i vile PAL Est Labets C878 ew Het ; fe } re ‘ ’ if * ; itt ‘ t : 1 nf a yt ‘ , Hy! + olbaeirlslt : Rota! f if stil Tegeteteiats Tester late it bieltt otet ti « te Hi 1 i water ttet sit rth } ; a fifi ¥ a SO {53 ptetee aS bf a ie ritls tiatetetaaery sant : : te Irethdetet abit fetaatet cutitat ritatalt , ith ie ' . 3 re + if iran ‘te : ane fet ah $s op )' aot § i font ie! utits? Pfister statelt the : ts if i Teleiel ele rtetatite ie : Teese tt pata? oT, ; hei ROCA ERR On Betien neha: 24° ? it bron seit rs Tetela’ + Het ant if a MS Ars oe) year! a siitrrttiitiite ‘ LY ne ii ’ She eth t 1% . tint ty te ‘ fh watt i ae . mba pence eatete Acworth net ins if ett, pat as ditties seit i ete eae f i a cats .. o a es Soy a en Soe rare tae ee ceiwret So it) by ibe rt * Di EE ie seta ae te sere eies fae * . I ite by a elt ert a ait SQ ey ys e= > D> > DATES. OF PUBLICATION OF THE PARTS. ee CEs © T= (OO. jn faeceweees tes ceca saaed on Wasaeaces June 15, 1808. Peer ol Pp. 69-194, Fl XV goo. 3.c 2. ener Feb. 22, 1899. Preeti. Eps LQ5=3h 25 bls ela y NON. co ccciws ous cnmaas Dec. 29, 1399. Pari. Vip. 353-530, Rl ANT XN, occ. scent Aug. 15, 1900. eee Ves reo. 537-050, ble eR Ske ee sa deanate July 20, Igor. Peat vile lp. O57—700, Pl. WI-EXEN, oh coche acne cee ay 1902. xe I. CONTRIBUTIONS TO A KNOWLEDGE OF THE LICHENS OF -MINNESOTA—III. THE ROCK LICHENS OF TAYLORS FALLS. Bruce FINK. THE COMPOSITION AND ORIGIN OF THE FLORA. The lichens listed in this paper were collected during two reg- ular annual excursions of the Summer School of the University of Minnesota. The date of collection of all numbers up to 92 is August 15, 1896, and the plants were collected by Professor Conway MacMillan and the writer, on the Algonkian igneous rocks of the Interstate Park, or on earth or branches in the crevices of the rocks. On the 14th of August, 1897, I accom- panied another excursion to examine the lichens growing on the Cambrian sandstone exposures near the park. Numbers g3 to 11g were collected by me during this second trip, and more spe- cies were noted on the sandstone, which had been collected the year before on the igneous rocks. In all, 24 lichens were found growing on both sandstone and igneous rocks, 22 on the igne- ous rocks only, 20 on the sandstone only, 10 on earth in the crevices of the igneous rocks and 2 on roots or branches in the crevices. The last lichen of the list was found on old boards and is recorded here because rare or difficult to detect and new to the State. It was my intention when I went to the park to publish what- ever might be found of interest with the last number of this ser- ies of papers. But after observing the field it became apparent that the locality is one of great interest both as to origin and present composition of its lichen flora and that these character- istic floral features could be presented best in a separate paper. A little observation showed that the tree lichens do not differ to any noticeable extent from those about Minneapolis, and I con- sequently confined my collecting to the igneous rocks and 2 MINNESOTA BOTANICAL STUDIES. sandstone. There are rocks near the park containing lime, but they did not seem to support any lichens of special interest. The collecting was all done on the Minnesota side of the river for the reason that the erosion of the valley has been such as to leave better exposures of rock here than on the Wisconsin side. I had collected from excellent exposures of igneous rocks in New England and have since visited similar ones in various parts of northern Minnesota; but I have never seen any other equally limited area of rock exposure that aroused so much in- terest, because of richness of lichen flora and evidence of mi- grations and struggle, as did this little area, set aside for an interstate park. During the trip of 1897 I noticed that the con- stant tramp of feet had begun to kill out the lichens in many places so that the impression of richness is beginning to fade, and the botanist must soon seek some place near by, if such ex- ists, where he may study this rich flora in its natural beauty. Professor E. E. Edwards, of Lancaster, Wisconsin, writes thus of the lichens found in the park: ‘‘ The rocks of the Dalles owe their beauty and variegated tints not alone to the metal oxides, or to the feldspar or hornblende chiefly composing them, but to the growth of minute lichens upon their surfaces, and these vary in color according to the dryness or moisture of the atmosphere. We have, therefore, in these, through sunshine and shadow and the varying seasons, an endless and almost kaleidoscopic play of colors that makes them alike the delight and despair of the artist.” The little area, being one of great natural beauty and set apart for an interstate park, will always attract thousands of visitors annually and I hope to present in this paper thoughts which will enable the botanist who has a fair knowledge of lichen species and their distribution to see in this wonderful lichen population something of far greater inter- est than mere beauty. Some comparisons between the locality now under considera- tion and others will best show its richness in rock lichens. The area examined covers only a few acres of surface and gave 66 lichens growing on rocks as a result of two days’ collecting. The whole region about Minneapolis when more thoroughly worked only furnished 30 saxicolous lichens, and the whole of Fayette county, Iowa, only 50. The latter region is surely better than the average for rock lichens, and I have studied it for six years. Probably however, the fact that I have not looked Fink: ROCK LICHENS OF TAYLORS FALLS. 3 so closely in the last two localities for lichens not strictly char- acteristic of rocks, but still growing on them occasionally, about offsets the limited time spent in examining the area form- ing the basis of the thoughts here presented, so that the numbers given above, save for difference in areas covered in the three instances, may still be taken to represent, approximately, rela- tive richness in rock lichens. But again, the Taylors Falls area, with one possible exception, gave me more such lichens collected in the two days than are recorded in any State list hitherto published. The igneous rocks at Taylors Falls are essentially like those which outcrop occasionally between this place and Lake Su- perior, and the fact that Taylors Falls is practically the south- ern limit of outcrop of these rocks furnished the first suggestion of the interest involved in an analysis of their lichen-flora. Of the 79 species and varieties collected, only 8 are plants not yet found further south in Minnesota or lowa. These 8 I have also found along Lake Superior, and they are species not com- monly occurring further south except at high elevations. These are Bratora rufonigra Tuck., Lecidea albocerulescens Scher., three forms of Bucllia petrea (Flot., Koerb.) Tuck., Unedzle- carta atllentd Tuck., Nephroma helveticum Ach. and Ephebe solida Born., all forms found on the igneous rocks and none of themon the sandstone. Subtracting the 8 species and varieties leaves 58 rock lichens, nearly all of which occur in other por- tions of the southern half of Minnesota, but not all on rocks. Those not known to flourish on rocks in other parts of the State grow on them here under unusually favorable conditions to be explained below. What has already been stated, especially the last paragraph above, merely suggests the problems of interest which I shall attempt to discuss and which involve a knowledge of geological conditions present and past, as well as acquaintance with lichen- species and their distribution in general. For the geological data concerning this area, I have relied largely on the researches of Dr. Charles P. Berkey, who has recently studied the region including Taylors Falls in detail, and who is therefore especially able to give the information needed for my purpose. The ques- tions which I shall consider below are those which thrust them- selves upon me as I observed and studied this extremely in- teresting lichen-flora; and though the area is a small one, the 4. MINNESOTA BOTANICAL STUDIES. questions involved are, it seems to me, none the less worthy of consideration when we notice that it is one of a series of similar areas where certain floral elements have become isolated and gradually killed out by others. The 8 species commonly found farther north have evidently migrated southward, and there are at least two views as to time and method or cause of migration worthy of consideration. First, the northern species might perhaps have migrated from Lake Superior along the exposures of igneous rocks extend- ing from the lake to four or five miles below Taylors Falls in quite recent times, long after the retreat of the last glacier. Second, they seem undoubtedly to be the remnant of a flora driven south, doubtless from some region far north of Lake Superior, by the advancing glaciers and left stranded on favor- able substrata at Taylors Falls as the southern extremity of a flora migrating south before the glacier or more probably mi- grating north on the return of post-glacial climate in the north temperate zone. The outcrops of the igneous rocks between Lake Superior and Taylors Falls are not frequent enough to make either theory seem very plausible; but the second is reasonable since, under the influence of slow decrease in temperature to the southward, migration would naturally follow increasingly favorable cli- matic conditions in that direction even if the outcrops were not more frequent than now. However, it seems probable that at the time of the first glacial advance the outcrops were much more numerous than now. Also, the rock lichens, now found on the igneous rocks only, doubtless found a foothold on the sandstones on the line of retreat under the more favorable cli- matic conditions of glacial times. The numerous boulders of the same igneous rocks, scattered over the ground by the gla- ciers could help in the advance southward of some of the species since glacial times, but hardly of those seldom or never found on boulders, as the Umbzlicarza listed. Onthe whole, it seems doubtful whether a single one of the 8 northern species has migrated southward in post-glacial times under increasingly unfavorable conditions as to climate and substrata. As the remnant of a lichen-flora driven south by glaciers, these plants must either have been stranded during a late gla- cial advance, as during a slight advance during, or more prob- ably after, the Wisconsin stage; or more probably have been Fink: ROCK LICHENS OF TAYLORS FALLS. 5 driven further south than their present position by each of the earlier stages, or ages as the case may be, and retreated with each return of interglacial conditions. The 8 northern species at Taylors Falls are thus either a few of a former flora which has doubtless partly died out and partly migrated northward, or possibly a few species which migrated to the locality from the mountains to the east and west during a late glacial stage, as stated above, or even after the final retreat of the ice. Igneous rocks are not supposed to have been exposed over the region covered by the glaciers south of the area now under considera- tion at the time of the first glacial advance, but sandstone no doubt outcropped frequently and probably further south than the glaciers extended. For a long period after each glacial retreat the surface was no doubt thickly strewn with rocks left by the melting ice, and these rocks would furnish abundant substrata for a retreat of the saxicolous lichens to the north. These same boulders, now largely covered, would partly remain at the sur- face during interglacial conditions and furnish sufficient foot- hold for the organisms to remigrate during a subsequent ad- vance of the ice, thus taking the place of the sandstone where it was covered by previous drift deposits. Thus several migra- tions, alternately southward and northward, probably followed in succession, and we are studying the last stage in the last northward retreat in this not yet completed series. Of course, it is apparent that the Umdzlzcarza and many other lichens not now found at Taylors Falls might have flourished on the sand- stone and later on the boulders at a time when the climate was more favorable for northern species than now, at and south of the area under consideration, both as to temperature and mois- ture, and that they could have migrated readily enough with the advances and retreats of glacial conditions. What species of the rock lichens were able to endure these cycles of migration and what were killed out is not easy to conjecture. However, it seems certain that the region was left barren of such life and repopulated several times, and it is extremely probable that enough species survived the migrations, or possibly in part flowed in from the east or west as stated above, to give an arctic or subarctic flora at Taylors Falls for a time after the close of the ice age. Since the time when this last northern lichen-flora became es- tablished in the region about Taylors Falls, there has been a 6 MINNESOTA BOTANICAL STUDIES. gradual change toward a lichen-flora characteristic of the north- ern United States at the present time. No doubt the 8 northern species now found on the igneous rocks are being rapidly re- placed by the more numerous species, which are better adapted to present