3.79 3135 2319 5.00 3.63 165 131 Chandler 4.24 4.01 87 32 22 28 5.75 4.13 260 216 York River 3.80 3.74 39 35 26 24 4.75 6.50 97 260 Petit Pabos River : 4.60 4.65 438 40 19 25 5.00 12.50 3800 250 West Branch Pabos River 3.81 31 26 3.63 182 Pecan River 4.06 3.90 34 25 25 30 4.88 38.13 250 250 Four Lakes (Marsoui) 3.85 25 25 12.50 40 Petite Riviére-a-Marthe 4.30 3.85 5035 1717 5.00 4.38 250 80 North Side Mt. Albert 6.45 .2.90 -20 50.30 30° 0.75: 3:75 350. -40 Lac des Américains Trail 5.15 4.20 48 40 20 23 1.25 38.75 350 250 East Side Mt. Albert 6.45 3.90 20 50 380 30 0.75 3.75 350 40 Lake Walker Area Valley 3.74 4.07 20 22 20 22 3.23 12.50 80 40 Mountain Top 4.06 3.75 40 8 18 23 12.50 4.13 80 24 East Slope 8.95 3.938 5 23 15 32 12.50 10.25 40 48 Philibert Slope 3.81 30 18 8.75 56 Appendix VI Per Cent or Dry WEIGHT OF THE MINERAL ELEMENTS IN THE NEEDLES FROM THE DIFFERENT LOCALITIES STUDIED Phosphorus Potassium Nitrogen Locality Inf Non Inf Non Inf Non Gaspé Park 0.148% 0.271% 0.765% Lake Ste. Anne 0.224% 0.175% 0.418% 0.298% 0.671% 0.673% Chandler 0.169% 0.115% 0.372% 0.344% 0.650% 0.622% © York River 0.164% 0.171% 0.236% 0.351% 0.631% 0.634% General Average 0.164% 0.175% 0.264% 0.317% 0.654% 0.615% 560 FarLtowlaA, Vou. 1, 1944 EXPLANATION OF PLATE I Fig. 1. Polystictus tomentosus (Fr.) Fr. Side-view to show the distinct stipe which is nearly central. Fig. 2. Polystictus circinatus (Fr.) Cke. Side-view to illustrate the sessile, applanate fruiting bodies characteristic of the species. _ Fig. 3. Map showing the distribution of P. circinatus. It will be observed that the species has a predominantly northern distribution. GOSSELIN: STUDIES ON POLysTICTUS CIRCINATUS 361 3 Map showing the North American distribution of Polystictus circinatus Fr, @ References from literature; @ Specimens from Farlow Herbarium; * Places where the author's studies were made. Priate [ 562 FarLowia, VoL. 1, 1944 EXPLANATION OF PLATE II Shadowgraph taken from seedlings of Picea rubra and reproduced at one-half of actual size. Specimens on the extreme left of upper and lower row show branching. Note that the inoculated seedlings (below) have longer internodes above the first year node, that the needles are more numerous and that the root system is more robust and for the most part more abundantly provided with secondary rootlets. GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS PiateE II 564 FarLowia, VoL. 1, 1944 EXPLANATION OF PLATE II Fig. 1. Cross-section of rootlet of Picea rubra inoculated with P. circinatus, to show the mantle of pseudoparenchymatous tissue and the Hartig net which sur- rounds all the cortical cells. Fig. 2. €ross-section of rootlet of Picea mariana from rootlet nest from which the sporophore of P. circinatus had arisen. Notice the distinct pseudoparen- chymatous mantle at the left, and the beginning of the formation of the Hartig net. Fig. 3. Cross-section of a small root of Picea mariana which has begun to show root rot. Note the accumulation of dark tannin globules in most of the cells and also the faintly showing mycelium that has penetrated into the interior of the central cells after having passed through the walls. GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS 565 PiateE III 566 FarLowia, VoL. 1, 1944 BIBLIOGRAPHY Allescher, A. (1886). Verzeichnis in Siid-Bayern beobachteter Pilze. p. 74. Ames, A. (1913). Consideration of structure in relation to genera of the Poly- poraceae. 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Raleigh, North Carolina Inst. for the Deaf, Dumb and Blind. p. 97. Dodge, B. O. (1914). List of fungi of Kewaunee County, Wisconsin, Journ. Wisconsin Acad. Sci. Arts and Let. 17 (2): 827, 829. Donk, M. A. (1933). Revisio der Niederlandischen Homobasidiomycetae-aphyl- lophoracea. II. Amsterdam. p. 1-278. Ellis, J. B. and B. M. Everhart. (1889). Some new species of hymenomycetous fungi. Journ. Myc. 5: 28, pl. 8. Ellis, J. B. (1890). Catalogue of the Plants of New Jersey. p. 483. Faull, J. H. (1922). Forest pathology, in Rept. Minister Lands and Forests, Ont. 1921: 259-266. Finn, R. F. (1942). Mycorrhizal inoculation of soil of low fertility. Black Rock Forest Papers (N. Y.) 1 (19): 116-117. Fries, E. M. (1821). Systema Mycologicum. 1: 341, 351. ————. (1836-1838). Epicrisis systematis Mycologici, seu synopsis Hymeno- mycetum. p. 474. (1848). Fungi natalenses. p. 8. . (1851). Novae Symbolae Mycologicae, in Acta Reg. Soc. Sci. Upsala, 3 (1): 70-71. 1851. Vol. date 1855. . (1863). Monographia Hymenomycetum Suecia. 2: 268. Haddow, W. R. (1941). On the history and diagnosis of Polyporus tomentosus Fries, Polyporus circinatus Fries and Polyporus dualis Peck. Trans. Brit. Myc. Soc. 25 (2): 179-190. Haskell, R. J. and J. I. Wood. (1930). Diseases of plants in the United States in 1929. Plant Disease Reporter. Suppl. 75: 66. Hatch, A. B. (1937). The physical basis of mycotrophy in Pinus. Black Rock Forest Bull. No. 6: 1-68. Hay, G. U. (1903). New Brunswick Fungi. Bull. Nat. Hist. Soc. New Brunswick 5: 117. -. (1905). Additions to the plants of New Brunswick. Bull. Nat. Hist. Soc. New Brunswick, 5 (3): 363. Hennings, P. (1898) in Engler & Prantl. Die Natirlichen Pflanzenfamilien, BOR: 167. 1755 10s Hesler, L. R. (1929). A preliminary report on Polypores of Eastern Tennessee. Journ. Tenn. Acad. Sci, 4: 9. eS Toe ee Bee ee ee ee et eee. ee) a A ye Se ee ey at Se ee GOSSELIN: STUDIES ON POLysTICTUS CIRCINATUS 567 Hester, J. B., J. M. Blume, and F. A. Shelton. (1937). Rapid chemical tests for coastal plain soils. Virginia Truck Exp. Sta. Bull. 95: 1433-1486. Hubert, E. E. (1924). The diagnosis of decay in wood. Journ. Agr. Res. 29: sol, 567. : . (1929). A root and butt rot of conifers caused by Polyporus circinatus Fr. Phytopath. 19: 745-747. (1931). Outlines of Forest Pathology. p. 351. Jiead: I. and J. G. Juul. (1938). Ratesopper pa levende naletraer. (Rot fungi on living conifers). Meddelelser fra det Norske Skogforséksveseen. 6 (3): 300-496. Karsten, P. A. (1889). Finlands Basidsvampar. p. 326. Kauffman, C. H. (1911). Unreported Michigan Fungi for 1910. Rept. Mich. Acad. 13: 230. (1921). The mycological flora of the higher Rockies of Colorado. Papers Michigan Acad. 1: 119. Konrad, P. and A. Maublane. (1926). Icones selectae fungorum. 5: pl. 457. Lloyd, C. G. (1907). Letter No. 14: 5. - . (1908). Myc. Notes Polyporoid Issue 1: 2, fig. 195-199, oo CLO)». Letter Nov 303 I,-2! 6) 11. ——, (1912). Letter No. 38: 6; No. 39: 4; No. 42: 5; No. 43: 4. ———. (1913). Letter No. 44: 1; No. 45: 3; No. 47: 1. ————, (1914). Letter No. 49: 1, 5. ————. (1915). Letter No. 54: 1; No. 58: 1; No. 60: 1, 4, 7. ————. (1916). Letter No. 61: 1, 6; No. 62: 3; No. 63: 2, 8. ———. (1920). Myc. Notes 6: 933, 949, 953, 964, 978, 979, 1014, 1018. ———. (1921). Myc. Notes 6: 1083. ————. (1922). Myc. Notes 7: 1127, 1160. (1923). Myc. Notes 7: 1187, 1188. Rompvoat B. O. (1904). ae . | enna a : ail m5 bd x si a a ie Pe al ly ; ? ; a a ne % zt ‘ian — nee | Lee : 4 . seas & = | See eA sith Se = ieee i Cx + r ENGES say oat oa fo 1290. Figures 1-9 . 604, FarLowia, Vou. 1, 1944 2. Pezizella culmigena Sacc., Ann. Myc. 12: 292. 1914. — Sacc., Syll. Fung. 24°: 1189. 1928. FIGURES 6-9. Apothecia subgregarious, rather numerous, sessile, attached by a broad central portion or at least appearing so, free at the margin, yellow, about 0.6-1.1 mm. diam. when dry, not perceptibly larger when moistened, when dry fairly thin, plane to saucer-shaped, when moistened somewhat thicker, cushion-shaped; receptacle smooth, when dry turned slightly off the substratum at the margin, slightly olive-brown toward the base, pale yellow above, “Chamois” to “Honey Yellow,” on moistening re- maining nearly the same color, lying closely adjacent the substrate; margin circular, even, rarely slightly lobed or angular, obtuse, rather thin, when dry even with the hymenium or slightly elevated, when moistened directed outward and downward against the substratum, con- colorous with upper part of receptacle; hymenium when dry plane, waxy, concolorous with receptacle and margin, becoming slightly convex on moistening; in section 170-200 y» thick from substrate to hymenial surface; medullary region very compact, of indeterminate structure, con- taining a large amount of crystalline material; ectal layer distinct, well differentiated, about 30 ,« thick at the base, somewhat thinner toward the margin, composed of compact, more or less angular, thin-walled cells 9-12 » diam., colorless except for those in the outer layer which in the basal region of the apothecium are brown and have walls slightly thickened, those at the margin of the disc clavate; paraphyses simple, colorless, thin-walled, non-staining, slightly clavate at the apex, 3-4 p diam.; asei originating from croziers, narrow, clavate, 65-68 x 6-7 p; ascospores biseriate in the upper part of the ascus, uniseriate below, cuneate, rounded above, tapering to an obtuse point below, straight or slightly curved, the sides sometimes slightly irregular in outline, 8.5-10 x 2.42.6 gw, 1-celled, lacking oil globules. Known only from the type: On decaying culms of Phragmites communis Trin., Weisskirchen, Mahren, Bohemia, May 1914, F. Petrak (FH). Saccardo in his original description indicates that this species is close to Helotium citrinulum. Superficially the apothecia of the type (Fig. 6) differ but slightly from those of H. citrinulum (Fig. 1) in being less bright- ly colored than most specimens of the latter and in being slightly grayish on the lower portion of the receptacle. Microscopically the species is distinct from H. citrinulum in its wedge-shaped spores (Fig. 9), long and narrow asci (Fig. 8) and different structure (Fig. 7). The hyphae of the medullary excipulum do not stain well, the walls are thin and appear to become confluent, and the tissue contains a large amount of obscur- ing mineral matter (Fig. 7). It is not certain whether the structure is fundamentally prosenchymatous or pseudoparenchymatous. In contrast, the corresponding region in H. citrinulum (Fig. 2) is rather loose and open and the hyphae are distinct. Also in Pezizella culmigena (Fig. 7) the ectal layer is more clearly defined, more rind-like, and more com- pact than in Helotium citrinulum, and the outer layer of cells in the basal WHITE: StupIEs In Genus HELotTium. IV 605 portion of the apothecium is brown, whereas in H. citrinulum the entire tissue is practically colorless. It is possible that the two species are less closely allied than their superficial resemblances would at first indicate. 3. Helotium midlandensis sp. nov. FIGURES 10-13. Apotheciis sparsis vel subgregariis, plerumque solitariis, rarius trinis vel octonis, stipitatis, flavis vel aurantiacis, 0.3-0.7 mm. in diam. et eaque altitudine; stipite sat zracili, cylindrico, levi, pallide flavo; disco apiriente depressione minuta citoque ap- planata, subcarnoso, remanente solido in siccis; receptaculo stipiti concolori, in siccis minute-ericeo-striolato de margine usque ad apicem stipitis, levi in humidis; hymenio plano vel patelliformi, rarius leviter convexo, flavo vel aurantiaco; margine levi, obtuso, circulari vel angulato ex pressione marginum aggregatorum attingentium; paraphysi- bus, simplicibus, interdum unum ramum prope basim gerentibus, cylindraceis, 3-5 sep- tatis, 2.43.2 y. in diam.; ascis e “croziers” natis, clavatis, 45-65 x 6.5—-6 .; ascosporis bi- seriatim vel subbiseriatim dispositis, unicellulatis, plus minusve ovoideis, 7-10x2.6—-3.2 ys. Apothecia known only from dried specimens, scattered to subgregari- ous, usually solitary, more rarely in clusters of three to eight, stipitate, in general aspect yellow or orange-yellow, 0.3-0.7 mm. across the disc and of about the same height; stipe rather slender, cylindric or essen- tially so, smooth, pale yellow; dise opening by a minute depression, very soon plane, subfleshy, retaining its original fullness on drying; re- ceptacle concolorous with stipe, when dry showing fine, silky super- ficial striae radiating from top of stipe toward margin, smooth when moistened; hymenium plane to patelliform, more rarely slightly con- vex, yellow or orange-yellow; margin smooth. obtuse, circular, or when crowded becoming angular from mutual pressure; in section: stipe about 180 » thick, very compact, composed of narrow, parallel hyphae about 3 » diam. with narrow lumen and thickened, confluent walls, those near the surface turning outward and ending in a thin indefinite zone 12-18 p» thick and composed of isodiametric cells 3-6 » diam.; medullary region of disc fairly compact, of thin-walled hyphae 2.5-3.5 p», much branched and interwoven; ectal layer compact, 40-50 » thick, composed of narrow, parallel hyphae 3—4.5 » diam., except at or near the surface where a few are of somewhat larger diameter and the cells shorter; subhymenium not differentiated from the medullary layer; paraphyses simple or occasionally once-branched near the base, cylindric, 3—5-septate, 2.4—3.2 » diam.; asci originating from croziers, clavate, 45-65 x 6.5—7 p; ascospores biseriate or subbiseriate, |-celled, irregularly obovoid, 7-10 x 2.6-3.2 p. Habitat: One specimen (Iowa, Grumbein) on petioles and larger leaf veins of old leaves of Quercus sp.; all others on old pods of Gleditsia triacanthos L, Distribution and specimens examined: Ohio: Morgan 801, reported incorrectly (Morgan, Journ. Mycol. 8: 184, 1902; Seym., Host Index, p. 416. 1929) as Helotium discretum Karst. (NY-E).— Iowa: Homestead, Sept. 26, 1931, G. W. Martin 5186, type (FH).— Hills, Dec. 3, 1933, G. W. Martin 5191 (FH).— (With Typhula juncea), North Liberty, Oct. 4, 1941, M. L. Grumbein (FH).— Kansas: Lawrence, 1890, W. C. Stevens 59, incorrectly labeled Helotium herbarum (NY-E). 606 Fartowia, VoL. 1, 1944 This species has much the aspect of Helotium immutabile Fuckel and other members of the H. epiphyllum group (cf. White, Farlowia 1: 137, 139-147. 1943) to which it belongs. It is distinguished among these species by its small size, the comparatively slight parenchymatous differ- entiation of the ectal layer of the disc, small asci, and small egg-shaped spores. Compare the following species. 