climatic conditions. With the exception of the Veph- roma, the 8 species persisting, all lichens confined wholly to rocky substrata, or essentially so, and being therefore favorably situated as to substrata, have persisted longest against un- favorable climatic conditions and the onslaughts of the species which are to-day surely replacing them. Vephroma, which is arboricolous as well as rupicolous, furnished only a few small, sterile specimens clinging to mossy rocks. phebe was seen in one spot only, and, so far as I could ascertain, Umédz- licaria persists only in a few cool, damp or shaded spots. The three species named above, not closely attached to the rocks, would naturally succumb to unfavorable conditions sooner than those named below, and all of the three former are sterile and apparently just on the verge of extermination in the locality. The other three species all grow closer to the rocks and are all abundantly fertile. Bzatora seems to be rare and is not strictly crustaceous as are the other two. uella is the most common of the 8 northern species and is one of two that would be expected to persist longest because of its strictly crustaceous habit. - Zeczdea is as thoroughly crustaceous, but not so common as the Buellva. It must be noted that the latter plant shows locally the strong tendency to vary so characteristic of organisms attempting to adapt themselves to change in environ- ment. Doubtless this variation has aided the plant somewhat in succeeding best of all the present or former more northern species of the locality against adverse climatic conditions. Whether or not the three forms of the species listed arose from one in this locality has no particular bearing in the matter as could be easily shown. Also the distribution of the three forms is so little known that knowledge as to which particular form is most common locally would not show whether it is one most commonly persisting in temperate regions or not. It is interesting to note the time involved in the establishment of the Arctic flora and the change from this to the present essentially temperate flora. According to Professor N. H. Winchell’s views as to the recession of St. Anthony falls, the final retreat of the glaciers from the region occurred about 8,000 Fink: ROCK LICHENS OF TAYLORS FALLS. 7 years ago. ‘Thus it seems that the succeeding 8,000 years must have sufficed for the establishment of a more or less rich Arctic flora and the gradual change to present floral conditions. The relative times involved in the establishment of the first flora and the gradual change to the present cannot be arrived at, since the richness of the first cannot be known, and we cannot yet be sure that a portion of the species migrating southward were not killed out in some portions of the series of migrations, so that some portion of the northern species that became established in the locality would have to migrate toward the center of the con- tinent from the southward-extending mountains already men- tioned. Light on this last supposition, which can only be fully obtained, it seems to me at present, by a study of the lichen flora of the British possessions far to the north of Minnesota, would be extremely interesting. The absence of the 8 northern lichens from the sandstone may be easily explained, since it seems that the present sand- stone surfaces exposed between Lake Superior and Taylors Falls are largely or entirely due to post-glacial erosion. If some of these surfaces are admitted to be as old as the time of the last glacial retreat, doubtless Arctic species grew on them at some time subsequent to that retreat. If this be true, it is yet easy to account for the failure of these lichens to persist on the sandstones as well as on the igneous rocks, since the lichen-flora of these porous and easily eroding surfaces must be a compara- tively changeable and transient one, so that whatever such species once inhabited them would now be replaced by species more characteristic of present climatic conditions. After the final retreat of the ice and the change to present conditions of temperature and moisture began, the rapidly eroding surfaces would begin to lose their northern species and be resupplied by those at hand on other substrata at once, while those on igneous rocks could be replaced, mainly, at least, only by a fierce and long-continued struggle between the Arctic and temperate floral elements. The large number of species found on the sand- stones is at first surprising, for while the igneous rocks are much richer in individual lichens, they show no appreciable advantage in species. The softer texture of the sandstone, which caused the more rapid destruction of the species growing under un- favorable climatic conditions, has also brought this condition. To be a little more explicit, while on account of their rapid ero- 8 MINNESOTA BOTANICAL STUDIES. sion a large number of individuals cannot become established on them and remain long enough to constitute a flora rich in in- dividuals, yet because of the porous character of the sandstone more of the species characteristic of temperate regions have doubtless already become established on them in one place or another than on the igneous rocks. While the 8 species and varieties so fully treated above are of special interest there are some thoughts concerning the other 70 (excluding the last one listed) that must not be lost sight of. As to distribution they are a heterogeneous group, 30 of them being pretty generally distributed over the United States and Canada, 24 being limited so far as their distribution is known to the territory east of the Rocky mountains, 7 being thus far found only in the northern United States and Canada and 5 oc- curring throughout the United States. The North American dis- tribution of the last 4 here considered is so little known that nothing can be definitely stated of it. Of these 70 all but 4 or 5 occur on rocks in some other portion of North America, but only 15 are strictly rupicoline. Of the other 55, some, though more characteristic of rocks, are more or less frequently found growing on other substrata; and others actually prefer other substrata and are growing on rocks here under unusually favor- able conditions. These lichens, like the others, are of course the descendants of a race that has migrated several times. Nearly all of them being species also occurring in Europe, it is certain that they were represented by like species during early Tertiary times, far to the north where our continent was then connected with the Eastern continent on both sides. The coming of a cooler climate and finally of glacial conditions inaugurated the series of migrations. Finally after the last retreat of the glaciers began, the 55 species, because of their adaptation to more than one substratum, would follow the retreat more surely and more rapidly and thus more certainly and sooner reach a given locality and begin to replace a flora growing under unfavorable conditions. ‘To just what extent the arctic flora would become established before these species would come in and begin to re- place it can not be stated since the rate of retreat of the glaciers relative to the rate of migration of essentially stationary organ- isms is not known. In the second paper of this series, I accounted for the com- parative scarcity of lichens about Minneapolis by dryness of Fink: ROCK LICHENS OF TAYLORS FALLS. : 9 climate and stated that, were it not for peculiarly unfavorable conditions as to rock-formations, this explanation would require a larger per cent. of the total number of lichens found there to occur on the rocks because of greater amount of moisture near the ground. The annual precipitation at Osceola Mills for the last six years has been 31.271 inches, while at St. Paul it has been 28.997 inches. The former place being only seven miles from Taylors Falls, the figures may be taken to show that the precipitation in the locality now considered is about 2.274 inches more per annum than that at Minneapolis. Hence we have at Taylors Falls essentially the same conditions as to mois- ture of atmosphere as at Minneapolis. However at the former place we have the extensive rock-formations necessary for the establishment of the plants, and we find further that the igneous rocks are favorably situated for lichen development in that they lie along a river course formerly better shaded than now and where moisture has been abundant in spite of comparative dry- ness of atmosphere. Also this flora was doubtless largely es- tablished when the climate was not so dry as now and is persist- ing against conditions less favorable than formerly existed. Moreover the 8 persisting northern species add to the number strictly characteristic of present climate and give the locality a further advantage over Minneapolis and vicinity. Doubtless study of the whole lichen-flora about Taylors Falls would show that between 30 and 50 per cent. of the lichens grow on rocks as compared with 12 per cent. at Minneapolis. The slight ad- vantage in annual precipitation of moisture for the former lo- cality, of course, adds slightly to the relative richness in rock lichens, but this is insignificant as a cause when compared with the elements considered above. Another objection of doubtful value to the first proposition suggested to account for the invasion of the northern rock-floral elements is that, though there is a continuous line of conifers from Lake Superior to Taylors Falls, the northern tree-inhab- iting lichens are wholly absent at Taylors Falls, or so scarce as to escape notice. The coniferous trees are not so conspicu- ous a part of the flora at Taylors Falls as in Pine county, fifty miles north. No doubt at least a part of the tree lichens char- acteristic of more northern regions, and now almost certainly to be found in Pine county, have extended down to this location in post-glacial times, as the conifers are abundant from Lake 10 MINNESOTA BOTANICAL STUDIES. Superior down to the southern part of this county, and with substrata abundant, they could do so in spite of unfavorable climatic conditions. They have apparently failed to advance as far as Taylors Falls, because of favorite substrata becoming somewhat scarce, and an increase of unfavorable conditions as to temperature and precipitation. JI am aware that the glaciers probably retreated slowly enough so that forests could spring up and furnish substrata for the retreat of species driven south in glacial times, before they would die out at the south on ac- count of the return of warm climate, and that whatever northern tree lichens exist in the pineries fifty miles north, could be ac- counted for, wholly or in part, as having migrated from the south. Yet I am quite convinced that there has been a circula- tion of arboreous lichen-floral elements, between Lake Superior and Pine county, in post-glacial time, which has not extended to Taylors Falls, to any easily observable extent surely, though conditions as to substrata are much more favorable for such lichens to move southward from the lake than for the rock lichens. Not a single species of northern lichen was found in the rock crevices or soil studied. I have noticed how in regions recently burned the soil becomes literally covered in places by lichens of various genera in five to fifteen years, and there can be no doubt that earth lichens took possession of the glacial drift rap- idly after the retreat of the ice began. However, from the very fact that lichens spring up rapidly on earth, the species charac- teristic of temperate climate would the more quickly take pos- session of the present limited amount of soil available for lichen growth, and whatever additional amount that was available when the strife began between arctic and temperate earth lich- ens, and the more rapidly kill out the northern species once in- habiting the drift. A consideration of the statements made in the last two para- graphs and various other portions of this paper points to the conclusion that a study of the whole lichen-flora of the area between Taylors Falls and Lake Superior is essential to a bet- ter understanding of the problems herein considered. In the next paper of this series, in which I shall consider the lichen- flora of the Lake Superior region, I shall be able to show ad- ditional reasons for the study of this territory. It is one of rapid transition in lichen-flora, and after a study of the areas to Fink: ROCK LICHENS OF TAYLORS FALLS. a Tf! the north and south of it, questions of extreme interest have been suggested to me which can only be solved by a study of this flora. The principal conclusions are as follows: (1) The region considered in this paper is an important one for the study of lichen-flora because of position, and geological relations past and present. (2) The flora considered is one of great interest as to origin and present composition and as to evidence of struggle between flora elements. (3) The present lichen-flora is composed of arctic, sub-arctic and temperate florae elements in which the last have long since gained the advantage and are killing out the others. (4) It is not supposed that the northern species migrated south in post-glacial times, but rather that this flora is one that followed the last retreat of the glaciers and was for a time essentially arctic, having since changed to its pres- ent composition. (5) Reasons for the above conclusions are as follows: (2) Southward migration would more naturally result from the decrease of temperature to the south inaugurated by the on-coming of a glacial cli- mate and would thus go on even though suit- able substrata might be somewhat scarce. (4) But during the glacial advances rocky substrata were doubtless more numerous than now, a condition adding to the ease of migration. (c) Under the influence of increasingly favorable cli- matic conditions to the south, the plants would take more easily to unfavorable substrata and migrate more readily on this account also. (d) The 8 northern lichens are all but one essentially rock lichens and are, therefore, the ones that would be expected to persist longest. (6) The northern floral elements considered may have been driven south during a late glacial advance not extending quite to the region, but more probably have been forced to migrate further south several times and migrate north as many times. 12 MINNESOTA BOTANICAL STUDIES. (7) The migrating plants may have been in part or wholly killed out in some part of the series of migrations south- ward in the Mississippi Valley, so that the present north- ern floral element would have to flow in from the moun- tains to the east and west, but more probably found sufficient substrata and were not killed out in the south- ward migrations in the valley. (8) The time involved in the change from arctic and sub-arctic to temperate flora is probably about 8,000 years. The re- lative times occupied in the establishment of the northern flora and the change to the temperature one can not be estimated at present. (9) The absence of the northern floral elements from the sand- stone is due partly to the fact that many of the surfaces of sandstone exposures are post-glacial. If some are as old as the last retreat of the glaciers from the region, the absence is still easily explained since the surfaces are easily eroded and porous so that floral changes go on rapidly on these rocks. (10) Lichens are not individually numerous on the sandstones because of this easily eroding nature which causes rapid change and destruction; yet a large number of species become established in one place or another on them be- cause lichens quickly gain a foothold. (11) The other 70 lichens of the rocks are not so character- istically rock lichens and would migrate more easily and rapidly, and the more quickly reach a locality and re- place an established flora existing under unfavorable conditions, because not confined to one substratum. (12) The rock lichen-flora of the locality is extremely rich be- cause of abundance of rocks, location in a river valley where shade and moisture have been plentiful and geo- graphically where the 8 northern species have persisted to increase the number more characteristic of present climatic conditions. (13) In substantiation of the method used to account for the present composition of the flora, I have attempted to show that the absence of tree and earth lichens from the local- ity tends to prove its correctness. (14) In view of work already done at Taylors Falls and along Lake Superior, the study of the region of rapid transition in lichen-flora between is greatly to be desired. » Fink: ROCK LICHENS OF TAYLORS FALLS. 13 I am under great obligations to Dr. Charles P. Berkey for in- formation concerning present and past geological conditions of the area studied. My thanks are also due to Professor Conway MacMillan for data concerning the distribution of the Conifers between Taylors Falls and Lake Superior and to Mr. Geo. H. Hazzard, of Taylors Falls, for the figures of annual precipita- tion of moisture in the vicinity of Taylors Falls. LIST OF SPECIES AND VARIETIES: 1. Ramalina calicaris (L.) Fr. var. farinacea Scuar. On igneous rocks and sandstone, no. 83. 2. Usnea barbata (L.) Fr. var. florida Fr. On igneous rocks and sandstone, no. 45. 3. Usnea barbata (L.) Fr. var. rubiginea Micux. On sandstone, no. 117. 4. Theloschistes lychneus (Nyv.) Tuck. On igneous rocks and sandstone, no. 64. 5. Theloschistes concolor (Dicxs.) Tuck. On sandstone, no. I02. 6. Parmelia perforata ( JAca.:) Acu. On igneous rocks, no. 71. Not previously reported from Minnesota. 7. Parmelia crinita Acu. On igneous rocks, no. 66. 8. Parmelia borreri Turn. On sandstone, no. I11I. 9g. Parmelia saxatilis (L.) Fr. On igneous rocks and sandstone, nos. 52 and 67. 10. Parmelia olivacea (L.) Acu. On igneous rocks, no. 60. 11. Parmelia caperata (L.) Acu. On igneous rocks and sandstone, no. 50. 12. Parmelia conspersa (Euru.) Acu. On igneous rocks and sandstone, no. 49. 13. Physcia speciosa (WuLF. Acu.) Nyt. On igneous rocks and sandstone, no. 63. 14 MINNESOTA BOTANICAL STUDIES. 14. Physcia aquila (Acu.) Nyw. var. detonsa Tuck. On igneous rocks and sandstone, nos. 42 and 86. Not previously reported from Minnesota. 15. Physcia pulverulenta (Scures.) Nyt. On igneous rocks and sandstone, no. 73. 16. Physcia stellaris (L.) Tuck. On igneous rocks, no. 2. 17. Physcia tribacia (Acu.) Tuck. On igneous rocks, nos. 55 and 77. 18. Physcia cesia (Horrm.) Nyt. On igneous rocks, no. 30. 19. Physcia obscura (Enru.) Nyt. On igneous rocks, nos. 5 and 47. 20. Pyxine sorediata Fr. On igneous rocks, no. 48. 21. Umbilicaria dillenii Tuck. On igneous rocks, no. 87. 22. Nephroma helveticum Acu. On igneous rocks, no. 26. Not previously reported from Minnesota. 23. Peltigera pulverulenta (Tayu.) Nyt. On earth among igneous rocks, no. 15. 24. Peltigera rufescens (NEck.) Horr. On earth among igneous rocks, no. 17. 25. Peltigera canina (L.) Horr. On earth among igneous rocks and on sandstone, nos. i6 and Baie 26. Peltigera canina (L.) Horrm. var. spuria Acn. On sandstone, no. 119. 27. Peltigera canina (L.) Horr. var. sorediata Scu@r. On sandstone, no. 96. 28. Pannaria languinosa (AcH.) KorErs. On igneous rocks and sandstone, no. 20. 29. Pannaria microphylla (Srv.) DE.is. On igneous rocks and sandstone, no. 35. Not previously reported from Minnesota. 30. Ephebe solida Born. (?) On igneous rocks, no. 59. The specimens were sterile and must be regarded as uncer- Fink: ROCK LICHENS OF TAYLORS FALLS. 15 tain as to species. I found the same plant in three or four lo- calities in northern Minnesota. Not previously reported from Minnesota. 31. Collema pulposum (BEeRNu.) Nyt. On earth among igneous rocks, no. 23. 32. Collema flaccidum Acu. On igneous rocks, no. 43. 33. Leptogium tremelloides (L. Fiz.) FR. On igneous rocks, nos. 18, 25, 29 and 61. Not previously reported from Minnesota. 34. Leptogium chloromelum (Srv.) Nyu. On sandstone, no. 109. Not previously reported from Minnesota. 35. Placodium elegans (Linx.) DC. On igneous rocks and sandstone, no. 53. 36. Placodium cinnibarinum (Acu.) Anz. On igneous rocks, no. 31 and 62. 37. Placodium aurantiacum (Licur) Narc. and Hepp. On igneous rocks and sandstones, no. 57. A form with scanty thallus and biatorine apothecia, appear- ing much like the next. 38. Placodium cerinum (Hrepw.) Nagc. and Hepp. var. sid- eritis Tuck. On igneous rocks, nos. 38, 39 and g1. 39. Placodium vitellinum (Euru.) Narc. and Hepp. On igneous rock and sandstone, no. 3. 40. Lecanora rubina (ViLu.) Acu. On igneous rocks and sandstone, no. 51. 41. Lecanora subfusca (L.) Acn. On igneous rocks and sandstone, nos. 12 and 115. 42. Lecanora subfusca (L.) Acu. var. coilocarpa, Acu. On sandstone, no. 108. 43. Lecanora atra (Hups.) Acn. (?) On igneous rocks, no. 4. Not previously reported from Minnesota. 44. Lecanora hageni Acu. On sandstone, no. 118. 45. Lecanora varia (Enuru.) Nyt. On igneous rocks and sandstone, no. 68. 16 MINNESOTA BOTANICAL STUDIES. 46. Lecanora varia (Euru.) Ny. var. symmicta, Acu. On sandstone, no 103. 47. Lecanora cinerea (L.) SomMERF. On igneous rocks and sandstone, nos. II, 19, 34 and 70. 48. Lecanora cinerea (L.) Sommerr. var. levata, Fr. On igneous rocks, no. 14. Not previously reported from Minnesota. 49. Lecanora fuscata (Scurapb.) TH. FR. On igneous rocks and sandstone, nos. 33, 56, 94, and 106. Not previously reported from Minnesota. 50. Rinodina sophodes (Acu.) Nyt. On igneous rocks and sandstone, nos. 59 and 105. 51. Pertusaria velata (TurN.) NYL. On sandstone, no. 95. 52. Pertusaria communis DC. On sandstone, no. I12. 53. Urceolaria scruposa (L.) Nyt. On igneous rocks and sandstone, no. 9. 54. Stereocaulon condensatum Horr. On sandstone, no. 93. Not previously reported from Minnesota. Also not listed before west of New England. 55. Cladonia mitrula Tuck. On sandstone, no. 99. 56. Cladonia cariosa (AcH.) SPRENG. On sandstone, 114. 57. Cladonia pyxidata (L.) Fr. On earth among igneous rocks, no. 65. 58. Cladonia squamosa Horr. On earth among igneous rocks, nos. 72 and 78. Not previously reported from Minnesota. 59. Cladonia cespiticia (PERs.) FL. On earth among igneous rocks and on sandstone, nos. 68" and IOI. 60. Cladonia fimbriata (L.) Fr. var. tubeformis Fr. On sandstone, nos. 104 and 110. 61. Cladonia gracilis (L.) Nyu. On earth among igneous rocks and on sandstone, no. 74. Fink: ROCK LICHENS OF TAYLORS FALLS. U7 62. Cladonia gracilis (L.) Nyv. var. verticillata, Fr. On sandstone, no. 116. 63. Cladonia delicata (EuRH.) FR. On old wood among igneous rocks, nos. 79* and 67. Not previously reported from Minnesota. 64. Cladonia rangiferina (L.) Horr. On earth among igneous rocks, no. 81. 65. Cladonia rangiferina (L.) Horr. var. alpestris L. On earth among igneous rocks, no. 82. 66. Cladonia macilenta (Euru.) Horrm. On old roots among igneous rocks, no. 79” and So". 67. Biatora rufonigra Tuck. On igneous rocks, no. I. Not previously reported from Minnesota. 68. Biatora coarctata(Sm. Ny.) Tuck. var. brajeriana ScHa@R. On sandstone, no. 92. Not previously reported from Minnesota. 69. Biatora myriocarpoides (NyL.) Tuck. On sandstone, no. 100. Habitat unusual, but I cannot distin- guish sufficiently between this and my wood specimens to separate them. Not previously reported from Minnesota. 70. Lecidea albocerulescens (WuLF.) ScHa@rR. On igneous rocks and sandstones, no. 27. Not previously reported from Minnesota. 71. Buellia spuria Arn. On igneous rocks, no. 22. Not previously reported from Minnesota. 72. Buellia petrea (FLot., Korrs.) Tuck. On igneous rocks, no. 58. Not previously reported from Minnesota. 73. Buellia petrea (FLor., Korrs.) Tuck. var. montagnei PLUCK. On igneous rocks, no. 89. Not previously reported trom Minnesota. 74. Buellia petrea (Fiot., Korrs.) Tuck. var. grandis FLOERK. 18 MINNESOTA BOTANICAL STUDIES. On igneous rocks, no. 89%. Thallus coarser, more crowded and lighter colored. Hypothallus deficient. Not previously reported from Minnesota. 75. Endocarpon miniatum (L.) Scua#r. On igneous rocks and sandstone, no. 41. 76. Endocarpon hepaticum Hrepw. On sandstone, no. 97. 77. Verrucaria fuscella Fr. On igneous rocks, no. 21. 78. Verrucaria muralis Acu. On sandstone, no. 107. 79. Thelocarpon prasinellum Nyv. On old boards, no. 113. Not previously reported from Minnesota. II. A METHOD OF DETERMINING THE ABUN- DANCE OF SECONDARY SPECIES. RoscoE PouND AND FREDERIC E. CLEMENTS. In determining the abundance of species, appearances are extremely deceptive. One who has worked over the prairies for many seasons comes to think that he can pick out instantly the most abundant secondary species. Long continued obser- vation in the field stamps a picture on one’s mind, and it seems a simple matter to pick out the several species and to classify them in the several grades of abundance with reasonable ac- curacy. As a matter of fact, this is not possible. After more than ten years of active field work on the prairies, it seemed to the writers that the mental pictures acquired was approximately sufficient to make the reference of the commoner secondary species of prairie formations to their proper grades an easy task. When actual looking at the prairies as the season permitted appeared to confirm the picture already formed, this seemed certain. Closer analysis of the floral covering proved that the conclusions formed from looking at the prairie formations and from long field experience, without actual enumeration of indi- vidual plants, were largely erroneous. ‘The psoraleas, prairie clovers and blazing stars would probably occur to all as among the most abundant of the secondary species in the vernal, estival and serotinal aspects of the prairies respectively. When we first addressed ourselves to the task of assigning to each of the various prairie species its proper degree of abundance, it oc- curred to us at once that we could take a certain species, or certain species, as types for each grade, and use these species as standards by which to measure the others. It proved in the end that the species selected, though of the commonest occur- rence and hence familiar from daily observation, were in many cases referred to wrong grades as compared with other species, no less common, but for some reason not so prominent. The difficulty is that the species which appear most prominent in the constitution of the prairies are not necessarily the most abundant. 