4. Helotium erraticum sp. nov. FicuRES 14-18. Apotheciis paucis, sparsis vel subgregariis, stipitatis, albis vel cremeo-flavis, usque ad 2.5 mm. latis et 1 mm. altis, siccando flavescentibus, cartilagineis; stipite crasso, plus minusve cylindraceo, levissimo, albo-hyalino, lutescente siccando; disco depres- sione minuta aperto, mox applanato, carnoso, levi, albo, siccando lutescente vel rube- scente; hymenio albo, plano, siccando lutescente vel rubescente, opaco vel plus minusve pellucido, cupuliformi; margine levi, obtuso, — Stipite consistente ex hyphis parallelis, densis, 2.5-5 y. crassis, tenuitunicatis, et ad basin et secundum superficiem expositam brevioribus et 4-10 y, crassis, membranis crassiusculis praeditis; excipulo ectali disci indefinito, admodum compacto, ad basin crasso, marginem versus tenuiore, ca. 40-50 p, crasso in parte media inter basin et marginem; regione medullari minus compacta; subhymenio vix distincto a medulla; paraphysibus longis, rectis, simplicibus vel rarissime semel furcatis prope basin, ad apicem clavatis, ascos longitudine paululum superantibus, 2.8-3.5 . crassis; ascis e “croziers” natis, clavatis, 90-120 x 9-11.5 wu; ascosporis biseriatim dispositis, subtus autem unam tantum seriem formantibus, ob- longis vel obovoideis, subcurvatis vel unum ad latus applanatis atque utrinque sub- attenuatis, interdum plus minusve crepiduliformibus, praecipue cum membrana asci iam compressae sunt, uni-cellularibus, 11-17 x 3.5-4 y, guttulis olei globulosis con- spicuis binis apicalibus instructis. Habitatio: Ad folia putridissima arborum frondosarum. Apothecia few, scattered or rarely subgregarious, stipitate; when fresh, white to creamy yellow, up to 2.5 mm. diameter and 1 mm. high, more often 0.5-1.0 mm. across the disc, and 0.6—0.8 mm. high; in drying, con- tracting slightly, becoming yellow or dull reddish yellow, “Chamois” to “Cinnamon-Rufous” or “Orange-Cinnamon,” cartilaginous and of more or less the same color and consistency throughout; stipe stout, cylin- dric or essentially so, entirely smooth, hyaline-white when fresh, drying translucent-yellow; dise opening by a minute depression, soon plane, fleshy, contracting slightly on drying, more or less saucer-shaped, re- maining thick; receptacle smooth, when fresh white, drying translucent- yellow below, similar toward the margin or of a brighter and more opaque yellow, or sometimes uniformly yellow or reddish; hymenium when fresh white or nearly so, plane, on drying becoming bright yellow Fig. 10-13, Helotium midlandensis: 10, Diagrammatic representation of sec- tion through apothecium, x 110, from type (FH); 11, Section through ectal ex- cipulum, Martin 5191, « 1290 (FH); 12, Asci and paraphyses, from type, x 1290 (FH); 13, Ascospores from type, x 1290 (FH). Fig. 14-18, Helotium erraticum: 14, Diagrammatic representation of section through apothecium, from type, x 78 (FH); 15, Section through ectal excipulum, from CU-P 27850 (FH); 16, Asci and paraphyses, from CU-P 29658, x 1290 (FH); 17, Ascospores, from CU-P 29658, « 1290 (FH); 18, Ascospores, from TRT /7611, _< 1290 (FH). en ea, > a | ca. WL. White del. Ficures 10-18 607 608 FartowiA, Vou. 1, 1944 to reddish, opaque or more or less translucent, saucer-shaped; margin even, obtuse, drying slightly elevated and very slightly undulate or wrinkled; in section: siipe composed of very compact, parallel hyphae, 2.5-5 pw diameter, thin-walled except at the base and along the exposed surface where the cells are shorter and thicker, 4-10 » diameter, with walls slightly thickened and confluent; ectal excitpulum of disc indefinite. very compact, thick at the base, thinner toward the margin, about 40-50 p thick half way between base and margin, the hyphae similar to those of the stipe, the outermost ones similarly somewhat larger in diameter and with walls slightly thickened, forming a thin rind layer extending nearly to the margin; medullary region slightly less compact, of thin- walled hyphae 2.5—-5 » diam., much branched and interwoven in all directions; swbhymenium scarcely differentiated from the medullary re- gion; paraphyses long, straight, unbranched, or very rarely once-branched close to the base, clavate at the apex, extending somewhat beyond the asci, 2.8-3.5 » diam.; asei originating from croziers, clavate, 90-120 x 9-11.5 4; ascospores biseriate above, uniseriate below, oblong or obo- void, slightly curved or flattened on one side, slightly pointed toward the ends, sometimes appearing more or less slipper-shaped especially when crowded in the ascus, 1-celled, 11-17 x 3.5—4 p», with a conspicuous and rather large oil globule in each end. Habitat: On much decayed leaves of various frondose trees (as listed below) ; one collection on old pods of Robinia Pseudoacacia L. Distribution and specimens examined: Quebee: On Acer sp., Populus sp., & sp. indet., Duchesnay, Aug. 25, 1938, H. H. Whetzel & T. Sproston, previously reported (Mycologia 31: 730. 1939) as Helotium phyllophilum and (White, Farlowia 1: 141, 159. 1943) as H. epiphyllum (CU-P 27850, FH).— Duchesnay, Aug. 26, 1938, R. F. Cain 11154, reported (Mycologia, l.c.) as H. immutabile and (White, ].c., p. 141, 147) as H. epiphyllum, spec. comm. H. S. Jackson (FH).— New York: On Acer sp., Coy Glen, near Ithaca, Oct. 3, 1938, H. H. Whetzel & W. L. White 3416 (FH).— On Acer sp. ?, Coy Glen, Oct. 3, 1938, Whetzel & White 3421, type (FH).— On Amel- anchier sp., Hamamelis sp., & Ulmus sp., Malloryville, near Ithaca, Oct. 18, 1941, H. H. Whetzel & J. Niederhauser (CU-P 29658, FH).— On pods of Robinia Pseudo- acacia L., Newfield Gorge, near Ithaca, Oct. 26, 1941, H. H. Whetzel & T. Sproston (CU-P 29666, FH).— Ontario: On Acer sp., Brewer Lake, Algonquin Park, Sept. 5, 1939, R. F. Cain, comm. Jackson, TRT 17611 (FH). This is another member of the Helotium epiphyllum group and must be cautiously separated from such species as H. epiphyllum, H. immuta- bile, H. carpinicola, and H. midlandensis. These species appear to be distinct and taxonomically satisfactory, but may be somewhat difficult for one who has had no experience with them. They cannot be sepa- rated with any degree of certainty macroscopically, but under the micro- scope are distinct on a number of characters, the final and most impor- tant being spore size and shape. 5. Helotium albidum (Rob. ex Desm.) Pat., Tab. Anal. Fung. 4: 173, fig. 382a-d.. 1885. Peziza albida Rob. ex Desm., Ann. Sci. Nat. 3: 16: 323. 1851. Helotium scutula (Pers. ex Fr.) Karst. var. albidum (Rob. ex Desm.) Karst., Myc. Fenn. I, Bidr. Kann. Finl. Nat. Folk 19: 112. 1871. WHITE: StTupIEs IN GENUS HELoTIUM. IV 609 Phialea albida (Rob. ex Desm.) Gill., Champ. France, Discom., -p. 105. 1879-1883. Hymenoscypha albida (Rob. ex Desm.) Phill., Man. Brit. Discom., p. 138. 1887. FIGURES 19-24. Apothecia arising from black areas l-several cm. long on the leaf petiole, 6-20 on each such area, scattered to subgregarious, stipitate, in the dried condition up to about 1.2 mm. high and 1.5 mm. across the disc, the larger ones expanding to 2 mm. when moistened, in the fresh condition creamy white or ivory white (Desm.) or with maximum mois- ture content more dilute in color, drying with stipe and outside of disc essentially concolorous, “Cinnamon” or “Pinkish Cinnamon,” and the hymenium “Orange-Cinnamon”; stipe cylindric, slender but not deli- cate, minutely scabrous when dry; disc subfleshy, spreading, saucer- shaped when dry; receptacle when dry finely and closely lined with superficial radiating striae; margin even, when dry conspicuously ele- vated above the hymenium; hymenium drying firmly waxy; in section: stipe composed of a broad central core, hyaline, of thin-walled hyphae about 4-6 » diam., and a thin, yellowish, cortical layer 4-5 cells thick and composed of short rectangular cells; medullary region of disc com- pact, composed of hyphae similar to those making up the central core of the stipe; hypothecium not noticeably differentiated; ectal layer not sharply distinguished, consisting of a zone about five cells thick made up of large hyphae 8-10 » or rarely up to 15 w diam., over which is a thin external covering of narrow more or less crushed or collapsed hyphae of smaller diameter; paraphyses simple, usually 3-septate, slightly en- larged toward the apex, about 3 » diam.; asei clavate-cylindric, not originating from croziers, 90-100 x 10-12 »; ascospores uniseriate below, biseriate above, 1-celled, 13-17 x 4—5 p, rounded above, very slightly tapering toward the lower end, slightly curved or flattened on one side, obtuse-pointed below, the content granular. On fallen, decaying petioles of Fraxinus excelsior in France. Known with certainty only from the material of Desmaziéres: Pl. Crypt. Fr., ed. 1, 2004; ed. 2, 1604 (FH). This is a member of the Helotium scutula complex in which many forms, varieties and species have been described. It is being allowed to stand as distinct from A. scutula or H. caudatum, the former on herba- ceous stems and the latter on leaves. It is more robust than H. caudatum and differs from both it and H. scutula in having spores which are smaller in size and more obtuse at the lower end. If it proves to be confined to Fraxinus petioles where it causes blackening, and if spore and other characters are constant, then it may be maintained as a species. The references listed above are only those pertinent to the synonymy. There are many European records, but they are based largely upon spcimens better referred to H. caudatum or H. scutula. 6. Helotium scutula (Pers. ex Fr.) Karst. var. fueatum (Phill.) in Rehm in Rabenh. Krypt.-Fl. 1°: 793. 1893. — Massee, Brit. Fung.-Fl. 4: 254, 1895. — Boud., Hist. Classif. Discom. Europe, p. 114. 1907. Peziza fucata Cooke & Phill.; Cooke, nom. nud., Grevillea 4: 132, pl. 65, fig. 300. 1876. 610 FarLowl1A, Vou. 1, 1944 Hymenoscypha scutula (Pers. ex Fr.) Phill. var. fucata Phill., Man. Brit. Discom., p. 137. 1887. Phialea scutula (Pers. ex Fr.) Gill. var. fucata (Phill.) Sacc., Syll. Fung. 8: 266. 1889.— Seaver, Bull. Lab. Nat. Hist. State Univ. Iowa 6°: 101. [Oct. 1911], spec. & descr. excluded. — Oud., Enum. Syst. Fung. 2: 1004, 1920. — Seym., Host Index, p. 292. 1929, based on Seaver’s report (l.c.). FIGURES 25-30. Apothecia scattered to subgregarious, fairly numerous, stipitate, hyaline-white when fresh, or sometimes yellowish-stained, 1-3 mm. high, up to 2.8 mm. across the disc though usually smaller, drying yellowish ochraceous, all parts approximately concolorous in both fresh and dried conditions, the disc contracting somewhat in drying, usually not over 1 mm. diam. in herbarium specimens; stipe slender, usually about equalling the diameter of the disc, originating underneath the epidermis of the substratum, scarcely noticeably erumpent, smooth, usually more or less broadened at juncture with disc; dise at first subglobose, open- ing by a pore, finally expanding, more or less saucer-shaped, of medium thickness; receptacle smooth when fresh, very minutely radiately fur- rowed when dry; hymenium becoming plane at maturity and with maximum moisture content, drying slightly to pronouncedly concave; margin thick, obtuse; in section: stipe with a rind about 30 » thick or about 5-6 cells thick, composed of longitudinally parallel hyphae 4-8 » diam. divided into rectangular cells 13-19 p» long with walls slightly thickened and confluent, this rind enclosing a large central core of parallel, compact hyphae 2.5-3.5 » diam. with walls slightly thick- ened, divided into cells 25-50 » long; disc composed of an ectal layer of hyphae similar to those of the stipe and a relatively reduced medul- lary region of more loosely interwoven hyphae; ectal layer about 50 p thick at the base, becoming thinner toward the margin, consisting of an outer zone similar to that of the stipe and a broader inner zone of radiately parallel hyphae similar to those of the central part of the stipe; paraphyses simple or frequently once or twice branched at or more often below the middle, usually near the base, 3—5-septate, slightly clavate at the apex and there 3-3.5 » diam.; asei originating from croziers which break rather easily, clavate, 118-135 x 12-15 »; ascospores biseriate, 1l-celled, obtuse and rounded above, flattened on one side or slightly curved, narrowed downward, acute at the lower end, 24-34 x 5—6.8 , each end beset with 1 to several small, inconspicuous cilium- like processes, content with numerous conspicuous refractive granules. Known from only three collections: New York: On old stems of Polygonum robustius (Small) Fernald (det. Fernald) lying in film of water in swamp, Cayuta Fig. 19-24, Helotium albidum, all from type material, Desm. Pl. Crypt. Fr., Ed. I, 2004 (FH): 19, Diagrammatic representation of section through an apothecium, < 40; 20, Section showing blackening of the exposed cell walls over the area of the petiole on which the apothecia are seated, the epidermis of the petiole having earlier sloughed off, « 1290; 21, Section showing structure of outermost tissue of stipe, exposed surface at left, taken from near but not at the base, x 1290; 22, Section showing structure of disc tissue, < 1290; 23, Asci and paraphyses, x 1290; 24, Ascospores, X 1290. wre, Reape Ne ae ts W.L. White del. Ficures 19-24 FarLowiaA, VoL. 1, 1944 612 Ficures 25-30 WHITE: STUDIES IN GENUS HELoTIUM. IV 613 Lake, Sept. 1, 1936, H. H. Whetzel & W. L. White (CU-P 25494, FH, US).— Same substrate and locality, Sept. 9, 1938, Whetzel, White, et al (CU-P 27863, FH, US).— England: “On dead stems of Polygonum lying in water.”, Shrewsbury, Phill. Elv. Brit. 120, type (CU-PD 11122, FH). It would be scarcely possible for three collections to agree more com- pletely than do those cited above. Although the substratum for the British specimen is insufficient for complete identification, there is no doubt about its being Polygonum, and the pieces of stem in the packets agree so closely with those of the American collections that one is tempted to guess that they represent a closely allied species. The evi- dence is that this fungus does exist as a distinct entity within the taxo- nomically difficult Helotium scutula complex of which it is a member. Its distinguishing characteristics are the presence of croziers at the bases of the asci, relatively large asci, large and robust spores, and occurrence on decaying, water-saturated stems of Polygonum. Though this variety may prove to be confined to Polygonum, this does not mean, however, that all members of the Helotium scutula group found on Polygonum should be referred here. On the contrary several collec- tions are at hand where the substratum is Polygonum and the fungi are typical Helotium scutula. A specimen on Polygonum from Iowa (CU-PD 553), reported by Seaver under the varietal name, represents typical material of the species. Cultures were made from both of the American collections, using mass ascospore discharge on potato dextrose agar. The spores germinated readily, becoming 1-3 septate in the process, and developing a mat of sterile mycelium. The cultures were maintained only in test tubes at room temperature, but there developed no conidial stage, spermatia, stromata or other. structures of taxonomic value. 