20 MINNESOTA BOTANICAL STUDIES. The prominent-flowered blazing stars and prairie clovers make a much greater impression on the eye than species which are far more abundant, and the same thing is true to a less degree of many other species. To insure accurate or even approximately accurate results, it is necessary to resort to some method of actual count. Actual count is usually practicable only when copious, gre- gario-copious or sparse plants are in question. But it is only with respect to such species, which are as a rule secondary in formations, that it is important to determine minutely the grade of abundance manifested. During the past season, in order to determine the actual quan- titative relations of the copious and gregario-copious species, we have made a large number of enumerations of the individual plants of each secondary species present in plots five meters square in characteristic formations of each of the four phytogeo- graphical regions represented in Nebraska. The plot used, five meters square, is as large as can be used to insure accuracy in counting. The deficiences resulting from the small size of the plots are corrected by taking a large number of plots at each station and averaging the results. There is a surprisingly close agreement in figures obtained from plots in widely separated stations in the same district, provided reasonable care is taken to locate them in typical situations. By way of illustration, a number of observations are given in full. These are not averages, but are the actual counts as taken in the field. The two immediately following were taken on the prairie 14 miles northeast of Lincoln in the prairie grass forma- tion (Sporobolus-Koeleria-Panicum). ‘The second was made about 400 yards distant from the first. Amorpha canescens : E 4 : ; ; 387 Aster multiflorus . : . : . ) 228 Antennaria campestris (16 Ratehess : : ; 209 Solidago rupestris . : : : : 5 » IGE flelianthus rigidus : : : : : d 97 Kuhnistera candida : ; ; ; 5 . ae Kuhnistera purpurea . ; ; ; 3 : 21 Brauneria pallida . ; ‘ : : : | 02a Solidago rigida . ; : : : : ; 19 Kuhnia glutinosa . : ‘ , ; ; : 8 Comandra umbellata . : : : ; é 7 Rosa arkansana ; ; : ; : Y E 2 Pound and Clements: METHOD OF DETERMINING SPECIES. 21 (2) Amorpha canescens ‘ j a ; : , 368 Aster multiflorus . , , : e305 Antennaria campestris (6 patches) . . : 154 Solidago rupestris . ; : . : - 005 flelianthus rigidus , : : ‘ - A 63 Kuhnistera candida 3 : : : ; : 26 Kuhnistera purpurea . : : : ; : 22 Brauneria pallida ‘ : : : : 19 Solidago rigida ’ : : F ; ; _ ¥6 Rosa arkansana 8 Solidago rigtdiuscula Gerardia purpurea Laciniaria scariosa NWN — Erigeron ramosus Linum rigidum was prominent, but did not occur in either of the plots, and in comparatively few of those laid out. In a large number of plots, Amorpha canescens averaged 309, Aster multifiorus 275, and Antennaria campestris 12 patches and 145 individuals. Enumerations were also made in the same formation in the transition area between the prairie region and the sand hill re- gion. The following example is one of a number made south of Broken Bow (Custer County). The formation is the ordi- nary prairie grass (Sporobolus-Koeleria-Panicum) formation, modified somewhat on account of the sandy soil. Amorpha canescens ; ‘ : ; : : 291 Aster multiflorus . : : F é 28228 Kuhnistera candida an ; : : . : 22 Solidago rupestris . ; , , - ; ; 21 Brauneria pallida - : - : : : 7 Helianthus rigidus . : : s . P é 12 Kuhnia glutinosa : . ; ; : : 5 The marked decrease in the number of secondary species and in the abundance of each is characteristic of this transition area. Another count, made where the prairie grass formation was giving way to the buffalo grass formation on the one hand, and to the bunch grass formation on the other, is interesting. While the prairie grasses (species of Sporobolus, Koelerta cris- tata and Panicum Scribnertanum) were controlling, there was a strong admixture of Bouteloua oligostachya, and two bunches pap MINNESOTA BOTANICAL STUDIES. of Andropogon scoparius occurred in the plot. The locality was about six miles northwest of the preceding. Amorpha canescens : ; 4 : . ; 192 Kuhnistera occidentalis (?) . E 4 : “S26 Solidago rupestris ( : ; : : : 18 Artemisia gnaphalodes . ; ; ; : wi Dg Solidago mollis. : : ‘ ‘ : ; 12 Solidago rigida ; ; : : : : ; 3 flelianthus rigidus . : : : ‘ : 2 The following count, made in the buffalo grass formation, about two miles beyond the one last set forth, shows the latter formation as affected by the near proximity of the prairie gras, formation. ‘The number of secondary species, small as it is, is very large for that formation. Where this count was mades the dominant grass was Louteloua oligostachya. ‘The only. other grass was Schedonnardus paniculatus, represented by two small patches. Plantago Purshit . d : ; , ; See: Solidago mollis . : : : : £ : ree Lygodesmtia juncea : f , a ; : 5 Lepachys columnarts (dwarf) Etriocarpum spinulosum . Grindelia squarrosa . Kuhnia glutinosa Lesquerella argentea = x S NN WC The constant diminution in the number and abundance of secondary species as one passes from the prairie grass formation of the prairie region to the buffalo grass and bunch grass forma- tions of the transition area and of the sand hill region is well illustrated by these figures. The difference betweeen the prairie grass formation in its ordinary situations and in the transition area, and between the buffalo grass formation of the ‘‘range” and the same formation in the transition area is better shown by figures obtained from such enumerations than in any other way. Many other examples of the efficacy of this method in representing changes in the floral covering as one passes from one district to another might be given. The method of actual enumeration of the individual plants present in plots of a given size makes accurate limitation of the several grades of abundance possible. Of course, this has noth- Pound and Clements: METHOD OF DETERMINING SPECIES. 23 ing to do with the mode of disposition of individuals. But given a copious, gregario-copious or sparse species, there still remains something more to be said before the abundance of the species is fairly indicated. Collation of the results of a large number of enumerations has shown that six grades of copious plants may be recognized readily. The first, in which the average number of individuals in a plot five meters square exceeds 200, corresponds to copious.’ As examples, there may be cited from the prarie formations Asmorpha canescens, with an average of 309 in the prairie region, Aster multiforus with an average of 275 in the prairie region and about 230 in the sand hill region; from the herbaceous layer of woody formations, Verdbesina alternifolia (which is almost gregarious at times), with an aver- age of 245. To the second degree (copious’) those species may be assigned in which the average number of individuals in a plot is from 150 to 200, such as Plantago Purshit (162) in the Peppergrass-Cactus formation in the transition area between the sand hill region and the foot hill region. Those species with an average ranging from 100 to 150 may be assigned to the third degree (copious’®). Examples are: Aster sagittifolius, which has an average of 133 in the herbaceous layer of the Bur- oak-Elm-Walnut formation in the Mississippi basin region and Solidago rupestris, which has an average of 104 in the Sporo- bolus-Koelerta-Panicum formation in the prairie region. In the fourth degree (copious*) those species may be included which have an average of from 50 to 100, such as Glycorhiza lepidota in the river valleys in the sand hill region, where its average is 83. All of the foregoing are of sufficient abundance to be included in the general term ‘‘ copious,” taking the latter to represent a quantitative idea as well as the manner of associ- ation of the individuals. Where the average falls below 50 and exceeds 5, we call the species ‘‘ subcopious.” Comparison and collation of statistics has shown that subcopious species fall into two groups, in one, which we call subcopious,' the average does not fall below 15. Examples are: Auhnistera candida in the Sporobolus-Koeleria-Panicum formation in the prairie re- gion, where it has an average of 18, Solidago mollis in the Peppergrass-Cactus formation in the transition area between the sand hill and foot hill regions, where its average is slightly over 20, and Artemzsta guaphalodes in the transition between the prairie and the sand hill regions, where its average is 16. 24 MINNESOTA BOTANICAL STUDIES. Where the average number in a plot is between 5 and 15, the species is called subcopious.” A glance at the list given above will show that these are often very striking components of the prairie formations. Finally, in case the average is below 5 and above .OI, or one individual in ten plots, the species is called “sparse.” Gregario-copious species may be treated in the same way, giving gregario-copious,' etc. ~I 48. 49: Holzinger : MUSCI OF THE INTERNATIONAL BOUNDARY. 43 Mnium cuspidatum Hrepw. Same station as the last. Mnium punctatum Hepw. On the road from Ely to Fall lake (June 8, 1897). At Camp 1, Fall lake (June 8-10, 1897). . Mnium serratum Brip. On a small point of land at the base of the United States peninsula, shore of Basswood lake (June 11, 1897). On Safety island, Lake Saganaga (June 16, 1897). . Timmia bavarica Hess. var cucullata (MicuHx.). On Grand Portage island, north shore of Lake Superior (June 23, 1897). . Atrichum undulatum P. B. On the point of land at the base of Kawasatchong falls, shore of Fall lake, near Camp 1 (June 8-10, 1897). Pogonatum alpinum RoE LL. On Grand Portage island, north shore of Lake Superior (june 235 560;7)). Polytrichum commune L. Same station as the last. . Polytrichum juniperinum WILLp. On the point of land at the base of Kawasatchong lake, shore of Fall lake, near Camp 1 (June 8-10, 1897). . Polytrichum piliferum Scures. On the prairie portage, shore of Basswood lake, near the rapids from Sucker lake (June 12, 13, 1897). . Fontinalis antipyretica Linn. In the river crossing the Grand portage about four miles north of Grand Portage village. Abundant (June 21, 1897). . Fontinalis duriaei Scu. On submerged rocks at the base of Kawasatchong falls near Camp 1 (June 8-10, 1897). Fontinalis holzingeri CARDOT. sp. nova in litt. At the second falls of Granite river ascending ‘from Lake Saganaga (June 17, 1897). ‘¢Du groupe Heterophylle, voisine du /. mzssourica Card., sed foliis rigidioribus, reti firmo, cellulis longi- 44 Bi MINNESOTA BOTANICAL STUDIES. oribus, haud vel vix flexuosis, valde chlorophyllosis, parietibus firmis, distincta.” . Fontinalis hypnoides Harr. ‘‘ forma foliis apice saepe denticulata.”’ In the stream flowing from North lake into Little Gun- flint lake. Abundant at the lower end of the stream (June 20, 1897). . Dichelyma pallescens B. S. At the base of alder trunks growing along the bank of Fall lake, near Camp 1 (June 8-10, 1897). . Neckera oligocarpa B. S. Near Camp 1 at the base of Kawasatchong falls, shore of Fall lake (June 8-10, 1897). Onasmall point of land near the base of the United States peninsula, Basswood lake (June 11, 1897). At the farther end of the portage across the United States peninsula, Basswood lake (June 12, 1897). On Safety island, Lake Saganaga (June 16, 1897). On Grand Portage island, north shore of Lake Superior (Juner235 1 507)). 58. Neckera pennata Hupw. On trees along the road from Ely to Winton, on Fall lake (June 8, 1897). Near Camp 1, on Fall lake (June 10-12, 1897). On the Prairie Portage, shore of Basswood lake, near the rapids from Sucker lake (June 12, 13, 1897). 59. Homalia trichomanoides jamesii (ScuIMP.). Near Camp 1, at the base of Kawasatchong falls, shore of Fall lake (June 8-10, 1897). On the portage from Fall lake to Newton lake (June 10, II, 1897). On a small point of land at the base of the United States peninsula, Basswood lake (June 11, 1897). Nore: This plant has leaves varying strongly toward the typical European form of the species. On Grand Portage island, north shore of Lake Superior (June 23, 1897). Norte: I have carefully studied the plants collected ; have compared them with Professor Macoun’s Canadian specimens sent out under No. 242; also with the /o- Flolzinger : MUSCI OF THE INTERNATIONAL BOUNDARY. 45 mata collected by Professor James, near Franconia, N. H., named //. jamese¢ Schimp., and probably typical material of Schimper’s species, also with HZ. tr7choman- oides from Denmark, collected by Dr. J. Hensen, near Hvals6, in 1883; and I fail to find a single good reason for separating our American forms of AHomadéa with ser- rate leaves from the European Homalia trichomanotdes as a distinct species. Not a single constant character can be established for our plant. I have made microm- eter measurements of the leaf cells of all the specimens examined, and I find them essentially of the same size in the same part of the leaf in all specimens. The American specimens that are fertile show not the slight- est difference from European specimens either in peri- chetial leaves or in perforation of the segments of the peristome along the keel. The only point of difference is the, on the whole, more obtuse apex of the leaves in our American forms. But it cannot fail to attract the at- tention of the student, especially when he reviews a large number of forms in different collections, that while in our American forms the apex is on the whole more rounded, some leaves may be found on every plant which have an apiculate apex. Also the European plants studied show some leaves more rounded at the apex than others. As for the description of the European plant, by European authors themselves, let me cite first from Muscineés de la France by M. Abbé Boulay (1884), p. 150. ‘*Féuilles largement oblongue-elliptiques, con- vexes-cultriformes par le bord superieur, un peu repliées en dessus par linflexion du bord inferieur, brevement apiculées, tres finement denticulées sur tout le contour (dont plus grandes et plus rapprochées ver le sommet ; long. 2, larg.I mm. * * * * * cellules moyennes 8—10 fois aussi 1. q.1.; vers les bords et au sommet, elles sont courtes, rhomboidales; * * * * * Janieres du peristome interne linéares, plus longues que les dents, peu ou millement ouvertes ver la carene.” In Limpricht’s Laubmoose II (1895), p. 715: ‘* Blatter gedrangt, zweizerlig-abstrehend, zuletst abwarts gebo- gen, flach ausgebreiten, unsymmetrisch, aus herablaufen- der, etwas verschmalerter Bariszungen-messerférmung, 46 MINNESOTA BOTANICAL STUDIES. stiimpflich, 1.8-2 mm. lang, und r mm. breit, am Rande der oberen Blatt halfte ausgefressen-gezahnt, am Grunde, an einer Seite eingeschlagen * * * Innerer Peristom * * * in der linie ritzenférmig durch brochen.” Now in Macoun’s Cat. of Can. Pl., 4 (1892), p. 163, the authors of Homalia macouni?, say of it: ** Very nearly allied to Hlomalia trichomanoides; differs in the leaves being longer, rather lingulate, the lowest basal cells yellow, the perichetial leaves more suddenly nar- rowed to a very short acumen, the segments of the per- istome cleft between the articulations.” This is quite all in the line of characterization. Among other localities it is credited to Lake Superior, Drummond’s specimens having been collected there. Both from actual comparison and from the circum- stance of locality, the Lake Superior plants collected by me are reasonably referred to the same plants upon ‘which Homalia macouni is founded. If this inference is correct then the only valid part of the above statements, which stand in place of description, is the first phrase ‘‘very nearly allied to A. ¢trichomanozdes.” ‘The leaf length varies according to European authors themselves. The ‘‘ rather lingulate” form of outline is ascribed by Limpricht to Homalia trichomanoides, when he makes the leaves ‘* zungen-messerformung,” z. é., *¢ lingulate-cultri- form.” As to the ‘‘ lowest basal cells yellow, the peri- cheetial leaves more suddenly narrowed to a very short acumen,” my own close observations fail to verify these two characters, which, if observed by the authors, must have been purely accidental. And as for ‘* the segments of the peristome cleft between the articulations,” this character, judging both the European specimens actually examined, and from the painstaking description of /o- mala trichomanordes made by European authors them- selves, as seen from citations above, is unconditionally conceded to belong to Homala trichomanordes Br. The only tangible difference, the slightly more obtuse leaves it certainly has in common with Dr. James’ own speci- mens of Homalia jamesiz. If now we turn to Lesq. and James’ Manual of Mosses of North America (1884), p. 285, we find not a single positive or new character as- 60. Gi. Holzinger: MUSCI OF THE INTERNATIONAL BOUNDARY. 47 signed to Homalia jamesti, except leaves ‘é striolate lengthwise when dry.” And this point is not borne out by the actual examination of James’ own material. It appears, therefore, that AYomalia jameszi is too close to Hl. trichomanoides ; that Hlomalia macounzz is identical with Homalia jamesiz; that the only difference is found in the more obtuse leaves of our species, which proves to be a variable character, and therefore that it should not stand as a distinct species, hardly deserving the name of a variety. As a variety it must be called: Homalia trichomanoides jamesii (Scurp.). Ff. jamesté Schimp. in Syn. ( yp. 473. FT. macount? in Mac. Cat. (1892), p. 163. The geographical distribution of this variety of Homalia trichomanordes, includes necessarily all the stations cited in Macoun’s Catalogue for AH. macounz? with those given for /7. jameszz in Lesquereux and James’ Manual. Myurella careyana SuLL. On Grand Portage island, north shore of Lake Superior (June 23, 1897). Myurella julacea Scu. At the base of Kawasatchong falls, shore of Fall lake, near Camp 1 (June 8-10, 1897). On Grand Portage island, north shore of Lake Superior (Ghmes23,<1897,). . Leskea nervosa Myr. On the Prairie portage, shore of Basswood lake, near the rapids from Sucker lake (June 12, 13, 1897). . Leskea polycarpa Euru. At the farther end of the portage across the United States peninsula, shore of Basswood lake (June 12, 1897). On the Prairie portage, shore of Basswood lake, near the rapids from Sucker lake (June 12, 13, 1897). . Leskea polycarpa paludosa Scu. On the way from Ely to Winton, shore of Fall lake (June 8, 1897). 48 66. 67. 68. 69. 70. "I. “I to MINNESOTA BOTANICAL STUDIES. Along the shore of Fall lake, near Camp 1 (June 8-10, 1897), abundant. . Anomodon attenuatus HartTm. Shore of Fall lake, near Camp 1 (June 8-10, 1897). On a small point of land, at the base of the United States peninsula, shore of Basswood lake (June 11, 1897). On the Prairie portage, shore of Basswood lake, near the rapids from Sucker lake (June 12, 13, 1897). Anomodon minor (P. BEAuv.) FURN. On the shore of Fall lake, base of Kawasatchong falls, near Camp 1 (June 8-10, 1897). Anomodon rostratus Scu. On the shore of Fall lake, base of Kawasatchong falls, near Camp 1 (June 8-10, 1897). On a small point of land, at the base of the United States peninsula, shore of Basswood lake (June 11, 1897). On the Prairie portage, shore of Basswood lake, near the rapids from Sucker lake ( June 12, 13, 1897). Pylaisia heteromalla Scu. On trees along the shore of Fall lake, near Camp 1 (June 8-10, 1897). On a small point, at the base of the United States penin- sula, Basswood lake (June 11, 1897). Pylaisia polyantha Scu. : On trees along the shore of Fall lake, near Camp 1 (June 8-10, 1897). On the Prairie portage, shore of Basswood lake, near the rapids from Sucker lake (June 12, 13. 1897). Platygyrium repens Scu. Shore of Fall lake, near Camp 1. On dead logs (June 8-10, 1897). Cylindrothecium seductrix SuLLiv. Same locality as above. . Climacium americanum Brip. Same locality as above. Portage from Mountain lake to Moose lake (June 20, LOOy) 73: 79: So. SI. Flolzinger : MUSC1 OF THE INTERNATIONAL BOUNDARY. 49 Thuidium abietinum Scu. Shore of Fall lake, near Camp 1 (June 8-10, 1897). On the Prairie portage, shore of Basswood lake (June Eo. r5, TOQy). On the portage from South-Fowl lake to Pigeon river (June 21, 1897). On Grand Portage island, north shore of Lake Superior (June 23, 1894). . Thuidium recognitum Linps. Shore of Fall lake, near Camp 1 (June 8-10, 1897). . Thuidium philiberti Liver. At camp, shore of Fall lake (June 8-10, 1897). . Brachythecium campestre Scu. Shore of Fall lake near Camp 1 (June 8-10, 1897). . Brachythecium flexicaule Ren. and Carp. On the Prairie portage, shore of Basswood lake (June E25 hay LOOT) . Brachythecium oxycladon (Brip.). Grout. At the base of the United States peninsula, shore of Basswood lake (June 11, 1897). On the Prairie portage, shore of Basswood lake (June Hpk LOOT) « On Grand Portage island, north shore of Lake Superior (June 23, 1897). This last is pronounced a ‘‘ slender form” by Mr. Grout. Brachythecium plumosum Scu.? On the Prairie portage, shore of Basswood lake (June 12 olaa LOOT). Brachythecium salebrosum Scu. Shore of Fall lake, near Camp 1 (June 8-10, 1897). Brachythecium starkei Scu. At the base of the United States peninsula, shore of Basswood lake ( June 11, 1897). . Eurynchium robustum (RoELL.),. At the base of the United States peninsula, shore of Basswood lake (June 11, 1897). On Basswood lake, at the farther end of the portage across the United States peninsula ( June 12, 1897). 50 aa: 84. 86. 87. 88. 89. go. gi. MINNESOTA BOTANICAL STUDIES. Mr. Grout remarks that these plants vary toward £. strigosum ; they are probably only large forms of this species. Eurynchium strigosum Scu. Shore of Fall lake, near Camp 1 (June 8-10, 1897). On the portage from South lake to Rat lake (June 20, 1897). On the portage from South-Fowl lake to Pigeon river (June 21, 1897). On Grand Portage island, north shore of Lake Superior June 23, 1897). Raphidostegium recurvans L. and J. Shore of Fall lake, near Camp 1 (June 8-10, 1897). On the point of land at the base of the United States peninsula, Basswood lake (June 11, 1897). On Safety island, Lake Saganaga (June 16, 1897). . Plagiothecium denticulatum Scu. On the road from Ely to Winton, shore of Fall (hice (June 8, 1897). At Camp 1, shore of Fall lake, near Kawasatchong falls (June 8-10, 1897). Plagiothecium muhlenbeckii Scu. At Camp 1, shore of Fall lake (June 8-10, 1897). Plagiothecium sylvaticum Scu. At Camp 1, shore of Fall lake (June 8-10, 1897). On Grand Portage island, north shore of Lake Superior (June 23, 1897). Amblystegium adnatum L. and J. At Camp 1, shore of Fall lake (June 8-10, 1897). Det. by G. Va Best. Amblystegium serpens Scu. On the portage from Mountain lake to Moose lake (June 20, 1897). Hypnum chrysophyllum Brin. At Camp 1, shore of Fall lake (June 8-10, 1897). Hypnum cupressiforme ericetorum B. S. At Camp 1, shore of Fall lake (June 8-10, 1897). At the lower end of Pipestone rapids, on Basswood lake (June 10, 11, 1897). Flolzinger : MUSCL OF THE INTERNATIONAL BOUNDARY. bl 92. Hypnum crista-castrensis L. On the road from Ely to Winton, on Fall lake (June 8, 1897). At Camp 1, on Fall lake (June 8-10, 1897). At the lower end of Pipestone rapids, on Basswood lake (June 10, 11, 1897). At the base of the United States peninsula, Basswood lake (June 11, 1897). Near Camp 3, at the farther end of the portage across the United States peninsula, on Basswood lake (June £2, 1907). On Prairie portage, shore of Basswood lake (June 12, 13, 1897). On Safety island, Lake Saganaga (June 16, 1897). 93. Hypnum filicinum trichodes Brin. On Grand Portage island, north shore of Lake Superior (June 23; 1607). Dr. Best remarks that this approaches the variety aczcu- linum C. M. and K. 94. Hypnum haldanianum Grey. On the road from Ely to Fall lake (June 8, 1897). At Camp 1, on Fall lake, near Kawasatchong falls (June 8-10, 1897). At the lower end of the Pipestone rapids, on Basswood lake. (june 205) 11,1897): At the base of the United States peninsula, on Basswood lake (June 11, 1897). On the portage from South lake to Rat lake (June 20, 1897). 95. Hypnum hispidulum Brin. On the road from Ely to Fall lake ( June 8, 1897). At Camp 1, on Fall lake (June 8-10, 1897). 96. Hypnum reptile Ricn. At Camp 1, on Fall lake (June 8-10, 1897.) At a small point of land at the base of the United States peninsula, Basswood lake (June 11, 1897). 97. Hypnum schreberi WILLp. At Camp 1, on Fall lake (June 8-10, 1897). At the lower end of the Pipestone rapids, on Basswood lake, near Camp 2 (June 10, 11, 1897). or ho MINNESOTA BOTANICAL STUDIES. At the base of the United States peninsula, Basswood lake (June 11, 1897). At the farther end of the portage across the United States peninsula, Basswood lake (June 12, 1897). On Safety island, Lake Saganaga (June 16, 1897). — At the east end of Gunflint lake (June 20, 1897). On Grand Portage island, north shore of Lake Superior (June 23, 1897). 98. Hypnum uncinatum Hepw. On the road from Ely to Fall lake (June 8, 1897). On the portage across the divide (June 20, 1897). 99. Holocomium splendens Scu. At Camp 1, shore of Fall lake (June 8-10, 1897). At the base of the United States peninsula, Basswood lake (June 11, 1897). On the Prairie portage, shore of Basswood lake (June 12, 13, 1897). On Safety island, Lake Saganaga (June 16, 1897). At the east end of Gunflint lake (June 10, 1897). On Grand Portage island, north shore of Lake Superior (June 23, 1897). 100. Hylocomium triquetrum Scu. On the road from Ely to Fall lake (June 8, 1897). At Camp 1, shore of Fall lake (June 8-10, 1897). At the farther end of the portage across the United States peninsula, shore of Basswood lake (June 12, 1897). On Safety island, Lake Saganaga (June 16, 1897). . At the east end of Gunflint lake (June 20, 1897). On Grand Portage island, north shore of Lake Superior (June 23, 1897). Vil. THE INFLUENCE OF GASES AND VAPORS UPON, THE, GROWTH OF PLANTS. Emit P. SANDSTEN. INTRODUCTION. In recent years considerable attention has been paid by phys- iologists to the influence of various chemical agents upon the growth of plants, and the results thus far obtained seemed to warrant further investigation along this line. The work here- tofore has been confined almost exclusively to the lower plant forms, which are more easily attacked by the difficult technique which is bound up in this kind of inquiries. The recent prelim- inary results of Johanssen (1) were announced shortly after this work was begun, and it was thought advisable to extend the work to cover the phases of the vegetative period as well as rest- ing seeds, etc. ‘To some extent the writer has had in mind the ultimate application of the reactions obtained in practical garden- ing though such results are reserved for verification and further trial. The work was done during the fall and winter of 1897 and 1898 in the laboratories of plant physiology in the Univer- sity of Minnesota under the direction of Dr. D. T. Mac Dougal, to whom the writer is greatly indebted for his valuable advice and kind criticism. MATERIAL AND METHODs. The experiments may be conveniently classified as follows : . The influence of gases and vapors upon seeds. . The influence of gases and vapors upon seedlings. . The influence of gases and vapors upon growing shoots. . The influence of gases and vapors upon resting bulbs, corms, etc. 5. The influence of gases and vapors upon plants growing in water cultures. The reagents used were alcohol, ammonia, carbon bisulphide, SB wWieN H D4 MINNESOTA BOTANICAL STUDIES. chloroform, ether, nitrous oxide and oxygen. Small quantities of alcohol (methyl), ammonia (hydrate), carbon bisulphide, ether and chloroform were placed in tubes inside of closed receivers and allowed to vaporize into the air enclosed. The nitrous ox- ide was the commercial mixture, N,O go parts, N 8.86, O 1.13. In certain experiments the pure gas which had been obtained from ammonium nitrate was used. Commercial oxygen from tanks was used. As a means of control and test of the actual efficiency of the reagents, leaves of Phzlotrza (Flodea) and hairs of Tradescantza, Tomato, Begonia, Pelargonium and Geranium were mounted in an Engelmann gas chamber and subjected to their action. These tests were carried on at a room temperature of 16 to 23° C., and the results noted below are quite in harmony with those given by previous writers. Oxygen. 