7. Helotium Dearnessii (Ell. & Ev.) White, Mycologia 34: 167. 1942, Peziza (Phialea) Dearnessii Ell. & Ev. nom. nud., N. Am. Fungi, second series 2624. 1891. Phialea Dearnessii Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 146. [Feb. 28] 1893. — Sacc., Syll. Fung. 11: 403. 1895. — Seym., Host Index, p. 558, (559 from error in host det.). 1929. — J. H. Miller, Pl. Dis. Rep. Suppl. 131: 47, Aug. 15, 1941, based on an incorrect determination. Hymenoscypha Dearnessii (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 37: 485. 1898. FIGURES 3]—34. Apothecia scattered to subgregarious, not crowded, appearing sessile or nearly so, in the dried condition ochraceous, all parts concolorous 0.3-0.7 mm. diam. across the disc, when moistened becoming more yel- Fig. 25-30, Helotium scutula var. fucata, all except fig. 30 from Phill. Elv. Brit. 120, type (FH): 25, Diagrammatic representation of section through apothecium, < 76; 26, Section showing structure of outermost tissue of stipe, x 1290; 27, Section showing structure of outermost tissue of disc, X 1290; 28, Asci and para- physes, x 1290; 29, Ascospores, 1290; 30, Ascospores germinated on potato dextrose agar, slightly more than x 1290 (CU-PD 25494). 614 Fartowia, Vor. 1, 1944 — lowish the largest reaching as much as 1.0 mm.; stipe obscure, scarcely long enough to raise the central part of the disc off the substratum, rarely longer, ochraceous, rarely darker at the base; dise rather fleshy, saucer-shaped when dry, becoming cushion-shaped when moistened; re- ceptacle smooth; hymenium opening by a minute depression, soon plane, remaining plane or becoming somewhat concave on drying, con- vex when moistened; margin even, slightly elevated when dry, when moistened expanded and recurved, wedge-shaped; in section: stipe originating underneath the cortex of the host, erumpent, cylindric, usually protruding only slightly beyond the cortex, very compact, homogeneous; ectal layer of disc thick, indefinite, in general composed of an inner layer originating at the top of the stipe, about 40 » thick at the base, tapering towards the margin, colorless, composed of compact parallel hyphae about 3 » diam. with walls scarcely or not at all thick- ened, these hyphae turning obliquely outward to form a thicker outer brownish zone and there becoming somewhat larger, shorter-celled, with walls thickened and confluent; medullary region thin, rather loose, the hyphae rather narrow, thin-walled, interwoven, becoming erect towards the hymenium; hypothecium not differentiated from medullary layer; paraphyses mostly simple, rarely once-branched below the middle, gradually enlarged from base to apex, about 3-3.5 » diam. at apex; asci not originating from croziers, clavate, 100-108 x 9-13 ,; asco- spores biseriate, elongate, pointed at the ends, attenuate below, 1-celled, containing a row of 2-9 oil globules, with a delicate cilium 2.5-3.5 p long at the lower end, sometimes also at the upper end, 30-35 x 44.5 not including the cilia. Known only from the following collections: Quebec: On old herbaceous dicotyle- donous stems, Aug. 25, 1938, J. VW. Groves (FH, OTB).— Ontario: On old stems of Steironema ciliatum (L.) Raf., London, May [24], 1890, J. Dearness 1713, type (FH-E).— Additional material from same place, June [13], 1890, J. Dearness 1713, Ell. & Ev. N. Am. Fungi, 2nd ser. 2624 (FH). According to a letter from Dearness to Farlow, he first collected the species May 24, 1890 and recorded the substrate as Monarda didyma. Ellis described it as on Monarda and it was so recorded in Seymour’s Host Index. Dearness apparently returned to the same place June 13, 1890, collected additional material which he sent Ellis for his N. Am. Fungi, and decided that the host was Steironema ciliatum. A very ample portion of the June 13th material, in addition to the portion in N. Am. Fungi, was sent to Farlow and is now in the Farlow Herbarium. Fig. 31-34, Helotium Dearnessii, all from Dearness, 1713, coll. June 1890 (FH) ; 31, Diagrammatic representation of section through apothecium, x 64; 32, Section showing details of structure of ectal excipulum, « 1290; 33, Asci and paraphyses, > 1290; 34, Ascospores, < 1290. Fig. 35-40, Helotium gemmarun, all from CU-P 251781 (FH): 35, Diagram- matic representation of section through an apothecium, * 60; 36, Section showing structure of outer layer of stipe, < 1290; 37, Section showing structure of medullary and ectal layers of disc, x 1290; 38, Hyphal tips from margin of disc, « 1290; 39, Asci and paraphyses, * 1290; 40, Ascospores, X 1290. FIGuRES Barr ey ieee WL. White del. a | 31-40 616 FarLtowiA, VoL. 1, 1944 The collection recorded by Miller (1. c.) from Georgia on Stetronema ciliatum, of which he kindly loaned me a fragment, proves to be the much more common and widespread species, Helotium scutula (Pers. ex Fr.) Karst., to which H. Dearnessii is surely closely related but clearly distinct in its long, attenuate spores. The two specimens of Helotium Dearnessii recorded above agree closely in their characters except that in the Ontario material the apothecia are - all so short-stipitate that on superficial examination they appear sessile, whereas in the Quebec specimen most of them are distinctly stipitate; also the cilia, which are difficult to see in the former are rather distinct in the latter. 6. Helotium gemmarum Boud., Bull. Soc. Myc. France 4: 81, pl. 17, fig. 2. 1888. — Boud., Rev. Myc. 11: 167. 1889.— Boud., Icones Mycol. 3: pl. 493. 1905- 1910. — Boud., Hist. Classif. Discom. Europe, p. 112. 1907. Phialea gemmarum (Boud.) Sacc., Syll. Fung. 8: 271. 1889. — Oud., Enum. Syst. Fung. 2: 61. 1920. — J. H. Miller, Pl. Dis. Rep. Suppl. 131: 47, 1941. FIGURES 35-40. Apothecia scattered to subgregarious, small, delicate, slender, stipitate, white, in the fresh condition up to 1.5 mm. high (up to 5.0 mm. fide Boud.) and 0.4-1.4 mm. across the disc; stipe slender, slightly en- larged upward, when fresh hyaline to dilute white, smooth at the base, minutely puberulent above, not changing perceptibly on drying; dise opening by a pore, at first infundibuliform, at length expanding, finally plane, thin, on drying quickly contracting to become more or less funnel- shaped with margin turned inward; receptacle minutely and sparsely puberulent, dilute white when fresh, remaining so on drying or vary- ing towards dingy cream-color; hymenium always about same color as receptacle, flat at full maturity and with maximum moisture content, otherwise more or less deeply concave; margin minutely puberulent, even with the hymenium in fresh, mature specimens, erect or turned in- ward in dried material; paraphyses not numerous, simple or more rarely once-branched below the middle, not enlarged at the apex; asci small, cylindric, clavate, 45-50 x 5-7 »; ascospores small, obpiriform, 6-9 x 2.5-3 py. Habitat: Appearing in early spring on the scales of buds of various species of Populus (listed below) which had fallen the previous season and lodged under and among debris on wet ground. Distribution and specimens examined: New York: On P. candicans Ait., Lloyd Preserve, McLean, Apr. 26, 1936, H. H. Whetzel, D. H. Linder, & W. L. White (CU-P 25181, FH).— On P. candicans, Lloyd Preserve, McLean, May 2, 1937, W. L. White et al (FH). — On P. candicans, Lloyd Preserve, McLean, May 18, 1940, H. H. Whetzel, W. L. White, & J. Niederhauser (CU-P 29086, FH).— Pennsylvania: On Populus sp. ?, May 5, 1935, H. H. Whetzel (FH).— Georgia: On P. deltoides Marsh., Agri- cultural Campus, Athens, Apr. 29, 1940, J. H. Miller (GA 583).— France: Type locality. Boudier indicated that it was frequent on the old bud scales of Populus nigra L. in the vicinity of Paris in February. Notes and drawings by Massee, made from Boudier material, are in the New York Botanical Garden but are not accompanied by a specimen. ee Be ek es ee Eee fe. See sae gs Ay RM ee PRS? Sree eT eee ak eke eee ee %,- { Ree Ln als 3 » a — 5 lial WHITE: Stupies 1n Genus HELotium. IV 617 There appears to be no European material of this species available in the herbaria of this country. However, it is well marked as to morphological characters, substratum, and season of occurrence. Com- parison of North American specimens with the illustrations of Boudier leaves no reason for doubt as to its identity. Two other species with similar characteristics — white, slender, with small spores, occurring in the spring in moist habitats — and possibly more or less closely related, are Phialea Cassandrae Kanouse, occurring in Michigan on the leaves of Chamaedaphne calyculata (L.) Moench (see White, Farlowia 1: 159. 1943), and Helotium amenti (Batsch ex Fr.) Fuckel (see Boudier, Icon. Mycol., pl. 496. 1905-10), of widespread occurrence on female catkins of Salix sp. Harvarp UNIVERSITY _ CAMBRIDGE, MASSACHUSETTS Pe sales a ae, ‘ oe : — ; oa : % coe Age Fa Tee oe cee ive ian panies! he steerer ace Perera, ae ate bre eu We oe : fa rae Be rote z te Vine = : ; ea) ‘a Pee 3 ioe es is ‘ ‘Ai by a ue we ne i iy ae mi 2 : “tg ay 3 2 — a sa oe a oa ee ae aa ae at saat up RS cal, dan slot a ee gy ee 4 te Me eee ‘Be ri tf ala brn ana = Ht FS Tay ea eee tr oem oe 1(4): 619-624 FARLOWIA July, 1944 REVIEWS UEBER DIE BEDINGUNGEN DER MYKORRHIZABILDUNG BEI KIEFER UND FICHTE. By Erik Bjorkman. Uppsala. Symbolae Botanicae Upsaliensis. Vol. VI, no. 2. 1942, When B. Frank (1885) likened the common fungus root of a tree to a factory op- erating for its own profit, he was not unmindful that the tangled and swollen root tips of a tree with their covering mantle of fungous skin, and their cakes of mycelia between the cells, look like witches brooms or other gall formations caused by a parasitic fungus. He considered that the fungus obtained its food from the host tree and in return provided the tree with water and nutrients for growth, which the fungus extracted from the soil through its widespread mycelial threads. This inter- pretation did justice to what one can see, and itis supported by much else: many nutrients are deficient in the raw forest humus (mar) (Stahl 1900); the fungus root is built like an organ for the exchange of substances (Mangin 1910); Neger (1913) found more sugar in roots from trees which habitually have fungus roots than in the case of ash and many other plants; Rexhausen (1920) found plenty of fungous starch (glycogen) in the mycelium of root fungi and more sugar or starch in in- fected roots than in other roots of the same tree; a number of large fungi of the forest known to form mycorrhiza (Melin 1922-25 et al) follow their host trees as a “dolphin follows a ship.” Lange (1923), also Thesleff (1919), Romell (1921) and Melin (1925) did not admit that the mycorrhizal agency can consume cellulose or the more common energy-giving foods, except sugar; mycorrhizae can be abundant in almost completely humus-free soil, such as volcanic ash (McDougall and Jacob 1927); soluble dyestuffs can penetrate the fungus membrane and mycorrhizal net right into the cells of the host tree (Endrigkeit 1937); conifer seedlings without or almost without mycorrhizae have shown a lower phosphorus content and most of them also a lower nitrogen and calcium content than plants with abundant mycorrhizae grown in the same soil (Hatch et al 1937). Frank soon abandoned his first explanation of fungus-infected tree roots. Led astray by the peculiar pine vegetation and by some experiments with tree seedlings which he could not explain correctly, he concluded that the fungus gets no nourish- ment from the root, but like other soil fungi lives on humus, and only lives in the mycorrhizal covering in return for an ample rental, especially of nitrogenous sub- stances. This interpretation was adopted more or less faithfully by most of those who wished to believe that mycorrhizae are useful to the tree. Fungi came to appear as inseparable thralls of the host plant, and one cared little about what pleasure they could get from being tenants in the roots. Nor did Stahl care either, although his explanation of fungus-infected roots of vegetables, after another manner, came near to the old theory of Frank. Quite generally one has hesitated to believe that fungi that do not penetrate into the root cells and there become digested, can do the tree any other service than common soil fungi do by forming useful substances or consuming harmful ones. Some have wished to attribute the benefits of root fungi to this (Tubeuf 1896, Rayner 1934, Burges 1936, and Lindquist 1939) but it leaves the formation of mycorrhizae unexplained, as Endrigkeit (1937) and Hatch (1937) have pointed out. Others have not been able to escape the thought that the tree’s mycorrhizal builders live on their hosts. It used to awaken doubt that the mycorrhiza can be of any use to the tree, since biologists have become increasingly unwilling to conceive of any peaceful exchange to mutual advantage outside the realm of human beings. Frank’s Danish pupil Sarauw (1893) and Stahl’s pupil Weyland (1912) could believe no more than R. Hartig, Ramann McDougall (up to 1927) and others, that the mycorrhizal partner of trees is anything more than a harmless sponger. Melin tried in 1925 to unite the current opinions by sifting out differences between | other kinds of mycorrhizae and the common tree mycorrhizae. He had to class fungi as parasites or saprophytes, but showed that they invade root cells more frequently 619 rrre rane 620 FarLtowliA, Vou. 1, 1944 than generally appears, and there they are consumed. Thereby the trees, in exchange for growth substances and carbohydrates, receive nitrogenous foods rich in carbon which the fungi have formed from substances in the raw humus that are resistant to decomposition. Thus neither of the partners needs to get the short end of the bargain, even if one calculates the calories exchanged. The theory failed of proof, for, with culture in mir, conifer seedlings with their mycorrhizae showed that they required ordinary nitrogenous nutrients; they could therefore not live on the nitro- genous staples of the mir (Hesselman and Melin 1927). In addition, Hatch (1937) found that seedlings get along just as well without fungi, with both the “more com- plicated” nitrogenous compounds (peptone and nucleic acid) which Melin had utilized in the experiment on which he had based his conclusions. Hatch in 1937 forgot what Melin’s experiment with different carbon sources had shown, and suggested that the mycorrhiza-forming fungi were consumed with small amounts of growth substances, “and perhaps sometimes carbohydrates”. Endrigkeit the same year expressed the