'The movements of protoplasm are greatly acceler- ated in an atmosphere of free oxygen for five to seven minutes, after which the movements gradually diminish until they cease entirely. Ifthe living cell is kept under the influence of free oxy- gen for considerable length of time it dies, but no apparent change in the structure or behavior of the protoplasm could be noticed. Nitrous oxide (N,O). This gas has the same general effect on living protoplasm as oxygen with the exception that it is less and does not kill the cell even when exposed for several days in an atmosphere of 90 per cent. nitrous oxide. The duration of active movements varies from three to five minutes (Moeller V.). Chloroform and Ether. ‘The action of these two reagents are about the same. Aqueous solutions containing 1/20000 part of reagents at first slightly increase the movements of the proto- plasm. By increasing the strength of the solution the rapidity of movement was also increased, but the reaction time was very much shortened. Strong solution causes vacuolization and par- alyzes the protoplasm. Ammonia. Weak aqueous solution containing from 1/30000 to 1/20000 parts of ammonia does not seem to modify the activity of the protoplasm when subjected to its action for a short time only. Stronger solution produces vacuolization and slightly ac- celerates the movements of the protoplasm for a minute or two. Carbon bisulphide.—The smallest possible quantity that could be introduced arrested all movements. Alcohol.—Aqueous solutions containing 1/20000 to 1/10000 parts of alcohol had no visible effect upon the protoplasm. A Sandsten: INFLUENCE OF GASES AND VAPORS UPON GROWTH. 55 2 per cent. solution excited rapid irregular movements which stopped inside of two minutes. Vacuolization followed rapidly and the cell was killed inside of ten minutes. In the experiments where seeds and seedlings were used, Zea mats, Victa and Phaseolus were employed exclusively. Straw- berry plants of the common cultivated kind were used in the ex- periments with growing shoot and proved well adapted to the work. The strawberry plants were taken from the bed on No- vember 6, 1897, and carefully selected with reference to vigor and equality. Two lots of plants were selected, one lot com- posed of plants one season old, the other composed of plants two seasons old. The plants were placed in three- and four-inch pots respectively November goth, and set in a cold frame where they remained until December gth, when they were taken to the green house and put under the experimental conditions described below. Dormant bulbs and corms of Arzsema, Warcissus, Hyacinth, Tulip, Freesca and Crocus were used for material in the resting stage. In the experiments with gases in nutrient solution in water culture seedlings of Zea mazs were used. : WYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 103 bundles and eight paired xylem bundles. Eventually closed rings of xylem and phloem are formed. At the corners of the stele in the pericycle groups of scleren- chyma are formed. The cells become very thick-walled and each group quite large. The pith is large. The cells are rather thick-walled. No definite perimedullary zone was distinguished. In the material examined cork formation had not commenced. Structure of Epicotyl. The epidermal cells are thin-walled, square in cross section, becoming at length much flattened. There is no collenchyma. The cortex is narrow; the cells are about the same size as those of the epidermis. The endodermis is composed of thin-walled cells. After secondary growth of the stelar tissues it cannot be definitely seen. The cells are about the same size as those of the cortex ; they contain starch. Starch is found at a later time in the various parenchymatous tissues. The number of vascular bundles is variable. Usually there are about twelve. These, at length, fuse to form closed rings of phloem and xylem. The pericycle develops a sheath of sclerenchyma which almost completely shuts in the phloem. The cells were not very thick- walled in the material examined. The pith is large, the cells rather » thin-walled. A perimedullary zone of small-celled parenchyma at length becomes differentiated. No material old enough to show cork formation was examined. Comparison of Structure of Hypo- cotyl and Epicotyl. The hypocotyl differs from the epi- cotyl in having a thicker cortex with a narrow small-celled parenchyma- tous sheath. The cells of the cortex are also larger. Baasieae In its primary stelar structure the aculeata differences are very marked. The Fic. 8. 104 MINNESOTA BOTANICAL STUDIES. hypocotyl has four phloem bundles and eight xylem bundles instead of a large number of conjoint bundles. It has four large groups of stereom instead of a narrow, almost continuous sclerenchymatous sheath. Cercis canadensis. Structure of Hypocotyl. The epidermal cells are rectangular in cross section; the radial diameter is the longer. These cells never become tan- gentially elongated. The cells of the outer cortex are smaller than those within. An indefinite sheath of small-celled paren- chyma similar to that in Parkznsonia can sometimes be recog- nized. The endodermis is small-celled; it remains distinct through the first year. Its cells contain starch. Starch is also present toward the close of the year in the pith. The stele is originally quadrangular. There are four xylem bundles and four phloem bundles. These, at length, develop into closed rings. Four small groups of sclerenchyma make their appearance in the pericycle at an early time and become, at length, consid- erably extended. The pith finally becomes cylindrical. The cells are large and thin-walled. The perimedullary zone is not clearly dif- ferentiated. Cork formation takes place in the cortex either next to or very near the endodermis. It begins sometime before the close of the season. Structure of Epicotyl. The epicotyl is somewhat quadrangular in the early stages. The epidermal cells are rectangular in cross section. ‘The tangential diameter is the greater. There is no hypoderma. ‘The cortex is thin. The cells are all about the same size. The endodermis was not definitely distinguished. In the youngest material examined the phloem forms a closed ring. There are four large primary xylem bundles. There are also some smaller ones. The latter have often only one or two xylem cells. A closed zone of xylem is soon produced. Nearly all the cells of the pericycle become, at length, scler- otic, thus forming an almost continuous sheath with but few parenchymatous cells. Ramaley: HYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 105 y The pith is large-called. A perimedullary zone was not dis- tinguished. Cork formation takes place in the second cortical layer as in Cercis stliquastrum (fide Moeller [1882] ). Comparison of Structure of Hypocotyl and Epicotyl. The epidermal cells of the hypocotyl, when seen in cross section, appear radially, not tan- gentially elongated as in the epi- cotyl. In the former region the endodermis is distinct, the cortex thicker and the sclerenchyma at first differently disposed. The stele of the hypocotyl has originally four phloem bundles and four xylem bundles. The young- est material of the epicotyl which was examined has a closed ring GON of phloem and four large xylem ©) Ss bundles, also a few small groups of == xylem. O Cork formation in the hypocotyl takes place in the lower cortex; in the epicotyl it takes place in the Cereis canadénsis second cell layer of the cortex. Fic. 9. Gleditsia triacanthos. Structure of Hypocotyl. The epidermis is composed of rather thick-walled cells which are oblong in cross section, the long axis being at right angles to the periphery of the section. These cells are eventually elongated in the tangential direction. The cortex is very thick. There is no differentiated hypo- derma, but three or four of the outer cortical layers are com- posed of smaller cells than those below. The endodermis is definite; it is large-celled. In some places it is two layers of cells in thickness. Starch, at first present only in the endodermis, is eventually widely distributed throughout all the parenchymatous tissues. The stele is cylindrical. There are in the young hypocotyl 106 MINNESOTA BOTANICAL STUDIES. eight paired xylem bundles and a large number of groups of phloem. The latter soon grow together, forming a complete ring, while the xylem bundles first fuse in pairs, afterward growing together into a closed zone. In the pericycle, alternating with the paired xylem bundles there are developed four large bands of sclerenchyma which ex- tend so far around that they nearly touch each other. By the end of the first season these become divided into a number of groups by the intercalation of parenchymatous cells. The pith, which is eventually of slight extent, is composed of large-celled parenchyma. Cork formation begins rather early the first season in the third or fourth cell layer of the cortex. Structure of LEpicotyl. The general shape of the epicotyl is originally somewhat hexagonally prismatic; it soon becomes cylindrical. The cells of the epidermis are originally nearly square in cross section. There are numerous, long, curved, pointed epi- dermal hairs. The outer two layers of the cortex become slightly collenchymatous. The other cortical layers are com- posed of parenchyma. The endodermis was not distinguished in material taken in the autumn but in the young epicotyl is quite distinct. The cells are rather large, similar to those of the cortical region but packed with starch. The phloem, in youngest material examined, forms a ring of tissue. There are about six principal xylem bundles which soon fuse. A broken sclerenchymatous ring is formed which resembles that of the epicotyl. No other stereom is, as a rule, produced the first year. The pith is large and composed of cells with unlignified walls. There is a small-celled perimedullary zone. Cork formation takes place in the hypoderma (cf. Moeller it6S215 p93). Comparison of Structure of Hypocotyl and Epicotyl. The hypocotyl differs from the epicotyl in the absence of epi- dermal hairs and of a collenchymatous hypoderma, in the pri- mary structure of the stele, and in its smaller pith. Ramaley : WYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 107 In the hypocotyl there are at first four pairs of xylem bundles and a number of phloem bundles. Four large groups of sclerenchyma soon make their appearance in the peri- cycle. In the epicotyl, on the other hand, a closed ring of phloem surrounds usually about six xylem bundles. No differentiated peri- medullary zone was distinguished in the hypocotyl. The structure of the two regions at the close of the first year differs only in the perimedullary region and pith; the formation of cork having removed the epidermis and hypo- derma. Gleditsia triacanthos PAPILIONACEA. Amorpha fruticosa. Structure of Hypocotyl. The epidermis consists of cells rather small, somewhat thick- walled, square or nearly so, in cross section, at length becom- ing flattened. ‘The cells of the cortex are large; those imme- diately below the epidermis somewhat smaller, but not forming a definite hypoderma. The endodermis of thin-walled cells containing starch re- mains distinct for some time. ‘Toward the close of the first, year its exact position cannot be determined, although it can be lo- cated approximately. A small amount of starch is scattered throughout the cortex, pith and pericycle as well as the endo- dermis, even in the youngest-stage. This is not the case in most species. Later the phloem and the medullary rays also come to be filled with starch. The stele is at first quadrangular, and remains so for a con- siderable length of time. There are four conjoint vascular bundles, and in addition there appear a few small patches of phioem. The bundles soon tend to unite in pairs. Xylem and phloem at length form complete zones. The medullary rays are very numerous; they are one cell in width. 