opinion that the mycelial net in the fully-developed mycorrhiza cannot obtain any nourishment from the host plant. The reviewer, on the other hand (Romell 1939) held that there was an exchange of substances ac- cording to Frank’s first belief, and supported this hypothesis with new reasons: a fungus does active duty in place of roots and root hairs and does not need to draw on the plant more than the ordinary root system would do. In order to be able to extract nutrients for growth from the soil as efficiently as possible in competition with all other fungi, the mycorrhizal fungi ought to have some kind of monopoly on easily digested energy-yielding food. Otherwise there seems to be no one since Rexhausen (1920) who has given Frank’s 1885 hypothesis any clear support. But here comes a young Swedish investigator and rejuvenates in new and im- proved form the very oldest theory for tree mycorrhiza as a useful structure. He does this by solving a riddle which has baffled investigators the world over during a gen- eration. The mystery for which Bjérkman has sought the solution is the peculiar variation in abundance of mycorrhizae. They are commonly found abundantly in moderately good and better mar humus, but sparingly in the most infertile mar and often also in mull, where they sometimes are absent. Recently only Hatch (1937) seems to have interpreted the fact, as far as it now goes, after the classical manner, namely that mycorrhizae are formed when the trees need them because they do not have enough mineral nutrients. This is no factual proof; Melin (1922-27) tried to obtain one and suggested that root fungi did not thrive in pronounced, but only in slightly acid mull (an unsupported supposition) and had insufficient “virulence” in bad mar. The word virulence is borrowed from medical science, literally meaning poisonousness, and connotes capacity to attack. But this power to attack is supposed to be explained by the word virulence. Of such a contorted phraseology Heribert-Nilsson says: “Such an explanation is as scant an aid as if a physician should stand up and declare that the cause of infantile paralysis lies in the susceptibility of man for infantile paralysis.” The word virulence is yet not altogether empty of meaning. Back of it lies the thought that the tree root and fungus are in a feud with each other. But the thought carried at once difficulties. Melin (1922) first supposed, as was natural, and as later McDougall did, that the fungus must be more virulent in order to penetrate the cells than to grow between them. The next year he had to contradict this and suppose something as strange as that a less virulent fungus goes between the cells, but a more virulent one can be forced by the host plant to hold itself in the cell walls. Melin’s specu- lative fabric has nevertheless appeared to be the best to be had, and M. C. Rayner has gratefully elaborated it further. Bjérkman sought first a foundation by collecting statistics on the abundance of mycorrhizae in different forest types. This yielded him as little as his less diligent predecessors had found, as far as providing a key or even any clue, and the fifty pages which deal with this serve best some academic requirement. Inoculated pure REVIEWS 621 cultures repaid his efforts better, but still the work would have had almost the same value without them. The scientific importance of the study lies partly in the ecologically planned experiments where uninoculated pine or spruce seedlings were grown in the same soil (rich mull, better and poorer mar and peat) with their natural content of soil life, or at least of sources of life, and partly in the theory to which the experiments led. It shows that Bjoérkman’s discovery is not a result of modern technique and the work of mycorrhiza investigators of recent times, but could have been made fifty years ago if someone then had been equally industrious and had had the vital spark. Six soils were tested in the experiment, some without additions, others under different degrees of illumination up to four soils with different additions of nutrients, and three soils under varying light and additions of nitrogen and phosphorus fer- tilizers. -Mycorrhizae were formed abundantly even in rich beech forest mull, after a moderate mixture of sawdust which decreased the nitrogenous food content. It is, therefore, not as Melin believed, that no mycorrhizae thrive in well-developed mull. Mycorrhizae are formed abundantly also in the soils showing the highest nitrogen content (far higher than are found in nature in the richest mull) if only phosphorus is low and light is strong enough. Lindquist (1932) also was mistaken when he thought that nitrate formation is to be looked upon as a “change in dominance” in the fungous flora of the soil of such a kind that mycorrhizal formation is thereby retarded. During the entire course of the experiments, three things were decisive: light, and the nitrogen and phosphorus contents of the soil. Thus mycorrhizae were formed in weak light (1/16 or sometimes 1% of full sunlight). With increased light there were steadily better seedlings with more mycorrhizae. On the contrary there were most mycorrhizae (but not the best seedlings) with moderate deficiency of nitrogen and phosphorus or both. Greater deficiency of nitrogen resulted in weak seedlings with few or no mycorrhizae. The same results were obtained in inoculated pure cultures when there was great deficiency of phosphorus. In soils richly fertilized with nitrogen and phosphorus the plants grew strongest, but had no mycorrhizae. The interaction between light and contents of nitrogen and phosphorus led to intricate correlations which are shown clearly in the report by means of three-dimensional curves. The action of light was known before, as a result of another study by Bjérkman (1940) together with Hesselman. The per cent of mycorrhizae in connection with light gives curves which appear logarithmic and call to mind Stalfelt’s photosynthesis curves for pine and spruce. Bjorkman guessed even in 1940 that there might be a connection between assimilation and mycorrhizal formation. This Melin (1927) had rejected as a possible cause for the stronger plants in Hesselman’s experiment having more abundant mycorrhizae; but he did not find the interpretation worth discussing seriously. Out of the guesses of 1940 eventually resulted the following: the mycorrhizae seek soluble carbohydrates in the root, and they consume them for energy, but they find them only if there is a surplus of carbohydrate in the root. Whether or not there is any surplus depends on the head-start photosynthesis (carbohydrate formation) has attained in the plant before the formation of albuminous substances. This ad- vantage will be small or nil if there is an abundance of all plant foods including nitrogen and phosphorus, so- that there will not need to be a shortage of materials when albuminous substances are built up. Then there will be little for the fungi to seek after in the root, and mycorrhizae will be formed sparingly or not at all. On the other hand, should there be a deficiency of nitrogen or phosphorus foods, there can easily arise a surplus of carbohydrates so that the formation of albuminous sub- stances falls behind photosynthesis. With a moderate lack of nitrogen or phosphorus or both, mycorrhizae can form abundantly, but if there is a serious lack of nitrogen or phosphorus, sooner or later photosynthesis will become weak also (phosphorus especially strongly influences carbohydrate formation in plants) so that the surplus of assimilated carbohydrate becomes less for that reason, and mycorrhizae are formed more rarely. Similar reactions occur if light becomes weak. Mycorrhizal formation ee we ee pee 622 FarLtowia, Vou. 1, 1944 is, expressed briefly, a result of and a sign that there is a certain surplus of energy- giving nourishment in the host plant. Bjorkman’s theory is this simple, but it is confirmed not only by his own involved curves, but also by what is otherwise known. A direct support for it is the higher sugar content that Rexhausen found in mycorrhizal roots. Bjorkman himself has found a still clearer direct support for his theory. In two experimental series he determined the amount of “reducing substance” in the roots of seedlings (in plain language — simple kinds of sugar; also many other substances according to Bjork- man). He who takes time to plot the data against the percentages of mycorrhizae (which Bjorkman should have done himself) finds a beautiful correlation. It is especially noteworthy that the relationship is so similar in two different soils. However, not even such a prop for a theory is any final proof of it, and questions remain. How does the fungus know that there is surplus sugar in the root? Does it go and feel here and there, or does the root give off sugar or something else tempt- ing when assimilation begins to satiate it? Still there does not seem to be room for any major error in Bjorkman’s theory. It is clear, although Bjérkman has not wished to say so, that Melin’s virulence theory has a reversed picture of reality. Whether mycorrhizae are formed or not depends more on the host plant than on the fungus, rather than as Melin and others have believed. The old scoffing belief that mycorrhizae are formed when the plant needs them came much nearer the truth, for a surplus of energy-giving food, which actually is the deciding factor, is conditioned in turn by a deficiency of mineral nutrients (c/N ratio). Among recent investigators Hatch came nearest the truth (as Bjorkman mentions rather cursorily) by his hypothesis that mycorrhizae are formed when there is too low a content in the plant of one or another of a number of nutrient substances. Bjorkman’s work throws new light not only on the problem with which he has been especially occupied. The solution he found to this question falls at once as a judgment of Solomon on the old strife between the “nitrogen theory” and “salt theory” of the ecology of mycorrhiza. It shows both theories to be more or less correct, but erects in their places a new, simple and beautiful theory, with an inner strength which no other mycorrhizal theory has had, for it gives an actual well- grounded explanation which appears axiomatic when it has once been clearly apprehended. The new theory should be called the Frank-Bjérkman, for it is a deepened and expanded form of Frank’s old one. According to it the mycorrhiza comes more than ever to appear like a factory for mutual profit. False must be called the whole view- point now for some time considered the only scientific one the world round: that altogether too human fable of the brutal burglar or the company formed by two gangsters who steal from each other. The exchange of substances in the mycorrhiza appears instead as a peaceful exchange where one party gains from the other’s un- necessary surplus, by offering goods he lacks in his activity, or if one will, by hiring and paying an industrious servant. Probably never before has anyone shown so clearly how the ways of nature can lead to a symbiosis between plants to their mutual advantage, and how this living together in many cases must inevitably take on this form just because an exchange serves its needs. Bjorkman’s work is destined to make a deep imprint on biological thinking in due time. His theory may furnish the key to the mystery of the symbiosis of alder and its nodule former, and perhaps also that of legumes and their root bacteria. A theory of such breadth as Bjérkman’s raises the question of exactly how it came about. One can read that it grew out of the hypothesis that the organisms forming mycorrhizae live on their host trees, and from Bjérkman’s interpretation of 1940 based upon it, of the effect of light upon mycorrhizal formation. However it is far from there to the complete theory. One thing and another suggest that Bjorkman almost brought out his work without its real point. Probably he came upon the solution when he read in the literature about the influence of phosphorus on the sugar con- REVIEWS 623 tent of beets and other plants, and about nodule formation on roots of legumes where the carbon-nitrogen ratio is at present considered to be the controlling factor. In such cases it is proof that there can be a certain value to the learning they are wont to have in Swedish doctorate dissertations. It in no wise follows from the above, however, that Bjérkman got his theory without effort, or even that almost anyone could have hit upon the solution by reading the literature. Others before this have brushed lightly past the key Bjorkman found, but have not seen it. If spruce seedlings die out on mull rich in nitrate, then it cannot reasonably be attributed to their failure to get the mycorrhizae they need (Lindquist 1932). When they do not have mycorrhizae it is on the contrary a proof that they do not lack any nutrient that is consumed in further elaboration of the first formed carbohydrates, and it is probably because they lack light. Black mycorrhizae are no sign that the soil is sick or cold so that a cutting must ~be made (Lindquist 1937). It indicates only weak light, that is, the stand is un- usually dense for northern Sweden. It must always remain a difficult task to judge the soil by the mycorrhizae, since poor mycorrhizal formation can depend either upon high nitrogen and phosphorus nutrition, or too little of either or both, or too scant light. Now perhaps it can be done in certain cases, thanks to what we know through Bjorkman. — Lars-Gunnar Romell in Svenska Skogsvardsféreningens Tidskrift. Haft IV. 1942, [Translated by H. I. Baldwin] RECHERCHES SUR LES CERAMIACEES DE LA MEDITERRANEE OCCIDEN- | ~ TALE. By G. Feldmann-Mazoyer. Algiers, Imprimerie Minerva, 1940 (1941). 510 pp. 191 text figs. 4 pls. In this important memoir, Mme. Feldmann-Mazoyer has presented the results of many years of research prosecuted in the Laboratory of Botany of the Faculty of Sciences of Algiers and in the marine biological stations of the Faculty of Sciences of Paris at Banyuls and Villefranche. The researches of the author deal with: 1) the cytology of the Ceramiaceae, 2) the morphology of their vegetative structures, 3) their reproduction, and 4) their tax- onomy. In these four series of researches, the author has obtained new results and has clarified questions which until now have remained obscure. In the parts devoted to cytology, the membrane, the nucleus, the cytoplasm and its elements, and the products elaborated by the cell (special floridean starch, os- mophilic granulations, basophilic bodies, . ..) are studied successively. The author makes many new contributions to the knowledge of the floridean cell, notably as it concerns the structure of the membrane, the morphology and the variations of the plastids, the structure and metabolism of floridean starch, etc. She has particularly studied the secretory cells, “bromuques,” which according to Sauvageau contain free bromine. By.new microchemical methods she has been able to establish the absence of free bromine in these cells. Also, the study of intercellular relations has permitted her to uncover in the axial cells, structures which are active in the translocation of dissolved nutrient substances, the structures playing the réle that had been attributed to the cells themselves. She has been able to confirm experimentally the role at- ‘tributed to. the synapses in these phenomena. In the portion devoted to the morphology of the vegetative structures, the author has been able to clarify structures heretofore poorly known. She has shown in par- ticular that the verticillate epidermis of species of Griffithsia correspond to the trich- oblasts of the Rhodomelaceae and has demonstrated their function as an absorp- tive layer. In the part treating with reproduction, the author has been able to describe the development of the reproductive organs in certain genera and species in which this was poorly known or even completely unknown. The systematic part constitutes a veritable monograph of the Ceramiaceae of the western Mediterranean. Using the results of researches enumerated above, the author 624 FarLowi4, Vou. 1, 1944 has ben able to establish a more rational classification of the family based principally on the structure of the procarp, in part on the manner of development of the gonim- oblast, and in part on the cytological structure. She has established three new tribes and modified the limits of many others, has created three new genera and suppressed four. She has described some new species and on the contrary has suppressed a large number, the study of very abundant living material having permitted her to determine that these so-called species represented only seasonal or ecologic variations. In each genus, very carefully prepared dichotomous keys permit the determination of the species which are later described in detail. Numerous illustrations facilitate further and render more certain this determination. The work of Mme. Feldmann will remain the basis of all studies of the Ceramiaceae, not only in the western Mediterranean, but of the entire world. Its great merit is that it was founded not only on herbarium material, but more especially on living plants, and that it has correlated, thanks to this fact, the numerous physiological and cytological characters with the purely morphological concepts of the earlier algologists. —R. Maire [Translated by D.H.L.] 1(4) 3 625-641 FARLOWIA July, 1944 INDEX TO VOLUME 1 INDEX TO AUTHORS AND TITLES Baker, E. E., E. M. Mrak and C. E. Smith. The morphology, taxonomy, and distribution of Coccidioides im- mitis Rixford and Gilchrist 1896, 199— 244. Barghoorn, E. S. and David H. Lin- der. Marine fungi: their taxonomy and biology, 395-467. Bartram, Edwin B.. Additions to the moss flora of northern Chile, 191-194; Burma mosses, 171-189; Mosses of Papua, New Guinea, 41-47; New and noteworthy Philippine mosses, 503-513. Chapman, V. J. The aims of future research in the algae, 5-8. Dixon, H. N. Alpine mosses from New Guinea, 25-40. Gosselin, Roger. Studies on Polystictus circinatus and its relation to butt-rot of spruce, 525-568. Greene, H. C. Notes on Wisconsin parasitic fungi, IV, 569-581. Herre, Albert W. C. T. A_ neglected field of study with the description of a new lichen from California, 391-393. Jackson, H. S. see Rogers, D. P. and H. S. Jackson. Khanna, L. P. On two new species of Megaceros with notes on M. arachnoi- deus, M. denticulatus, M. giganteus, and M. grandis, 515-523. Linder, David H. Foreword, 1-3; The genera Kickxella, Martensella, and Coemansia, 49-77; see also Barg- hoorn, E. S. and D. H. Linder. May, Valerie. Marine phytogeography, 491-493. Merrill, Elmer D. An index to Rafi- nesque’s published technical names for the cellular cryptogams, 245-262. Mrak, E. M. see Baker, E. E., E. M. Mrak and C. E. Smith. Nickerson, Walter J. Studies in the genus Zygosaccharomyces. I, Transfer 625 of pellicle-forming yeasts to Zygopich- ia, 469-481. Papenfuss, George F. Notes on algal nomenclature. III. Miscellaneous spe- cies of Chlorophyceae, Phaeophyceae and Rhodophyceae, 337-346. Prescott, G. W. New species and va- rieties of Wisconsin algae, 347-385. Prince, Alton E. Basidium formation and spore discharge in Gymnosporan- gium nidus-avis, 79-93. Rogers, D. P. The genus Pellicularia (Thelephoraceae), 95-118. Rogers, D. P. and H. S. Jackson. Notes on the synonymy of some North American Thelephoraceae and _ other resupinates, 263-336. Seeler, Edgar V., Jr. colous fungi, 119-133. Smith, C. E. see Baker, E. E., E. M. Mrak and C. E. Smith. Smith, Gilbert M. Microaplanospores of Vaucheria, 387-389. Svedelius, Nils. Galaxaura, a diplobi- ontic floridean genus within the order Nemalionales, 495-499. Whelden, Roy M. Notes on New Eng- land algae III. Some interesting algae from Maine, 9-23. Whetzel, H. H. A new genus of the Sclerotiniaceae, 483-488. Whiffen, Alma J. A discussion of tax- onomic criteria in the Chytridiales, 583-597. White, W. Lawrence. Studies in the genus Helotium, III. History and di- agnosis of certain European and North American foliicolous species, 135-170; Studies in the genus Helotium, IV. Some miscellaneous species, 599-617. Wiedling, Sten. Morphologic and physiologic investigations in diatoms, 501-502. Several fungi- 626 FarLtowia, VoL. 1, 1944 InpEX TO GENERA AND SPECIES New names and the final members of new combinations, and the pages on which they occur, are in bold-faced type; in taxonomic papers, where the main treatment of a genus or species is given, the page number only is in boldface. Synonyms and the pages on which they occur are printed in italics. Acanthocladium baculiferum 186; ben- guetense var. latifolium 187 Acanthorrhynchium papillatum 46 Achlyogetonaceae 585 ; Acia ferruginea 272; tomentosa 271 Aciella tomentosa 271 Acinaria 251; coccifera 251; flexuosa 251; latifolia 251; salicifolia 251 Acroporium diminutum 39; oxyporum 45 Acrosporangiatae 498 Actigea 255, 259; multifida 255; sicula 255 Actigena multifida 255; sicula 255 Actinomorpha aurantiaca 255 Actinothuidium Hookeri 185 Actinotrichia 496, 498 Actycus siculus 255 Aecidium citrinum 255 Aedycia 249, 255; alba 255; rubra 255 Aerobryidium aureo-nitens 182 Aerobryopsis 36; longissima 181 Agaricus 293; aterrimus 255, 256; az- ureus 255, 256; ellipticus 255; miptica 256; surrectus 256 Aglaozonia 6 Alcyonium Bursa 253 Aleurodiscus 95, 265, 267-271, 276, 307; albo-roseus 268; albus 306, 307; amor- phus 269; amylaceus 291, 300; api- culatus 267, 268; aurantius 307; Bertii 268, 307; botryosus 299; Burti 268; candidus 268; crassus 268; cremeus 268; croceus 304; diffissus 270; dis- ciformis 279; Grantii 269; helveolus 269; javanicus 267; lepra 268; macro- sporus 307; Micheneri 303; mirabilis 267, 268; Oakesii 268; pallide-roseus 268; paraphysatus 297; peradeniae 267; Peteloti 268; polygonioides 300; roseus 299; sajanensis 270; salmoneus 268; sinensis 268; spinulosus 267; sub- cruentatus 307; subgiganteus 303; succineus 270; usambarensis 267; Zell- eri 270, 319, 324 | Allophylaria phyllophila 158 Alternaria maritima 405 Alveolinus 256 Alytosporium croceum 304 Amanita aterrima 255, 256; azurea 253, 256 Amasperma 251; flocculosa 251; monilia 251; torulosa 251 Amaurodon viridis 277 Amblystegiaceae 193 Amerosporae 407 Amoebochytrium 592, 596 Amphinema sordescens 276 Amphiroa 337; tribulus 338 Amphisphaeria maritima 411, 440, 442- 447, 450, 451, 453, 455, 457, 458 Anabaena 10; aphanizomenoides 373; flos-aquae 10; flos-aquae var. minor 10; minutissima 10; wisconsinensis 373 Anastomaria campanulata 256; dimidiata 256 Anema Dodgei 393 Aneuriton 251; marginatum 251 Angstroemia orientalis 504 Annularia 256 Anoectangium Stracheyanum 174 Anomobryum cymbifolium 176; gemmi- gerum 171, 176 Anomodon toccoae 506 Anthoceros 392 Aphanistis 591, 993 Aphanizomenon flos-aquae 373 Aphanocapsa 392 Aporemia 251 Arthrocormus Schimperi 29 Arthrodesmus quadridens 16 Arthrodia 251; linearis 251 Artocreas Micheneri 303 Ascochyta Thaspii 580 Ascophyllum 7 Aspergillus 101, 446, 471; fumigatus 205 Asterodon ferruginosus 271 Asteromella astericola 572; Asteris 572 Asterophlyctis 591, 592, 594 Asterostroma 265, 271, 272; andinum 271, 272; bicolor 271, 272; gracile 271; ochrostroma 271; spiniferum 271, 272 Asterostromella 264, 293; dura 310; ef- fuscata 290; epiphylla 323; granulosa 283; investiens 292; rhodospora 294 Astrocitum dimidiatum 256; multifidum 256; quinquefidum 256 INDEX TO Astrycum dimidiatum 256; multifidum 256; quinquefidum 256 Athelia citrina 304; epiphylla 323; sericea 308, 309; strigosa 8 muscigena 275 Atractylocarpus comosus 28; dicranoides 28 Atrichum flavisetum 189; undulatum 189 Auricularia aurantiaca 319 Barbella burmensis 182; enervis 182; javanica 182; pendula 182; spiculata 182; Stevensii 182 Barbula consanguinea 175; constricta 175; indica 174; replicata 191; Wis- selii 30 Bartramia cubica 33; Halleriana 177; quadrata 34 Bartramiaceae 33, 177, 506 Battarrea 261 Belonium sulphureo-tinctum 169, 170 Bescherellea Eyrtapus 35; Cyrtopus var, papuana 35 Blastocladiales 593 Blastomyces 211, 213 Blastomycoides 213, 219; dermatitidis 213; immitis 213, 220; Tulanensis 213 Blastosporidium 219; Schooii 212, 220 Blyttiomyces 590-593, 595 Boletus hematoporus 256; pusillus 256 Bombardia 409 _ Botryobasidium 95, 96, 97, 264, 272, 273; coronatum 107, 272; flavescens 105, 106, 272; granulatum 107; isabellinum 99, -272; ochraceum 105, 106, 272; Solani 113, 272; subcoronatum 104, 272; vagum 110, 272 Botryocladia Skottsbergii 344 Botryohypochnus 96, 100 Botryophialophora 403; marina 404 Botrytis 577; cinerea 484, 487, 577 Bourdotia deminuta 294 Brachyhymenium acuminatum 176; nepa- lense 33, 176 Brachytheciaceae 37, 44, 185 Brachythecium Buchanani 185; longi- cuspidatum 185 - Braunfelsia dicranoides 28 Breutelia Romeri 34 Brotherella erythrocaulis 187 Brotherobryum Dekockii 28 Bryaceae 32, 44, 176, 192 Bryonopsis 251 Bryopsis 251 Bryum argenteum var. lanatum 176; coronatum 44, 176; crispo-capillare VOLUME 1 627 33; flexisetum 192; megalothecium 192; nitens 176; pachycladum 33; por- phyroneuron 176; ramosum 176; scle- rodictyon 33 Bytholotrephis 492; gracilis 492 Caldesiella 272; vaga 309; viridis 277 Callicostella Karnbachii 44 Calocera cornea 88 Calothrix 391 Calycella callorioides 601; Jlicis 160 Calycellina 160, 166; Roipuliies 161, 169; punctiformis 161, 166 Calycina aurantiaca 140; epiphylla 140; fastidiosa 150; Ilicis 160; naviculaspora 151; populina 160; saprophylla 151 Calymperaceae 30, 42, 173, 504-506 Calymperes daruense 43; Ebaloi 505; Hampei 174; moluccense 43; Motleyi 43; peguense 173; porrectum 505; sala- kense 43, 505; serratum 505; stricti- folium 505; tuberculosum 505 Calyptothecium burmense 183; himan- tocladioides 171, 183; Ramosii 183; Urvilleanum 183 Campylodontium flavescens 185 _Campylopus austro-subulatus 28; gracilis 172; laetus 172; novae-guineae 28; umbellatus 172 Carteria ovata 6 Catenaria 215, 251, 592, 596; Anguillulae 216; arenaria 251; concatenata 252; vagabunda 252 Catenarioideae 592 Catenochytridium 591-593, 595 Ceramium 252; callithamnion 252; opac- um 252; squamosum 252 Ceratobasidium 116, 264, 272, 327; atrat- um 272; cornigerum 116; fibrillosum 327; nlambede 272, 273; feriginat cum 116 Ceratostoma vitreum 125 Cercoseptoria Blephiliae 578 Cercospora Alni 579, 580; Barbareae 579; Boutelouae 579; briareus 581; gentianicola 581; incarnata 581; Nas- turtii var, Barbareae 579; Violae 581. Cercosporella Pycnanthemi 578 Ceriospora 398, 408 Ceriosporopsis 408, 411; halima 409, 440-442, 444-451, 453-455, 457 Cerocorticium albissimum 314 Cerophora 256; capitata 256; clavata 256; dichotoma 256; fastigiata 256; globosa 256; globularis 256; minuta 256; pyri- 628 . Fartowia, Vou. 1, 1944 formis 256; ramosa 256; thamnoides 256 Chaetangiaceae 498, 499 Chaetangiae 498 Chaetangium 496, 498; Hystrix 342 Chaetocladium 54 Chaetocorticium fusisporum 320 Chaetomitrium 30; acanthocarpum 44; orthorrhynchum 44; torquescens var. barbatum 37 Chaetonema 351 Chaetophoraceae 350, 351 Chantransia 498; dichlora 252 Chara 251-253; capitata 253; fetidissima 252; patens 252 Characias 252 Chlamydomonadaceae 348, 349 Chlamydomonas polypyrenoideum 348; variabilis 6 Chledipole 252; lobata 252; tubulosa 247, paivs Chledripole 252 Chlorociboria 135 Chlorophyceae 10-19, 348, 349 Chlorophyta 348-361 Chlorosplenium 135 Chondrus capensis 343 : Chromosporium pactolinum 297; viride 292 Chroococcaceae 371, 372 Chroococcales 371, 372 Chrysophyceae 7, 363, 364 Chrysophyta 361 Chrysymenia Skottsbergii 344 Chytridiaceae 584, 590, 591 Chytridiales 583-597 Chytridioideae 590-593, 595, 596 Chytridium 587, 590-595; Lagenaria 587 Ciboria 135, 483 Cirriphyllum cameratum 38; cirrosum 37; novae-guineae 37; subenerve 38 Cladochytriaceae 592, 595, 596 Cladochytrioideae 592 Cladochytrium 584, 592, 596; hyalinum 596 Cladophora 5 Cladopodanthus 43; heterophyllus 43; muticus 43 Claopodium nervosum 184 Clastobryum caudatum 186; caudiforme 186; Dickasonii 186 Clathrus 249, 257; columnatus 257; trun- catus 257 Clavaria bicolor 256; byssacea 278, 279; citrina 256; citrinofusca 256; contorta 270; driophylla 256; dryophylla 256; fistulosa 270; Himantia 278; lepido- rhiza 257; oxantha 257; rubescens 257; tricolor 257 Cleistostoma ambigua 171, 179 Closteridium 10; siamensis 11 Closterium 12-14, 18, 251; angustatum 12; macilentum 9, 12; Ralfsii var. hy- bridum 12; rostratum 12; setaceum 12; striolatum 12 Coccidioides 212-214, 216, 217, 219, 220; braziliensis 214; esferiformis 214; im- mitis 199-244; immitis var. metaeuro- peus 214, 219, 220, 223; immitis var. Pipkini 214, 219, 220; immitis var. typicus 214, 219, 220; pyogenes 212, 213, 220 Coccidioidiaceae 218 Codiaceae 491 Codium dichotomum 338; prostratum 338; tomentosum 338, 491; Vaughani 338 Coemansia 49-57, 59, 61, 68; aciculifera 57, 65, 67; braziliensis 57, 64; brevi- ramosa 57, 62; ceylonensis 56, 57, 60; erecta 57, 62, 63, 65; guate- malensis 57, 64, 65; interrupta 57, 62; kamerunensis 57, 63, 65; pecti- nata 51, 55, 57, 66-68; repens 68; reversa 53, 55, 57, 60, 61; scorpioidea 55-57, 66; spiralis 57, 60, 61; Thax- teri 57, 66, 67 Coemansiella 50, 57; alabastrina 57 Coenomyces 592, 596 Coleosporium 81, 85 Colonnaria 249, 250, 257; truncata 249, 