108 MINNESOTA BOTANICAL STUDIES. About the time that the epicotyl has reached its full length four small areas of sclerenchyma appear in the pericycle, one adjoining the phloem of each vascular bundle. These increase somewhat in size, and are still visible in two-year-old material. Numerous isolated sclerenchymatous elements are found scat- tered through the phloem. The pith is composed of rather large cells with thin walls, which soon become lignified. As the plant grows older the pith becomes almost obliterated. No perimedullary zone was dis- tinguished. Cork formation takes place in the outer pericycle, at length cutting off all tissues outside, leaving the bundles of scleren- chyma which are at the inner limit of the pericycle. Structure of Epicotyl. The epidermis is composed of cells nearly square in cross section. These abut directly upon a large-celled, few-layered cortex. There is no hypoderma. A definite endodermis was not distinguished at any time al- though in a very young stage certain starch containing cells were recognized as having the appearance of endodermis; a continuous ring of them was not traced. With the exception of the endodermal and medullary region, starch does not occur until the plant has developed a number of internodes above the epicotyl. The cells of pericycle and phloem are at length filled with starch. The stele is cylindrical from the first. Owing to fusions the number of vascular bundles is variable. There are, however, generally about five or six bundles. The phloem and xylem eventually form closed rings. There is a narrow interrupted ring of stereom at the outer edge of the pericycle. Toward the end of the first season numerous small patches of thick-walled fibers appear in the phloem and the pericycle. The pith is large-celled; it does not decrease appreciably in size as the stem grows older. In the lower part of the epicotyl cork formation takes place in the pericycle below the ring of stereom mentioned above, thus cutting off the cortex and epidermis which soon die and disappear. In the upper part it takes place in the cortex (cf. Moeller [1882], p. 383). This plant shows a distinct ‘* re- gion tigellaire” in two-year-old material. Ramaley: HYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 109 Comparison of Structure of Hypocotyl and Epicotyl. In very young plants the hypocotyl shows a few slightly dif- ferentiated layers of smaller cells in the outer cortex. The epicotyl has nothing of the kind. i The endodermis of the hypocotyl / my is distinct in the early stages, but was | not definitely located in the epicotyl. Starch appears earlier in the hypocotyl and the four large groups of stereom are not represented at all in the epi- cotyl which, however, has an_ inter- rupted circle of the same material. In the former region the stele is at first quadrangular, while always cylin- drical in the latter. Cork formation in the hypocotyl is pericyclic, but is cortical in the epicotyl. The final structure of the two regions is very similar, save in the arrange- ment of sclerenchyma. Amorpha fruticosa Robinia pseudacacia. Structure of FHypocotyl. The epidermal cells are oblong in cross section, radially elongated at first, later becoming elongated in the tangential direction. A few straight multicellular hairs are present. There is no differentiated hypoderma. The cells of the cortex are all about the same size. The endodermis is small-celled and contains starch. It is not easily distinguished in material taken at the close of the grow- ing season. Starch is also found in some of the pericyclic cells in early stages. Later nearly all the parenchymatous tissues have starch. m The stele is originally quadrangular. There are eight phloem bundles and four xylem bundles. These soon produce closed zones. In the pericycle opposite each of the original xylem bundles a group of stereom appears. These groups, at length, become quite large. In addition to these, at the close of the first year, there are some small patches of stereom irregularly disposed just outside the phloem. 110 MINNESOTA BOTANICAL STUDIES. The pith is composed of parenchymatous cells which acquire thick lignified walls. There is a well-differentiated perimedul- lary zone three or four cells in width. The cells are small and have thick lignified walls. Cork formation, according to Flot [1890], takes place rather deep in the cortex. Structure of Epicotyl. The cells of the epidermis are square or oblong in cross sec- tion, and become in time greatly flattened. There are numerous epidermal hairs. A narrow collenchymatous hypoderma is present. The cells of the cortex are about the same size as the epidermal cells. The endodermis was distinguished only in very young stages. The cells are rather small and closely packed with starch. Starch is found at a later time in the various parenchymatous tissues. Troschel [1879] states that in year-old twigs starch is present in some of the elements of the wood but disappears the next year. In young material the epicotyl is elliptical in cross section. The stele follows this closely in shape. The phloem forms a closed ring surrounding a variable number of xylem bundles. There are usually more than eight of these bundles. The xylem also soon forms a complete zone in which medullary rays are prominent. The pith cells become, at length, thick-walled. There is a well-de- fined perimedullary zone. Cork arises in the fourth, fifth or sixth layer of the cortex (cf. Moel- ler [1882], p. 384). = Comparison of Structure of Hypo- NG cotyl and Epicotyl. ) The hypocotyl is without the col- ' lenchymatous hypoderma of the epi- cotyl; it has fewer epidermal hairs ; the cortex is thicker; there are four Robinia large groups of stereom with some pseudacacia ~~ very small ones instead of a broken Fic. 12. ring of medium-sized bundles. Ramaley: HYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 111 In its primary structure the stele of the hypocotyl differs con- siderably from that of the epicotyl. There are four xylem bundles instead of eight or more and eight phloem bundles in- stead of a closed ring of phloem. The cork, although of cortical origin in both regions, arises in the hypocotyl in deeper layers. RUTACEZ. Ptelea trifoliata. Structure of Hypocotyl. The cells of the epidermis, when seen in cross section, appear nearly square. They, at length, are flattened. There are a few short, blunt, unicellular hairs. The cortex is large-celled. There is no distinct hypoderma differentiated. The endodermis is large-celled and contains, at first, very little starch. It later becomes closely packed with starch. The various parenchymatous tissues at length also contain starch in the cell cavities. Numerous lysigenous reservoirs are pres- ent in the outer part of the primary cortex. The stele is originally four-angled. There is one phloem bundle and one xylem bundle in eachangle. The phloem soon forms a closed ring surrounding the now greatly enlarged xylem bundles which enclose, at this stage, a somewhat cruciform pith. The xylem bundles also finally fuse. Four very small groups of sclerenchyma appear, toward the end of the season, in the pericycle. They are equidistant. There are about six cells in each group. Some sections do not show all these groups, as the sclerenchymatous elements do not form continuous strands in the hypocotyl. Some sections show no sclerenchyma at all. The pith is eventually quite small. The perimedullary zone is not well developed. Si The formation of cork begins early in the outermost cortical layer of cells. Structure of Epicotyl. The epidermis is composed of cells which appear slightly rec- tangular in cross section. They are elongated in the tangen- tial direction. Numerous epidermal hairs are present. There is a narrow collenchymatous hypoderma. The cells of the in- ner cortex are very large. 1 ae MINNESOTA BOTANICAL STUDIES. The endodermis is distinct and can be recognized in year-old material. Starch is present from the first. ‘The various paren- chymatous tissues at length have a small amount of starch. Secretion cavities develop in the cortex. The stele is small; much smaller than is usual in most spe- cies. Inthe youngest material examined the phloem forms a complete ring surrounding a small number of xylem bundles which eventually fuse. Numerous groups of elements inthe pericycle become scler- otic so that they form an interrupted ring of sclerenchyma sur- rounding the phloem. The pith is small, unusually so for an epicotyl. There is a definite perimedullary zone of small-celled parenchyma contain- ing starch. The formation of cork takes place in the outermost layer of hypoderma (cf. Moeller [1882], p. 326). Comparison of Structure of Hypocotyl and Epicotyl. The hypocoty! does not have the epidermal hairs and the collen- chymatous hypoderma of the epicotyl. The cortex of the former region, though very thick, is but little thicker, in pro- portion, than that of the epicotyl. In the stele of the hypocotyl there are four phloem bundles and an equal number of xylem bundles, while in the epicotyl, in the youngest material ex- amined, the phloem forms a closed ring surrounding about six xylem strands. The sclerenchyma of the hypocotyl is in four somewhat irregular columns — in the pericycle, while in the epicotyl it forms more nearly a closed sheath. n the former region also the peri- Fic. 13. medullary zone is poorly developed. Ptelea trifoliata SIMARUBACES. Ailanthus glandulosa. Structure of Hypocotyl. The epidermis consists of small cells, square or nearly so, in cross section, and considerably bulged when young. A few Ramaley : HYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 113 short, unicellular hairs were seen, but none noted in very young plants. There is a hypoderma of one or two layers of some- what larger and thicker-walled cells. The rest of the cortex is parenchymatous and large-celled. The endodermis consists of cells somewhat smaller than those of the adjacent cortical layer. The endodermis was not recog- nized in older material. Starch is present from the first in the endodermis, pericycle and pith, but does not appear in the cor- tex till nearly the close of the first season. The stele is at first quite small. There are four xylem bun- dles arranged in pairs and four phloem bundles similarly dis- posed. The phloem soon forms a complete ring, surrounding the now considerably enlarged xylem bundles, which also even- tually form a closed ring. Opposite each of the four original xylem bundles there ap- pears in the pericycle a group of sclerenchymatous cells. These groups become, at length, somewhat divided so that the old hypocotyl may have a considerable number of smaller groups. There are numerous sclerenchymatous fibers scattered in small and large patches through the phloem and pericycle. The pith is thin-walled; toward the end of the first season it becomes lignified. The perimedullary zone, described by Flot [1893], as an important feature of the stem structure is first definitely noted at this time. ; The oleoresin canals described by Trécul [ 1867 ] as occurring at the outer border of the pith, and by Van Tieghem [1884] as in the inner xylem of the stem, were not distinguished in the hypocotyl. Crystal rosettes of calcium oxalate occur singly in certain cells of the phloem area. Single oleoresin cells are found here and there in the cortex and phloem. Cork formation, as noted by Flot [1889 and 1890] takes place in the layer of cells just below the epidermis. Structure of Epicotyl. The epidermis resembles that of the hypocotyl, but there are numerous, somewhat long, curved or hooked epidermal hairs. Most of these are unicellular. The hypoderma is, as previously described for the stem by De Bary ({1884], p. 404), collenchymatous. The cells are small; toward the inside the hypoderma gradually shades into the ordinary cortex. 