257; urceolata 249, 257 Colophermum floccosum 252 Coltricia tomentosa 526, 527 Conferva isacella 252 Coniophora 116, 265, 273-280, 287; albo- flavescens 277, 280; arida 280; arida subsp. fumosa 267, 274; atrocinerea 266, 273, 274, 280; avellanea 275, 300; Betulae 280; brunneola 273; byssoidea 275, 314; centrifuga 286; cerebella 276; corrugis 276; corticola 277, 280; crocea 277; cyanospora 277; dryina 277, 279, 325; Ellisii 273; flava 278; flavomar- ginata 278; fulvo-olivacea 274; fumosa 273, 308; furva 273; fusca 274; fusi- spora 105; Harperi 277, insinuans 310; Karstenii 274; laeticolor 277, 278; leucothrix 273; mustialaensis 277; ochracea 105; ochroleuca 102, 103; Pe eee Sa ery TE ee er Ce en Mee Rags ys, © InDEX TO VOLUME 1 629 olivacea 266, 273, 274, 280, 308; Poly- poroidea 277, 279, 310; puteana 276; sibirica 274; Sistotremoides 274, 280, 308; sordulenta 309; suffocata 280, 308; sulphurea 308; umbrina 273, 275, 280; vaga 105, 106, 280 inum 277; effuscatum 290; effusum 290; Eichlerianum 314, 315; Ellisii 273; epiphyllum 323; ermineum 291, 300, 312; evolvens 318; fenestratum 105, 106, 291; fibrillosum 307, 308; flavescens 105, 107; flavissimum 304, vy eh ee Re ee ae eee esha Bag 305; frustulosum 105, 107; frustulosum var. intermedia 105; fumesum 273, 308, 309; furfuraceum 285,°286; fuscostrat- um 291; fusisporum 103, 105; Galzini 285; geogenium 281, 291, 306, 311; glebulosum 284; Greschikii 310, 311; ulaqueatum 281, 292; illaqueatum fa. rhizophorum 281; inaequale 311; in- canum 292; incrustans 285, 299; in- vestiens 292, 299; involucrum 293, 294; isabellinum 99, 107-109; jamaicense 294, 310; javanicum 267; Koleroga 112, 113, 294; lacteolum 286; lacteum Coniophorella 274; atrocinerea 273; bys- soidea 276; laeticolor 278, 279: ochro- leuca 102; olivacea 274; olivacea subsp. fulvo-olivacea 274; umbrina 274; um- brina var. olivacea 274 Conjugales 360, 361 Coprotinia 484; minutula 484—487 Corallina Flabellum 337 Coralliochytrium 591, 592, 595 Cordyceps 398 Coronella 49-51, 57; nivea 49, 50, 57 Corticium 59, 95, 264, 265, 279, 280-306; sect. Athele 292; sect. Botryodea 96; abeuns 280, 281; albo-cremeum 295; alboflavescens 277 ; albo-ochraceum 279, 290; albo-ochraceum subsp. amianthin- um 289; albo-stramineum 281, 311, 312, 321; albulum 312, 316; album 116, Allescheri 313; alutaceum 59; alutarium 292, 293; amianthinum 289, 290; amor- _phum 269; amylaceum 291; apiculatum 281, 282, 292; arachnoideum 282, 286-288; araneosum 99; areolatum 113-115, 281, 282; argillaceum 296; Atkinsonit 282; atratum 272; auranti- acum 270; Berkeleyi 282, 283, 326; bicolor 304, 305; bisporum 286, 288; bombycinum 283, 295, 296, 325; botry- oideum 107, 108, 111, 284; botryosum 108, 110, 111; brunneolum 273; bys- sinum 304, 305; byssinum var.. micro- sporum 304; byssoideum 275; caerule- um 273; calceum 284, 285, 317; calo- thrix 316; calotrichum 318; canum 285, 286; centrifugum 282, 286, 287; cen- trifugum subsp. bisporum 287; cen- trifugum subsp. fugax 287; centrifugum var. macrospora 287; centrifugum var. soredioides 286; centrifugum var. tenuis 287; Chusqueae 282; citrinum 304; confine 288; confluens 275, 276, 296, 300, 306; consimile 286-288; contiguum 288, 289, 301, 302; cornigerum 116; coronatum 107, 109; coronilla 286, 288, 292, 328; corruge 2763; cremeum 314, 315; cremoricolor 327; cretaceum 284, 285, 325; creceum 304; crustaceum 288, 289, 302; crustulinum 289; de- cipiens 286-288; deglubens 327; dry- 294-296; lacunosum 275; laeticolor 277; laetum 296; laeve 318; Jatitans 282; leucothrix 273; Litschaueri 281, 300; livido-caeruleum 281; Macounii 305, 306; maculatum 305, 306; micro- sclerotia 113-115; microsporum 286: miniatum 319; Murrayi 290; mustia- laense 277; mutatum 313; myxosporum 282; ochraceum 107, 289; ochrofarctum 296; ochroleucum 112, 113, 281, 291, 292; octosporum 287; oleosum 313; olivaceum 273; Overholtsii 296, 297; pactolinum 297; pallescens 310; pal- lidum 296; paraphysatum 297, 307; pectinatum 298; peradeniae 267; per- tenue 322; Pezizoideum 286-288, 298; pilosum 298, 299, 302; Pini-canaden- sis 323; Polyporoideum 279, 310; porosum 281, 289, 299, 300-302, 305; praetermissum 322; praticola 98, 115; pruinatum 107-109; Pruni 299, 301, 302; puberum 325; punctulatum 281, 295, 320, 321; puteanum 276; racem- osum 298, 299; radiosum 294, 295; rhizophorum 281; roseopallens 299: roseum 299, 300; Roumeguertt 324; rubellum 275, 300; rude 310; Sambuci 325; sanguineum 319; scariosum 327; secedens 327 ; septentrionale 300; serum 325; setigerum 282; soctatum 291, 300; Solant 111, 113; sordidum 314, 315, 321; sordulentum 309; sphaerosporum 286, 292; spinulosum 268; sterigmat- icum 116; Stevensti 112, 113, 300; stramineum 299, 300, 301, 305; sub- album 298, 299, 302; subapiculatum 630 324; subcinereum 289, 302; subcoron- atum 104, 105, 302; subgiganteum 291, 303; subincarnatum 303, 324; sub- pallidulum 285; subsulphureum 303; suecicum 285; suffocatum 280; sul- phurellum 277; sulphureum 99, 113, 303, 304, 305, 308, 309; tenue 322; terrigenum 116; tessulatum 281, 292; teutoburgense 105; Thelephoroides 309 ; Torrendii 312; trigonospermum 307, 308; Tsugae 296; tuberculatum 295; Tulasnelloideum 292; umbrinum 275; usambarense 268; vagum 108, 110, 111, 113, 115, 305, 308; vagum subsp. Solani 113; vagum var. Solani 113, 305; vel- lereum 295; versatum 326; vesiculos- um 301, 302, 305; vinososcabens 305, 306 Cosmarium 19; bioculatum 13; sub- turgidum 13; subturgidum fa. minor 13 Cratoneuron filicinum 193 Crossidium Roseae 192 Cryptococcus 213 Cryptophaeella Heteropatellae 120 Cutleria 6 Cyanophyta 371-373 Cyathela 257 Cyathella 257 Cyathicula 166 Cyathophorella 507 Cylindrochytridium 593, 595 Cylindrochytrium 593 Cylindrosporium Gei 577; triflori 577 Cynicus 257 Cyphella amorpha 269 Cyrtopodaceae 35 Cystopus 328 Cystoseira australis 343; caudata 343 Dacryomyces 259, 262 Daedalea 257 Daltonia aristifolia 509; armata 508, 509; Macgregorii 37 Dangeardia 591, 593, 594 Dasyscypha puberula 169 Dedalea 257 Deloxus 252 Deraphytus 252 Derbesia marina 6 Derepyxis amphora 362 Desmatodon involutus 175 Desmotheca apiculata 31; cymosa 31 Diaphanodon thuidioides 179 Dicarphus 257; rubens 257 Dichostereum durum 310 FarLowiA, Vou. 1, 1944 Dicnemonaceae 28 Dicnemos 29; calycinus 29; rugosus 29; undulatifolius 28 Dicoccum nebulosum 578 Dicranaceae 26, 172, 504 Dicranodontium uncinatum 172 Dicranoloma fragile 173; laevifolium 28; novo-guinense 28 Dicranoweisia alpina 27; fastigiata 27; papuana 27 Dicranum gymnostomum 173 Dicteridium 257 Dictilema 252; glomerata 252; xantho- sperma 252 Dictyota dichotoma var. implexa 338, 339; dichotoma var. intricata 338, 339; im- plexa 338 ; interrupta 339; subarticulata 339, 340 Didymosporae 408-413 Didymostilbe 122 Dilophospora Alopecuri 577 Diphas 250, 262 Diphascium 250 Diphascum 250, 262 Diphyscium 262 Diplodia orae-maris 403 Diplonema sordescens 275 — Diplophlyctis 587, 591, 592, 595 Diplophlyctoideae 590, 591, 592, 595, 596 Dipodascus 217 Dispira cornuta 53 Ditrichaceae 26, 172 Ditrichum Colijnii 26; spinulosum 26; spirale 26 Draparnaldia Judayi 351 Draparnaldiopsis alpinis 351 Druparia globosa 257; rosea 257; violacea 257; volvacea 257 Drupasia globosa 257; rosea 257; violacea 257 Dycticia clathroides 257 Ecklonia biruncinata 341; exasperata 341; radiata 341 Ecpexis fluviatilis 252 Ectocarpus confervoides 493; siliculosus 6 Ectosperma longiuscula 252 Ectospermia 252 Ectropothecium aureum 40; dealbatum 187; dentigerum 39; eleganti-pinnatum 40; laticuspes 39; longicapillare 46; Micholitzii 46; Moritzii 40; plumosum 40 OF EE OE ATO MEL ee eae Sie INDEX TO VOLUME I 631 Eleutheris 123 Eleutheromycella 121; mycophila 119, 123 Eleutheromyces 119, 121; Geoglossi 122, 126; longisporus 121; subulatus 119- 122, 123 Empusa 86, 211 Endacinus 257; tinctorius 257 Endaematus albus 257 Endematus 257 Endochytrioideae 591 Endochytrium 591, 592, 595, 596 Endocoenobium 592, 595 Endoconia leucomela 257; stuposa 257 Endomyces 212 Endomycetaceae 469 Endonema 252 Endonius 257 Endosperma aggregata 252; globosa 252 Endotrichella arfakiana 36; Campbelliana 36; nematosa 507; rigida 180 Enteromorpha 5, 7; prolifera 6 Entodon flavescens 186; plicatus 186 Entodontaceae 45, 185 Entomophthora 84, 88, 89 Entomophthoraceae 88 Entophlyctaceae 590, 591, 592, 595, 596 Entophlyctis 586, 587, 591, 592, 595 Entophlyctoideae 590-592, 595, 596 Entosthodon wichurae 175 Epidrolithus 250, 262 Episperma micramnia 252 Epixyla clavata 258; cylindrica 258; lineata 258 Eriosperma alba 258; fugax 258 Erysiphe graminis 569; scandens 112 Erythrodontium julaceum 185 Euastrum 13, 18, 19; humerosum 13; ob- longum 13 Eucamptodon 29; novae-guineae 29 Eucapsis 9-11; alpina 9; alpina var. minor 10 Eudacnus 258 Euglena breviflagellum 364; elastica 365; minuta 365; oxyuris var. minor 366; spirogyra var. marchica 369 Euglenaceae 364-371 Euglenophyta 364-371 Eunotia pectinalis yar. minor 501 Exoascaceae 469 Fabroniaceae 193 Favaria 258 Favolus 260; europaeus 124 Fissidens 250, 262; Brassii 41; bryoides 41; diversiretis 504; geniculatus 26; kurandae 26; lagunensis 504; micro- cladus 41; nobilis 172; papuensis 41; rupicola 26; subspathulatus 25; sylvaticus 172, 503; Wichurae 504; Zwickeyi 503 Fissidentaceae 25, 41, 172, 503, 504 Fleischerobryum longicollis 178 Floribundaria aurea 182; floribunda 182 Fomes annosus 86; circinatus 527, 528; Pini 530 , Foreauella orthothecia 188 Forsstroemia rigida 35 Fucus 6, 7; compressus 343; elongatus 341; filicinus 254; fungosus 253; im- plexus 338; interruptus 339; loreus 341; ovalis 344; ovatus 344; radiatus 341; radiatus B exasperatus 341; to- mentosus 253, 338; vermicularis 343, 344 Funaria hygrometrica var. calvescens 175 Funariaceae 175 Fungi Imperfecti 402 Fusarium 577; moniliforme 578; tricinct- um 578 Fusisporium Kuhnii 287 Galaxaura 337, 495-499; corymbifera 496; Diesingiana 496, 497; tenera 496, 497 Garckea phascoides 172 Garovaglia papuana 36; Pauerlenii 36; rigida 507; subintegra 36; Zwickeyi 507 Gastroconium ovale 344: ovatum 344 Geaster 261 Gelatina 258; alba 258; aurantiaca 258; candida 258; flava 258; foetidissima 258; lutea 258; rubra 258 Gelatinaria 258 Geminella crenulatocollis 349; muta- bilis 350 Gemmularia 258, 261; albida 258; levius- cula 258; rimosa 258; rugosa 258 Geoglossum 126, 258 Geotrichum 213, 219; dermatitidis 220; immite 220; louisianoideum 220 Gigartina bracteata 342; Hystrix 342; polycarpa 342; Radula 342; Radula var. Hystrix 342; Radula var. Radula 342 Glaucocystis cingulata 372; duplex 371; nostochinearum 10, 372; oocystiformis 372 Glenospora 219; metaeuropea 220, 223 Gloeocapsa 391, 392 Gloeocystidiellum 302; porosum 301 ‘ti Ne ee 632 Fartowia, Vou. 1, 1944 Gloeocystidium 264, 302, 306; albe- stramineum 306, 312, 321; albo-stra- mineum var. causseanum 321; albo- stramineum subsp. cremicoler 321; albo-stramineum subsp. Kichleri 321; albo-stramineum subsp. sphaerosporum 321; alutaceum 295; argillaceum 296; Bourdotii 312; caliciferum 322, 323; clavuligerum 302; contiguum 289, 301, 302; coroniferum 282; cremicolor 321; Eichleri 321; furfuraceum 302; inaequale 311, 312; Karstenii 289, 302, 306; luridum 312; ochroleucum 281; oleosum 313; pallidulum 313; pallidum 296, 313; pallidum subsp. argillaceum 296; porosum 301; praetermissum 322; praetermissum var. Bourdotii 312; rude 311; stramineum 301; tenue 322; tenue subsp. inaequale 311; tenue subsp. praetermissum 322; Torrendii 312 Gloeocystis 348 Gloeopeniophora Allescheri 313 Gloeosporium 571,572; cinctum 571, 572; Oncidii 572 Gloeotulasnella 265, 306; calospora 328; opalea 306; traumatica 306 = * Gloiophloea 496, 498 Glossadelphus hermaphroditus 510; sim- ilans 509 Gonatobotrys pallidula 313 Gracilaria concinna 343 Grandinia granulosa 282, 283; granulosa var. ochracea 283 Grateloupia Hystrix 342, 343 Gymnodiniaceae 371 Gymnodinium caudatum 371 Gymnogongrus capensis 343; vermicularis 343, 344 Gymnopus 258 Gymnosporangium 85, 89; juniperi-vir- ginianae 86; nidus-avis 79-90 Gyrophana 287 Halicystis ovalis 6 Halimeda 337 Halophiobolus 398, 415; cylindricus 416, 442-447, 450, 451, 454; halimus 416, 419, 420; longirostris 416, 418, 453, 463; maritimus 416, 419; medusa 416, 419; opacus 416, 417, 418, 440, 442, 444-448, 450, 451, 453-455, 458: salinus 416, 419, 440, 442, 444-448, 451, 453-455, 458, 462 Halosphaeria 412; appendiculata 412 Hansenula 480 Haplosporangium parvum 218, 219 Haplosporidia 212 Hectocerus 258; clavatus 258; dichotomus 258; globosus 258; pyriformis 258; thamnoides 258 Hedwigiaceae 179 Helicodendron tubulosum 444 Helicoma Curtisii 458; maritimum 405, 407; Miilleri 458; salinum 406, 409, 440-442, 446-448, 454, 458 © Helicomyces scandens 444 Helmyton glomeratum 252; spiralis 247, 252 Helotium 135-170, 599-617; albidum 154, 599, 608; albopunctum 139, 154, 155- 157, 159; album 601; amenti 617; aurantiacum 140-142; callorioides 601, 602; carpinicola 137, 138, 141, 142, 145, 146, 608; caudatum 137, 139, 150, 151, 154, 157, 159, 609; citrinulum 600-602, 604, 605; citrinulum var. Seaveri 600, 601; citrinum 599; con- formatum 136, 145-147, 162; Conocarpi 137, 138, 148; contortum 138, 147; cyathoideum 599; Dearnessii 137, 613, 616; discretum 605; epiphyllum 135- 139, 141, 142, 144-147, 159, 162, 599, 606, 608; epiphyllum var. acarium 142; epiphyllum var. candidum 142, 146, 147; epiphyllum var. Cerasi 141; epi- phyllum var. Ellisii 142; epiphyllum var. Illicina 162; epiphyllum var. prae- ponens 142; erraticum 606; fastidi- osum 139, 150, 157; flexucsum 600, 601; fraternum 137; fructigenum 599; gemmarum 616; herbarum 599, 605; Ilicis 160, 162, 164; immutabile 136- 138, 141, 142, 145-147, 150, 606, 608; Linderi 139, 154, 160; lucellum 166, 167; lutescens 137; midlandensis 605, 608; naviculasporum 151, 152; pal- lescens 137; phyllogenon 139, 155, 156, 157, 159; phyllophilum 135, 136, 139, 141, 152, 156, 157, 159, 162, 608; phyl- lophilum fa. albescens 152; populinum 138, 159, 160, 162, 164, 166, 167, 170; puberulum 138, 163, 167, 1693; puncta- tum 160; punctiforme 160, 162, 167, 170; punctiforme fa. Tiliae 162; sapro- phyllum 151, 152, 155; scutula 137, 154, 599, 609, 613, 616; scutula var. albidum 152, 608; scutula var. caudatum 151; scutula var. fucatum 6093 sor- didatum 152, 155; sparsum 15], 152; translucens 139, 149, 150; trapezoide- bgt neiraced INDEX TO VOLUME 1 633 um 147; tumidulum 139, 165-167; vir- gultorum 599 Helvella infula 128 Hepataria 258; cuneata 258; erecta 258 Hericium 256, 258; grande 258 Herpetineurum toccoae 506 Heterodon 250, 262; bryoides 250, 262 Heterolagynion Oedogonii 364 Heterophyllum brachycarpum 509; San- tosii 509 ‘Himanthalia elongata 341; lorea 341 . Himantia sulphurea 304, 309; umbrina 273 Himantocladium cyclophyllum 37 Histoplasma 211 : Holomitrium Griffthianum 27, 173; noyae-guineae 27 Homaliodendron exiguum 184; flabel- latum 184; microdendron 184 Homoeostrichus 340; flabellatus 341; multifidus 340; Sinclairii 341; spiralis 341 Hookeriaceae 37, 44, 508 Hookeriopsis utacamundiana 37 Hormidiopsis crenulata 350; ellipsoide- um 350 Hormogonales 372 Hyalinia tumidula 165 Hyalocomiaceae 188 Hyalopyenis 119, 128; blepharistoma 124, 125; hyalina 124, 125; vitrea 124, 125 Hyaloscypha lachnobrachya 167; puber- ula 169; punctiformis 160 Hyalotheca 361 Hydnochaete setigera 271 Hydnum 257; aurantinum 258; auranti- um 258; barbatum 259; caerulescens 259; citrinum 259; dilatatum 259; fer- rugineum 272; ferruginosum 272; fusco- atrum 279; granulosum 282; Himantia 278, 279; puniceum 259; setosum 279; Sobolewskii 277; subfuscum 278; to- mentosum 272 Hydrodictyaceae 356, 357 Hydromycus 259; aquosus 259; tremel- loides 259 Hygroamblystegium austro-fluviatile 193; crassicostatum 193; irriguum var. spinifolium 193 Hygrohypnum peruviense 193 Hylocomiaceae 188 Hymenochaete 271, 315; Eillisii 273; pu- bera 325 Hymenoscypha albida 609; albopuncta 155; citrinula 600; Dearnessii 613; epi- phylla 140; glagosa 167; phyllogena 156; phyllophila 158; populina 161; puberula 169; punctiformis 161; scutula var. fucata 610; tumidula 165 Hyophila involuta 174 Hyphochytrium 215, 216; infestans 216 Hyphoderma asperum 282; puberum 325; spiculosum 282; sulphureum 308 Hypnaceae 39, 46, 187 Hypnodendraceae 34 Hypnum austro-fluviatile 193; plumae- forme 187 Hypnochnella violacea 116 Hypochnopsis mustialensis 277; ochro- leuca 112 Hypochnus 95, 265, 293, 306-310; albo- stramineus 312, 321; albus 306; argil- laceus 99; bisporus 286, 287; Brefeldii 107; byssoideus 275; centrifugus 286; chaetophorus 316; confluens 275; coro- natus 107, 108, 110; cremicolor 320, 321; epiphyllus 323; fibrillosus 307; filamentosus 113-115, 308, 309; flave- scens 105; flavo-brunneus 280, 308; flavus 99; fumosus 304, 308, 309; fusig- er 103, fusisporus 102, 103, 105; gran- ulatus 107; isabellinus 99, 107, 108, 309; Langloisii 101, 309; longisporus 315; muscorum 275; mustialaensis 277 ; obducens 275; ochroleucus 112, 113; olivaceus 273, 274; pallescens 294, 309, 310; Peniophoroides 294, 310; Poly- poroideus 279, 310; puberus 325; rose- - us 300; Sambuci 325; Schroeteri 107- 109; sericeus 308; serus 325; setosus 275; Solani 113, 115; sordidus 315, 320, 321; sparsus 99; strigosus 6 fila- mentosus 275; subtilis 325; sulphureus 308; Thelephoroides 310; Tulasnello- ideum 292; umbrinus 273, 275; vagus 309; violeus 328 Hypolepia 259; difformis 259; igniarias 259; ignarius 259 Hypomyces 124, 128; Geoglossi 119, 126 Inonotus hispidus 297; Leei 297 Isaria 61; brachiata 119, 120; felina 61; odora 259 Isophlis concentricus 252 Isopterygium albescens 188; minutirame- um 46; Textori 188 Jania 337; rubens 338 Juratzkea seminervis 193 634, Kickxella 49-52, 54-57; alabastrina 49, 50, 54-57, 58 Kickxellaceae 49-56-77 Kneiffia Allescheri 313; aurantiaca 270; cremea 314; Eichleri 320; glebulosa 284; globifera 317; laevis 318; latitans 282; longispora 315; muscorum 276; polonensis 316; pubera 325; Roume- guerti 324; sera 325; setigera 282, 284; setigera var. trachytricha 283; subas- condita 324; tenuis 322; Tomentella 276 Kneifhiella aspera 282; latitans 283; seti- gera 282 Labyrinthula macrocystis 395 Lachnella puberula 169 Lactarius pargamenus 124 Lagynion reductum 363, 364; triangu- laris var. pyramidatum 364. Lamarckia vermilara 253 Lambertella 483, 487 Laminaria 253, 405; radiata 341 Lamyxis 259 Langloisula 292, 293; spinosa 292, 293 Laternea 249; columnata 249 Leiacina 251, 253; capitata 253; lucida 253 Lentescospora 411; submarina 411 Lenzites 123 Leotia lubrica 124 Lepocinclis glabra fa. minor 370 Leptodontium erectifolium 30; kinaba- luense 30; Warnstorfii 30 Leptogium 250, 262 Leptohymenium tenue 188 Leptopora 259; difformis 259; nivea 259; stercoraria 259 Leptorima nivea 253; oculata 253; un- dulata 253 Leptosphaeria muralis 413; orae-maris 413 : Leptuberia amorpha 250, 262 Leucobryaceae 29, 43, 173, 504 Leucobryum 29; arfakianum 29; Bow- ringii var. sericeum 44; Hollianum 173; neilgherrense 173; scalare 173; steno- phyllum 29 Leucodontaceae 35 Leucophanes candidum 44; Massarti 504; octoblepharioides 504 Liagora 337 Lichiton mamillaris 253; phyteloides 253, 254; tuberculatum 253 Limacium penarium 124 biruncinata 341; Fartowla, Vout. 1, 1944 Linderia 249, 250, 257 Linospora 417 Lloydella Karstenii 326 Lycoperdon 260; coccineum 259; com- planatum 259; violacinum 259, 260 Lycopodium violacinum 259 Lyngbya gigantea 373; Hummelii 373; latissima 372, 373 Lyomyces 300; byssinus 304; isabellinus 99; mustialensis 277; roseus 300; Sam- buci 325, 326; serus 325; sulphureus 308 Macrochytridioideae 591 Macrochytrium 591; botrydioides 216 Macromitrium 195-198; angustifolium 44; Blumei 195;_ brachystele 31; Dickasonii 178; falcatulum 179; goniorhynchum 179; leucoblastum 31; lonchomitrium 31; longipes 31; nepa- lense 178; ruberrimum 31; salakan- um 196, 197; semipellucidum 44; sub- megalocladum 31; sulcatum 179 Macrothamnium macrocarpum 188; sub- macrocarpum 188 Mallomonadaceae 363 Mallomonas pseudocoronata 363; ur- naformis 363 Martensella 49-52, 54, 57, 58, 59, 61, 68; Corticii 55-57, 59, 61; pectinata 50, 51, 55, 57, 58, 59, 68; spiralis 60 Martinia 483, 487; panamaensis 487 Massarinula 410 Mastopoma Armitii 38 Megaceros 515-523; alatifrons 519; amoe- nus 519; aneuraeformis 519; arachno- ideus 515-517, 519; caledonicus 516, 519; callistictus 519; carnosus 519; celebensis 516, 519; columbianus 519; crassus 516, 519; cristisporus 519; den- ticulatus 515-517, 519; endiviaefolius 516, 519; flagellaris 519; flavens 519; fuegiensis 515, 516, 519; giganteus 516, 519; gracilis 519; grandis 517, 519; guatemalensis 519; Hodgsoniae 515, 516; jamaicensis 519; Jamesonii 516, 519; lacerus 516, 519; laciniatus 519; leptohymenius 519; longispirus 516, 519; martinicensis 515, 516, 519; mexicanus 519; minarum 519; monan- drus 516, 519; monospirus 519; muricu- latus 519; Novae-guineae 516, 519; Novae-zelandiae 516, 519; Nymanii 519; pallens 516, 519; parvisporus 516, 519; salakensis 519; schizophyllus 519; solidus 516, 519; Stahlii 519; tjiboden- InpEx TO VOLUME 1 635 sis 519; tosanus 519; Vescoanus 517, 519; vincentianus 517, 519; Wiemanii 519; Zotovii 515 Megachytriaceae 592 Megachytrium 592, 596 Megalocystosporidium 212 Megalosporidium 212 Melanospora 128; vitrea 125 Melobesia 337 Merasperma 253; bifurcata 253; cylin- drica 253; dichotoma 253 Merisma candidum 268 Merulius undulatus 259 Mesasperma 253 Meteoriaceae 181 Meteoriopsis ancistrodes 183; reclinata 183; squarrosa 183 Michenera artocreas 303 Micrasterias 13, 14, 18, 19; oscitans 14; pinnatifida 14; rotata 14 Microdictyon 252 Microdus brasiliensis 26; papuanus 26; pomiformis 26 Micropodia populina 161 Micropyxis 119, 122, 125; Geoglossi 126 Mitochytridium 591, 592, 595 Mniaceae 177, 506 Mniodendron Hellwigii 34 Mnium coriaceum 177; rostratum 177; Thomsoni 177 Molendoa burmensis 174; Roylei 174 Mollisia 166; tumidula 165, 166 Mollisiella citrinula 600 Mollisina acerina 164, 167, 170 Moniliales 403-407 Monilinia 483 Morchella cava 259; odorata 259 Mortierella 53; tuberosa 51, 66 Mucoraceae 217 Mucronoporus 526; Andersoni 297; cir- cinatus 526, 527; tomentosus 526, 527 Mutinus 255 Mycastrum 259; siculum 259, 260 Myceliochytriaceae 596 Mycoacia 279; Himantia 278, 279 Mycoderma 213, 219, 480; immite 213, 220 Myriosydrum 253 Myrsidrum 253; Bursa 253; clavatum 248, 253; dilatatum 253; effusum 253; ra- mosum 253; vermilara 253 Myuriaceae 180 Myurium 46; rufescens 180 Myxophyceae 9 Neckera crenulata 183; himalayana 183 Neckeraceae 37, 183, 507, 508 Neckeropsis nitidula 507 Nemalectra calida 253; fuscata 253; ‘plumosa 253 Nemalionales 495-499 Neokneiffia aspera 283 Neolindbergia falcifolia 35 Neottiospora arenaria 574 Nephrochytrium 591, 592, 595 Netrium digitus 1] ’ Nidularia 259 Niptera sulphureo-tincta 169 Nitzschia closterium fa. minutissima 501; communis 501; Kiitzingiana 501; Kutz- ingiana var. exilis 501; ovalis 501; palea var. debilis 501; palea var. trop- ica 501; subtilis var. paleacea 501 Nodularia 269; balsamicola 269 Nostoc 253, 391, 392 Nostocaceae 373 Nostocus 253 Nowakowskia 591, 592, 594, Nowakowskiella 584, 592, 596 Obelidioideae 590, 592 Obelidium 591, 592, 594, Octoblepharum albidum 173 Odontia 277; Acerina 283; arguta 314; Brinkmanni 288; fusca 308; Himantia 278; setigera 283; Sistotremoides 273; tenerrima 99, 100; tenuis 309; vaga 308; vesiculosa 283 Oedogoniaceae 352-356 Oedogonium crenulatocostatum var. cylin- dricum fa. major 352; Howei 353; - inconspicuum 353; Kjellmanii 352; Kjellmanii var. granulosa 352; Koz- minskii 355; latviense 354; margar- itiferum 352; microgonium 353; oelandicum var. contortum 355; ovi- forme var. gracile 352; polyandri- um 355; Sawyeri 354; sinuatum fa. seriatum 354; Smithii 353; sol 354; spheroideum 353; _ spiripennatum 354; undulatum 355 Oidium 97, 208, 212, 213, 219; candicans 108; Citri 113, 115; coccidioides 212, 220; Curtisii 111; immite 212, 220; pyogenes 220; protozoides 212, 220; tomentosum 110 Olpidiaceae 585 Olpidium 13, 587 Omalycus 259; erosus 259; violacinus 259, 260 636 FarLtowlA, Vou. 1, 1944 Ombrophila 139; clavus 139; purpuras- cens 146; subaurea 137, 141, 142 Onnia 525; circinatus 525-527; tomentos- us 525-527 Onygena 260 Oocystaceae 357-359 Oocystis apiculata 357; pyriformis 357 Oospora 208 Ophiobolus 395, 416; Aalimus 395, 419; maritimus 419; medusa 419 Ophiocytiaceae 362, 363 Ophiocytium cochleare 363; desertum . var. minor 362; elongatum 363; elon- gatum var. major 362 Opospermum nigrum 253 Orbimyces 399, 404; spectabilis 404 Orimanthis foliacea 253; vesciculata 253 Orthomniopsis japonica 506 Orthomnium trichomitrium 177 Orthorrhynchium philippinense 37; Rid- leyi 37 Orthotrichaceae 31, 44, 178, 192 Orthotrichum Johnstoni 192; Lyellii 192 Oscillatoria princeps 372, 373 Oscillatoriaceae 372 Ovulinia 483 Oxydontia 279; Himantia 278 Oxytrema 253 Ozonium croceum 304, 309 Pachydisca immutabilis 145 Pannaria lepidiota 392 Papillaria Deppei 181; fuscescens 181; lanosa 181 Paracoccidioides 214; braziliensis 205, 214 Peckiella Geoglossi 126 Pediastrum biradiatum var. emarginatum fa. convexum 356; muticum var. crenulatum 356; quadricornutum 356; tetras 357 Pellicularia 95, 96-118, 264, 272; ansosa 97, 102, 103, 104; asperula 97, 100; chordulata 97, 98; cystidiata 97, 101, 102; filamentosa 98, 111-113- 116, 272, 282, 305; flavescens 98, 102, 103, 105, 106, 114, 116, 272, 280, 291; isabellina 97, 99, 272, 309; Koleroga 96-98, 112, 294, 300; Langloisii 97, 101, 309, 319; lembospora 98, 109; ochroleuca 97, 102-104; pruinata 98, 107, 108, 272, 284; subcoronata 97, 102, 104, 272, 302; vaga 98, 108- 110-111, 272, 305 Penicillium 446 Peniophora 95, 264, 265, 285, 293, 310- 327; admirabilis 310; affinis 318; alba 310, 311, 324; albo-straminea 264, 281; 291, 306, SLL, 321: albula 282, 295, 312, 316, 323; Allescheri 313, 318; alutaria 313, 314, 319; arachnoidea 314, 324; argentea 315; argillacea 325; aspera 282-284, 326; asperipilata 315; attenuata 266; aurantiaca 270, 271, 296, 298, 319, 324; Burkii 312, 316; byssoides 275, 276, 308, 314; byssoides subsp. Tomentella 276, 314; calothrix 316; canadensis 316; can- dida 295; chaetophora 103, 3163 cin- erea 289; coccineo-fulva 326; crassa 320, 326; cremea 313, 314, 315, 324; cremea subsp. Allescheri 313; cremea var. Allescheri 313; cremea var. Eich- leriana 314; cremea var. glauwcescens 314; cretacea 284; crocea 278; delec- tans 316; diffissa 270; dissoluta 316; dryina 277-279, 325; duplex 296, 297; KEichleri 320; Eichleriana 314, 315; filamentosa 293; flammea 316, 317; flava 278; fusispora 102, 103, 105; gigantea 317, 328; glebulosa 284, 317; glebulosa subsp. Pirina 316; globi- fera 317; gracillima 317; Greschikii 105, 310, 311, 324; hastata 311; hetero- cystidia 317, 318; incarnata 276; 298; Karstenii 326; Kauffmanii 317, 318; laevis 318; laevis var. affinis 318; laminata 313, 314, 319; latitans 283; lepida 270, 271, 319; limonia 319; livida 285; longispora 315; longispora var. clavispora 315; longispora var. cyl- indrospora 315; longispora var. gloeo- cystidiata 315; longispora var. myceli- alis 315; ludoviciana 316; luna 320, 326; lurida 266; magnahypha 101, 319; media 266; miniata 319; Molleriana 324: muscorum 276; mutata 313, 318; ochroleuca 102; Odontioides 313; odo- rata 320, 327; pallidula 313, 314, 319, 324; pallidula var. regenerans 313, 314; Peckii 320, 321; pertenuis 322; phyllophora 307, 323; Piceina 323; pilosa 298; Pirina 316; polonensis 316; praetermissa 322; pubera 325; rimicola 273; Romellii 284, 285; rosea 300; Roumeguerii 324; rudis 311; Sambuci 325; sanguinea 319; san- guinea subsp. anaemacta 319; setigera 283; Sheari 270, 271, 324; sordescens 276; sordida 314, 315, 321, 324; sor- didella 315, 321; sphaerospora 321; ee ey en re ys 2c ee ee ee Ae ee a ee ae an CGM ee, Wee ee ae, ye a Eee ee ee ne eee TEETED HGS hy Se PRE ete VOCRT Ge tr orem OmvRr Mametgy (Tee Wer a tart) dls tee eee InpEx To VOLUME 1 | 637 stratosa 324; subalba 298, 302; sub- alutacea 103, 266; subapiculata 311, 313, 324; subascondita 324; subcremea 311, 324; subcremea var. subuncinata 311; subincarnata 303; subsulphurea 303, 304, 324; subtilis 325; tabacina 278, 325; tenuis 264, 322; tenuissima 325; Thujae 325, 326; Torrendii 312; trachytricha 282, 326; velutina 315; versata 326; verticillata 320, 326 Peniophorella pubera 325 Penzigiella cordata 171, 180 Peritrichospora 398, 414, 448; integra 414, 415, 440, 442-447, 449, 451, 454, 455, 458; lacera 415 Peronospora alta 580 Pexisperma 253; amplectens 253; de- pressa 253; dichrosperma 253; lutes- cens 247, 253; sputo 253; truncata 253 Pezicula phyllophila 158 Peziza albida 608; alborufa 260; amorpha 269; atrata 260; caudata 151, 152; cupularia 260; cupularis 260; Dear- nessti 613; depressa 260; epiphylla 140; epiphylla var. populina 156, 157; fucata 609; glagosa 167; globulosa 260; lachnobrachya 162; lupularia 260; ochrochlora 260; phyllophila 158; pu- berula 169, 170; pulcherrima 260; punc- tiformis 160, 163, 164, 166, 169, 170; smaragdina 260; smeraldina 260; smir- aldina 260; tumidula 162, 165 Pezizella 137, 166; albopuncta 155; citrin- ula 600; culmigena 604; Dactylidis 600; epiphylla var. populina 156, 157; glagosa 139, 167; populina 160; pu- berula 162, 169, 170; punctiformis 160, 162, 164, 165; punctiformis fa. folii- cola 162, 164; punctiformis fa. fruti- cola 165; punctiformis fa. petiolicola 162; tumidula 160, 162, 165-167; tumi- dula fa. Castanae 162 Phacidium coronatum 163 Phacus anacoelus var. undulata fa. major 366; asymmetrica 366, 367; Birgei 367; chloroplastes 367; chloroplastes fa. incisa 368; costata 368; crenulata 368; hispidula 367; pseudoswirenko 368; Raciborski 367; Segreti var. ovum 369; setosa var. crenata 368; spirogyra 369; spirogyra var. maxima 369; striata 369; Swirenko 369 Phaeociboria 138, 483 Phaeoseptoria 576; Calamagrostidis 576 Phallus 260 ~ Phanerochaete odorata 320 Phaxantha lichenoides 254 Pherima 260 Phiala 260 Phialea albida 609; Cassandrae 139, 159, | 617; Dearnessii 613; gemmarum 616; phyllogena 156; phyllophila 158, 162; phyllophila fa. fagicola 156; phyllo- phila var. Jaapii 151, 152; scutula var. fucata 610 Phialina puberula 161, 169 Phialophora 402 Phialophorophoma 402; __litoralis 403, 442, 444, 445, 447, 450, 451, 454, 463 Philonotis angusta 178; aristifolia 177; evanidinervis 506; falcata 178; Griffith- jana 177; imbricatula 178; perlaxi- folia 178; speciosa 178; Thwaitesii 34; Turneriana 178 Phlebia 319; coccineo-fulva 326; vaga 308, 309 Phlebriella vaga 309 Phlyctidiaceae 586, 590, 591 Phlyctidioideae 590, 591 | Phlyctidium 591, 594 Phlyctochytrium 590-595 Phoracis filicina 254 Phorima 260; betulina 260; boletoides 260; coccinea 260; difformis 260; min- uta 260 Phragmosporae 413 Phycerus bifidus 254. Phycopteris angustata 339; cuneata 339, 340; Harveyana 340, 341; interrupta 339 Phylictis bifurcatus 254: cervicornis 254; cuneiformis 254; dichotomus 254; Jati- folius 254; polypodioides 254; sub- fistulosus 254; undulatus 254 : Phyllachora 575; graminis 575; oxalina 573 Phyllops 260 Phyllosiphon 387, 389 Phyllosiphonaceae 388, 389 Phyllosticta 572; guttulatae 573; Heli- anthi 573; solidaginicola 573, 574; solidaginis 574 Phyllotylus australis 342 Physidrum aggregatum 254; hyalinum 254; pisiforme 254; rubescens 254 Physocladia 592, 596 Physoderma 569; Claytoniana 569 Physorhizophidium 591, 594 638 Physotris capitata 254; glomerata 254 Physydrum 254 Phytelis atra 254; granulata 254; macro- carpa 254; radiata 247, 254; sulcata 254; tuberculata 253, 254 Phyxalium 254 Pichia 475, 477, 480; farinosa 477 Piemycus 260 Piesmycus 260; nigrescens 260; violaceus 260 Pilaira 215 Pilobolus 215 Piloecium 46; pseudorufescens 45 Pinnatella ambigua 508; Kuhliana 508; mucronata 508 Piptocephalis 53, 54 Pisolithus 257; tinctorius 257 Plagiotheciaceae 186, 509 Plagiotheciopsis oblonga 45 Plagiothecium neckeroideum var. javense 509 Plaxarthrus 254 Pleurosporangiatae 498 Pleurotus 124 Podochytrium 591, 594 Pogonatum gymnophyllum 189; inflexum 189; Junghuhnianum 189; macrophyl- loides 510; microstomum 189 Polychytrium 592, 596 Polyphagoideae 590, 592, 594, 596 Polyphagus 587, 592, 595 Polyporus 528; adustus 155; balsameus 525; betulinus 123; circinatus 525-528; circinatus var. dualis 528; circinatus var. triqueter 528; dualis 525-528; frondosus 123; peakensis 526; Schwein- itzii 525; tomentosus 525-528; tomen- tosus var. americanus 527, 528; to- mentosus var. circinatus 527, 528; tri- queter 526, 528 Polysaccum 257; pisocarpum 257 Polysiphonia violacea 6 Polystema 260 Polystictus 525; circinatus 525-568; cir- cinatus var. triqueter 528; tomentosus 527-530, 532, 536; versicolor 123 Polytrichaceae 189, 510 Polytrichum 13; xanthopilum 189 Poria 259; subacida 525, 537 Poronea 260 Porula 254 Posadasia 214, 219; esferiformis 220; sphaeriformis 212 Potarcus bicolor 254 Pottia flavipes 191; Heimii 192 Fartow1a, Vou. 1, 1944 Pottiaceae 30, 174, 191 Priapus 260; niveus 260 Prionitis 342; Andersoniana 342; austral- is 342; filiformis 342; linearis 342 Prionodontaceae 35 Protomyces 213, 216-218 Protomycopsis 217, 218 Pseudococcidioides 213, 219; Mazzai 213, 216, 220 Pseudohelotium puberulum 162, 169,170; punctiforme 160 Pseudohypnella verrucosa 510 Pseudopiloecium 45, 46; scabrisetum 45 | Pseudopohlia bulbifera 171, 176 Pseudoscinaia 496, 498 Pseudosymblepharis pervaginata 171, 174 Psilopezia mirabilis 267, 268 Psorospermia 211 Pterobryaceae 36, 180, 507 Pterobryopsis acuminata 181; crassicaul- is 180; caudata 180; flexipes 181; orientalis 181 Pterogoneurum Roseae 192 -Ptilium crista-castrensis 188 Puccinia 85; Andropogonis 580; Andro- pogonis var. Polygalina 580; Eleochar- idis 580; Helianthi 571; Liatridis 570, 571; rubigo-vera 577 Pycnodon asper 283 Pycnopeziza 138, 149; pachyderma 149; Sejournei 149 Pyrenium 260 Pyrenomycetes 407—420 Pyrisperma 260; hypogea 260 Pyrrophyta 371 Quadrigula 360 Radulum investiens 292, 293; Pini-cana- densis 323 Ramaria 260 Ramularia Equiseti 578 Remispora 398, 409, 411; maritima 410, 440 Rhacopilaceae 179, 506 Rhacopilum magnirete 506; Schmidii 179, 506; spectabile 506 Rhaphidostichum 46; loriforme 45; Pullei 38 Rhinosporidium 214 Rhizidiaceae 586, 590-594, 596 Rhizidioideae 590, 591, 594, 596 Rhizidiopsis 591, 594 Rhizidium 587, 591, 592, 594 Rhizochrysidaceae 363, 364 INDEX TO VOLUME 1 Rhizoclosmatium 591, 592, 594 Rhizoctonia centrifuga 286, 287; micro- sclerotia 115; Solani 115 Rhizophlyctis 586, 591, 592, 594. Rhizophydium 587, 591, 593, 594. Rhizopus nigricans 51 Rhizosiphon 591, 592, 595 Rhodobryum giganteum 177 Rhodura 497 Rhynchonectria 122 Rhynchostegiella’ menadensis 45; papu- ensis 44 -Rhynchostegium menadense fa. gracile 38 Riella 261 ' Rimella 261; obovalis 261 Rivularia 391 Robergea 417 Rosenscheldia 573; Heliopsidis 572, 573 Rotula 261 Roya 251 Rozea pterogonioides 185 Rugosaria 258, 261 Russula 128; adustus 124 Rutstroemia 135, 139, 146, 149, 483, 487 Saccharomyces 212, 475, 477; farinosus 477 Saccomyces 591, 592, 595 Samarospora 407, 408 Samarosporella 407; pelagica 408 Sarcanthia 261 Sarcomelas 261 Sarconemus 261 Scenedesmaceae 359, 360 Scenedesmus opoliensis var. contacta 359 Scherffeliomyces 591, 592, 595 Schistomitrium heterophyllum 43 Schizochlamys compacta 348; delicatula 349 Schlotheimia Campbelliana 32; emargin- ato-pilosa 27, 32; longiseta 32; Mac- gregorii 32; rubiginosa 32; subrubi- ginosa 32 Sciadocladus celebensis 34; Kerrii 34; novae-guineae 34; Wisselii 34 Scinaia 496-498; furcellata 493 Scinaiae 498 Sclernax lutescens 254; truncata 254 Scleroderma 255, 259; flavidum 255; geaster 255; violacinum 260 Sclerospora 84, 89; philippinensis 88 Sclerotinia 483 Sclerotiniaceae 483-488 Sclerotium quercinum 163 639 Scolecosporeae 415—420 Scopulariopsis 219; americana 214, 220 Scyphophorus 250, 262 Scyphorus 250, 262 Scytonema 391, 392 Sebacina 265, 266, 327, 328; calcea 285; deglubens 327; deminuta 293; ftbril- losa 327; incrustans 327; polyschista 327; scariosa 327 Sematophyllaceae 38, 45, 186, 509 Sematophyllum revolutum 38; subhumile 187; tristichulum 187 Septochytrium 592, 596; variabile 596 Septogloeum oxysporum 576, 577 Septoria 575; Andropogonis 576; Andro- pogonis var. Sorghastri 576; Ane- mones 980; Cacaliae 581; cylindrica 580; emaculata 576; Nabali 581; Sene- cionis-aurei 581; Senecionis-sylvatici . 581; Thlaspii 575° Septosperma 591, 594 Septotinia 483 Serpula 287 Siphonaria 591, 592, 594 Siphorus alternus 254; fasciculatus 254 Siphytus filiformis 255; hexodon 255; obconicus 248, 255 Sisostrema 261; globularis 259, 261 Sistotrema 261, 264, 328; coronilla 328; _ subtrigonospermum 328 Solutoparies 591, 594 Speira pelagica 407 Spermipole effusa 255 Sphaeria 261; Helvellae 127; maritima 413; subulata 122 = Sphaerococcus vermicularis 343 Sphaeromyxa subulatum 122 Sphaeronema 120, 121, 128; dblephari- stoma 119, 124, 125; cylindricum 121; Helvellae 127 ; oxyspora 122; subulatum 119, 120, 122; vitreum 124, 125 Sphaeronaemella 119, 121, 127, 128; Helvellae 119, 121, 127, 128; oxyspora 122, 123; subulata 122; vitrea 125 Sphaeropsidales 402, 403 Sphaeropsis 261; demersa var. foliicola 574; foliicola 574 Sphaerulina orae-maris 413 Sphagnaceae 25, 172 Sphagnum sericeum 25; subsecundum 172 Spicaria croceum 304 Spinoclosterium curvatum 13 Spinocosmarium 16; quadridens 16 Spirillum desulfuricans 471 640 FarLowia, VoL. 1, 1944 Spirogyra floridana 361; illinoisensis 361 ; pseudofloridana 360 Splachnaceae 175 Spongia dichotoma 338 Sporodesmium cellulosum 407 Sporonema 575 Sporophlyctidium 591, 592, 594 Sporophlyctis 592, 595 Sporotrichum 211; croceum 304; flavis- simum 304 Stagonospora 575; Paspali 575; simplicior fa. Andropogonis 575 Staurastrum 16; Johnsonii var. depauper- atum 17; muticum 14; punctulatum 17; Rotula 17, 18 Stemastrum 261; Boscii 261 Stephanoma tetracoccum 127 Stereonema 255 Stereophyllum tavoyense 186 Stereum candidum 268; crustaceum 288; duriusculum 310; insinuans 309; Kar- stenii 320, 326, 327; Karstenii fa. in- crustata 326; lepra 268; Murraii 290; Murraii fa. tuberculosum 290; oderatum 320, 327; Pini 297 Stericium 26] Stigeoclonium Jubricum 351; pachy- dermum 350 Stipitococcaceae 361, 362 Stipitococcus apiculatus 361; capense 362; crassistipatus 362 j Stipiza 261 Stromatinia 483 Stypnion fluitans 255 Syncephalastrum 52, 55, 56; racemosum 54 Syncephalis 52, 54-56, 58; pycnosperma 54, 55; reflexa 55, 56; tenuis 54, 55; Wynneae 54, 55 Synchytriaceae 217, 585 Synchytrium aureum 580 Syrrhopodon albovaginatus 30, 42; asper 30; Beccari 30; bornensis 30; hispido- ciliatus 504; Mulleri 43; perarmatus 42, 504; philippinensis 43, 505; stricti- folius 505 ; tuberculosus 505; Wallisii 30 Taphridium 217, 218 Taphrina lutescens 569 Taxithelium capillarisetum 39; dimorpho- phyllum 39; glossoides 39; instratum 46; nepalense 39, 187; nitidulum 39; novae-guineae 39; selenithecium 39 Tayloria indica 175 Tetradesmus Smithii 360 Tetraédron 11; asymmetricum 357; bifurcatum var. minor 358; cruciatum var. polyfurcatum 358; cruciatum var. reductum 358; enorme 358; enorme var, pentaédricum 358; irregulare 358; limneticum var. gracile 358; lobulatum var. crassum 359; main- ensis 11; regulare var. granulata 359; regulare var. incus fa. major 359; tortum 11 Tetraspora 253; lamellosa 348 Tetrasporaceae 348, 349 Thamnium subseriatum 184 Thelephora amorpha 269; aspera 282, 284; aurantiaca 261; bolaris 319; bom- bycina 295, 296; byssoides 275; calcea 284; calcea * glebulosa 284, 317; can- dida 268; candidissima 268; centrifuga 286; citrina 304; confluens 275; cre- tacea 285; epiphylla 323; evolvens 318; fumosa 308, 309; granulosa 282-284; incrustans 327; insinuans 309; isabel- lina 99, 100; lactea 295, 296; laevis 318; Menieri 308; miniata 319; Mur- raii 290; nigrescens 261; odorata 320; olivacea 266, 273, 275; pallescens 309; _ pubera 325; puteana 276; revoluta 261; rosea 299; Sambuci 325; san- guinea 319; semicampanulata 261; sera 325, 326; setigera 282, 284; Sisto- tremoides 273, 280; subochracea 292; sulphurea 304, 308, 309; umbrina 273, 275, 318; umbrina var. lignatilis 274 Thuidiaceae 184, 506 Thuidium cymbifolium 185; glaucinoides 184; investe 184; Meyenianum 184; tamariscellum 184 Thyridium 30; gracile 43 Tiarospora 409 Timmiella anomala 174 Tomentella 264, 279, 306; sect. Tomen- tellastrum subsect. Botrytes 96; albo- straminea 321; byssina 304; fibrillosa 307; flava 99; fugax 287; granulata 107, 109; isabellina 99; Menieri 308; obducens 275; ochraceo-viridis 99, 100; sulphurea 308; testaceogilva 99; trig- onosperma 307, 308; umbrina 274 Tortella cyrtobasis 174 Trachelomonas hexangulata var. repanda 370; mammillosa 370; __ peridini- formis 370; superba var. spinosa 370 Trachyphyllum inflexum 185 Trachypodaceae 35, 179 Trachypodopsis auriculata 35; crispatula 180 [INDEX TO VOLUME 1 Trachypus 36; bicolor 179; perplicatus 35 Trametes Pini 530 Tranzschelia Pruni-spinosae 576 Trechispora 264, 328; Brinkmanni 287, 288, 328; coronifera 282, 288; sub- trigonosperma 328 Trematodon longicollis 172 Tremella 258 Tremellodendron candidum 268 Trentepohlia 391 Treuberia 359 Tribonema 364 Trichia 261 Trichidium 261 Trichoglossum 119; Farlowi 126, 127; Walteri 127 Trichosporium pedrosoi 205 Trichosteleum Boschii 46; hamatum 46 Trichostomum angustatum 174 Tuber 260; rufescens 261 Tuberium 261 Tucahus 258, 261; albidus 258, 261; leviusculus 258, 261; rimosus 258, 261; rugosus 258, 261 Tulasnella 265, 266, 328; bifrons 310; calospora 328; Eichleriana 328; metal- lica 272, 273; Violae 328; violea 328 Tylostoma 261 Typhula juncea 605 Udotea flabellata 337; Flabellum 337, 338 : Ulothrix aequalis 349; cylindricum 349; moniliformis 350 Ulotrichaceae 349, 350 Ulva tomentosa 253 Urceolella puberula 169 Uredo citrina 261 Uromyces Fabae 576 Vararia 264, 293, 299; alutaria 292; ef- fuscata 290, 294; granulosa 283; in- vestiens 292; pallescens 309, 310; pectinata 298; Peniophoroides 294, 310; phyllophila 323; racemosa 299 Vaucheria dichotoma 255; flexuosa 255; Gardneri 387-389; stricta 255 Vermilaria 255 — Vesicularia elegantula 40 Vibrio estuarii 471 Vibrissea 398 Volvaria coccinea 261 Volvox fuscus 255; ovalis 255 Volvycium 261; coccineum 261, 262 weg Ee a? Lt ere ee Se Cn ee 2 eee Leen fe ee ee eh ee Seer i 641 Warburgiella 38; leptocarpa 38; papuana 38, 39; subpapuana 38 Webera rubripila 32 Xanthidium 14; antilopaeum 14, 15; anti- lopaeum var. truncatum 16; antilopae- um var. Tylerianum 16; armatum 14; cristatum 14; pseudobengalicum 14; subhastiferum 14; Taylorianum 16; tetracentrotum 14, 15; tetracentrotum var. quebecense 15; Tylerianum 16 Xanthocapsa 392, 393 Xanthochrous 297; circinatus 526, 527; tomentosus 526 Xanthophyceae 361-363 Xerocarpus alutarius 292, 293; crustaceus 288; laeticolor 277; odoratus 320, 326, 327; subsulphureus 303 Xylissus 262; -cylindricus 262; lineatus 262; oblongus 262 Yeast G 476 Zarnardinula 342; Andersoniana 342; australis 342; filiformis 342; line- aris 342 : Zonaria 255; cuneata 340; dichotoma var. intricata 338, 339; flabellata 341; Harveyana 340; interrupta 339, 340; multifida 340; spiralis 341; sub- articulata 339, 340; Tournefortii 340; Turneriana 339 Zonilia 262 Zygnema pectinata 6 Zygnemataceae 360 Zygodesmus argillaceus 99, 100; pubidus 276 Zygohansenula 480 Zygopichia 469-475-481 ; Chevalieri 471- 475, 477-479; Chevalieri var. Ander- sonii 472-476; Chevalieri var. Bark- eri 474-476; Chevalieri var. fermentati 479; chiantigiana 473, 474, 478; fari- nosa 473-475, 477 Zygorhizidioideae 591 Zygorhizidium 591, 594 Zygosaccharomyces 469-481; bisporus 476; Chevalieri 475-477; farinosus 477: Guilliermondii 471, 473, 474, 478; japonicus 479; japonicus var. Soya 4.79; major var. salsus 479 Zymonema 213; immitis 220 Zythia compressa 122, 123; subulata 122 MANUSCRIPT Contributions from anyone will be considered, preference being given to papers dealing with phases of the taxonomy of the Fungi, Algae, Musci, Hepaticae, and Lichens. © Contributors should strive for a clear concise style of writing. A limit of 100 printed pages is set for any one article, but it is hoped that a paper of such length will be so arranged as to be divisible in two approximately equal parts. Writers are responsible for the accuracy of references to literature cited and to text illustrations. 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