114 MINNESOTA BOTANICAL STUDIES. A definite endodermis was not distinguished at any stage, al- though, since starch is present in the region of the pericyle and endodermis from the first, those regions can be located approx- imately. Starch is found later in pith and cortex; also in many of the inner xylem elements. There are, at first, eight to ten conjoint vascular bundles. Eventually the phloem and xylem form closed rings. Scattered sclerenchymatous elements are found in the phloem, pericycle and cortex. The pith is irregular in outline. The first formed xylem ele- ments project into it. The perimedullary zone is not conspic- uous the first year, being composed of a few cells with unligni- fied walls. The cork cambium is formed in the outermost hypodermal layer (cf. Moeller [1882], p. 327). Comparison of Structure of Hypocotyl and Epicotyl. The hypocotyl has a few, the epi- cotyl a considerable number, of epi- dermal hairs. The hypocotyl does not have the collenchymatous hypo- derma found in the epicotyl. The pith is smaller and circular instead of scalloped ; the perimedullary zone is better developed. The endodermis is distinct in the hypocotyl for a considerable time, \,\ while in the epicotyl it was not *) definitely distinguished at all. The hypocotyl has, at first, four xylem | and four phloem bundles; the epi- }) ) cotyl eight to ten conjoint bundles. iy At the close of the year the only dif- ae means ferences are those noted in the me- Fic. 14. dullary and perimedullary regions. ANACARDIACEZ. Schinus molle. Structure of Hypocotyl. The epidermal cells are square or oblong in cross section, becoming, at length, flattened. There are numerous short epi- Ramaley: UYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 115 dermal hairs. No hypoderma is developed but the cells of three or four outer layers of the cortex are smaller than those of deeper layers. The endodermis is small-celled and easily recognized in young stages, although at that time the cells are without starch. Later starch appears in small quantities in these cells and in those of the pith and phloem. : The stele is originally four-angled and remains so for some time. In each angle there is a single xylem bundle and two groups of phloem; these form a crescent-shaped mass _border- ing a group of cells which later develop into a resin duct. After a time secondary vascular bundles are intercalated be- tween the primary bundles. All finally fuse to produce closed zones of phloem and xylem. A few small groups of sclerenchymatous cells develop at the outer border of the phloem. The pith remains somewhat four-sided. The four original xylem bundles project into it at the angles. The pith cells have thin, unlignified walls. A perimedullary zone of small cells was distinguished. Material old enough to show cork formation was not obtained. Structure of Epicotyl. The epidermis resembles that of the hypocotyl. Trichome structures seem to be no more abundant. There is no collen- chymatous hypoderma developed. The cells of the cortex are all about the same size. The endodermis is not easily recognized owing to the fact that in young stages it contains no starch. Later when starch is present the cells have been compressed and displaced by pressure from the subjacent tissues. The stele contains a variable number of vascular bundles. Usually there are about eight. _In connection with each bundle is a small resin passage, at first pointed out by Trécul [1867]. In older material these resin passages become quite large and somewhat flattened. The phloem and xylem then form closed zones. Groups of sclerenchyma, usually consisting of only a few cells, are found at the periphery of the phloem. These are often located near the resin passages. The pith is nearly circular, not quadrangular, and is com- 116 MINNESOTA BOTANICAL STUDIES. posed of large, thin-walled cells. There is a distinct perime- dullary zone. The region of cork formation was not determined. Comparison of Structure of Hypocotyl and Epicotyl. In their primary structure the steles of the hypocotyl and epi- cotyl show important differences. That of the former region is quadrangular; it has four ‘primary vascular bundles and ata later time other secondary bundles are intercalated. These latter do not have resin canals. In the epicotyl there are about eight vascular bundles each with a resin canal. The pith of the hypocotyl is four- sided, that of the epicotyl circular in outline, when seen in cross section. RHAMNACEZA. Berchemia racemosa. Structure of FHypocotyl. The cells of the epidermis are nearly square in cross section, sometimes radially elongated, but becoming at. length considerably flattened. No hypoderma is developed, although the cells of the outermost layer of cortical tissue are considerably smaller than those below. There are about five layers of cells in the cortex. This tissue is extremely loose, having many in- tercellular spaces. The endodermis is quite distinct until nearly the time that cork formation begins. The cells are smaller than those of the cortex but larger than the pericyclic elements. Starch is present in the endodermis from the first, but does not appear in the cortex at alJ, nor in the pith and phloem till about the close of the season. The stele is originally four-angled. There are four xylem and four phloem bundles. These are paired. They soon fuse so that there are two crescent-shaped bundles, and by further growth closed rings of xylem and phloem are produced. While the bundles are in the crescent form four small groups FIG. 15. Ramaley: HYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 117 of sclerenchyma appear in the pericycle, one opposite each of the original xylem groups. The pith is composed of large cells, whose thin walls be- come, at length, somewhat lignified. A perimedullary zone of about two layers may be distinguished but is not always con- tinuous the whole way around the pith. The cork has its origin in the inner cortex or in the endoder- mis. Some of the layers of cork carry a brown pigment. Structure of Epicotyl. The epidermis, composed originally of small cells, square or pentagonal in cross section, eventually becomes strongly cuticu- larized and the separate elements very much flattened. There is no hypoderma developed. The cortex is rather large-celled, but very narrow, being only three or four layers of cells in thickness. During the second year the walls of these cells become conspicuously pitted. Many large crystals, chiefly cubical in form, are found in this region. The endodermis, composed of flat cells containing starch is distinct till near the close of the first season. Except in the en- dodermis starch is absent until about the end of the first year’s growth, when it appears in the pith and medullary rays. Even in very young stages the phloem forms a closed zone _ surrounding a ring of from six to ten, but generally about eight, xylem bundles. These soon become fused. A narrow band of sclerenchyma, for the most part only one cell wide, is found at the outer limit of the phloem; it does not form a closed ring, but is more or less irregular and broken. Small patches of sclerenchyma are found in the phloem of two-year-old seed- lings. The pith is large-celled ; the walls are thin but slightly ligni- fied. No perimedullary zone was distinguished. The region of cork formation was not distinguished. Two- year-old material was examined, but the cork cambium had not begun to form. Comparison of the Structure of Hypocotyl and Epicotyl. The epidermis of the hypocotyl remains thinner-walled and exhibits less cuticularization than that of the epicotyl. This is to be expected, since in the former region cork is produced the first year, while inthe latter not till a later period. 118 MINNESOTA BOTANICAL STUDIES. The hypocotyi has at first four xylem and four phloem bun- dles, the epicotyl a closed ring of phloem and about eight xylem bundles. In the former area there are but four groups of scleren- ce chyma, while in the latter there is an interrupted circle of thick-walled ele- cc ments just outside the phloem. The pith of the hypocotyl is smaller than that of the epicotyl. Rhamnous purshiana. Structure of fypocotyl. The cells of the epidermis are origi- nally square or pentagonal in cross section. They become, at length, much Berchemia = flattened. The cortex is thick and very large-celled. No hypoderma is differentiated. The endodermis consists of small, thin-walled cells contain- ing starch. It remains distinct until cork formation takes place. Starch is found, late in the season, in the perimedullary zone and phloem, but not in the cortex. The stele is originally somewhat four sided and has four vas- cular bundles which soon fuse and produce closed zones. The pith is composed of very large cells. The perimedul- lary zone is rather ill-defined. It is one or two cells in width. The cells are small and contain starch. The cork is of endodermal origin. Its formation begins to- ward the close of the growing season. BIG. a6: Structure of Epicotyl. The epidermal cells, at first square or pentagonal in cross section, become at length, considerably flattened. ‘There are numerous short, curved and pointed hairs. A poorly developed hypoderma is present in year-old material. The cells of the cortex are all about the same size. The endodermis was distinguished only in young material. The cells are small and contain starch. The perimedullary zone, cortex and phloem have at a later time, small amounts of starch. Ramaley: WYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 119 The phloem, in the youngest material examined, forms a complete ring. There are about six xylem bundles. These soon fuse. Considerable masses of stereom develop at the periphery of the phloem forming a broken sheath. The pith is large-celled. There is present a definite peri- medullary zone of small cells containing starch. The cell walls are lignified. Cork is produced in the outermost hypodermal layer as in other species of /?hamnus (cf. Moeller [1882], pp. 292 et seq.). Comparison of Structure of Hypocotyl and Epicotyl. The hypocotyl has a thicker cortex than the epicotyl; it does not havea hypoderma; epidermal hairs are absent; the scleren- chymatous ring found in the epicotyl is here absent. The endodermis of the hypocotyl remains distinct for a greater time than that of the epicotyl. The former region has originally four vascular bundles; the latter has, in the youngest material examined, a zone of phloem and about six xylem bundles. Cork formation in the hypocotyl is endoder- mal while in the epicotyl it is hypoder- mal. VITACEA. Vitis cordifolia. Structure of Hypocotyl. The cells of the epidermis are nearly square in cross section, but become at length considerably flattened. Many of them are somewhat prolonged, forming short, blunt papilla. A thick cuticle is present. In cross section it Srey arellians appears minutely notched. Fic. 17. Three or four of the outer layers Soe of the cortex are small-celled, but not collenchymatous. The cells of deeper layers are larger and all about the same size. The endodermis is small-celled, and is for a long time read- ily distinguished because it contains starch. Starch is gen- erally absent from the other tissues, but, at a later time, ap- pears in the pith. 120 MINNESOTA BOTANICAL STUDIES. There are four primary vascular bundles. Other secondary bundles soon become intercalated and finally complete rings of phloem and xylem are produced. A single group of sclerenchymatous elements is formed at the outer edge of each primary vascular bundle. By the end of the season other smaller groups are also present. The pith finally becomes very small. Sometimes, by the projection into it of two of the vascular bundles, a line of xylem extends nearly across it. There is no perimedullary zone. The cork cambium, as in other species of Vztes (cf. Flot [1889]), is formed about the close of the first season in the pericycle. Structure of LEpicotyl. The epidermal cells are square in cross section, becoming at length flattened. The cuticle is like that of the hypocotyl. A well-differentiated collenchymatous hypoderma is present. The other cells of the cortex are parenchymatous. The endodermis contains starch and is, therefore, easily dis- tinguished. ‘Toward the end of the season starch is also found in the pith. The number of primary vascular bundles is var- iable. Usually there are more than eight. At an early time closed zones of phloem and xylem are produced. A broken ring of sclerenchyma is developed toward the end of the first year in the pericycle. The pith is thin-walled and large- celled. A definite perimedullary zone was not distinguished. The cork, as in the hypocotyl, is of pericyclic origin (cf. Moeller [1882], p- 207) Comparison of Structure of Hypo- cotyl and Epicotyl. The hypocotyl has a thicker cortex than has the epicotyl. It has but four primary vascular bundles instead of eight or more. It has four large masses of sclerenchyma in the peri- cycle and a few smaller ones instead of a more nearly continuous scleren- Fic. 18. chymatous ring. Vitis cordifolia FRamaley: HYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 121 The pith of the hypocotyl becomes, at length, nearly ob- literated. ELHZAGNACEHA. Eleagnus umbellata. Structure of Hypocotyl. The cells of the epidermis are more or less oblong in cross section. There are no epidermal hairs. The outermost layer. of the cortex becomes somewhat thick-walled but not collenchy- matous. ‘The cortex has about six cell rows. ‘The endodermis is small-celled. It remains distinct for a time, but in sections of material gathered at the end of the season it was not distinguished. Very little starch is found in any of the tissues save in the endodermis. The stele is originally four-angled. A single phloem bundle and two xylem bundles are placed in each of the angles. The phloem soon forms a closed ring while the xylem bundles fuse in pairs and increase in size. This leaves a cruciform pith. The continued growth of the xylem produces a complete zone surrounding, at length, a circular pith. Secretion cells in the phloem are numerous. At the outer edge of the xylem, in old material, are a few patches of sclerenchyma forming a very much interrupted ring. The pith is of considerable extent and is surrounded by a perimedullary zone of small cells containing starch. Cork formation takes place far down in the cortex. Structure of Epicotyl. The epidermis consists of cells which are oblong in cross section with the tangential about twice the radial diameter even in very young material. The peculiar stellate trichome struc- tures, well known in this genus, are abundant. The outer cortical cells are nearly circular in outline, when seen in cross section. They are somewhat smaller than the cells of the epidermis. The inner cortex is composed of large cells which eventually are very much flattened owing to pres- sure of the growing parts within. The endodermis is small-celled and contains starch. In old material it was not recognized. The stele is circular from the first. In the youngest material examined the phloem forms a complete ring surrounding about 122 MINNESOTA BOTANICAL STUDIES. six xylem groups. The xylem bundles soon fuse producing at the end of the season quite a thick zone. An interrupted sclerenchymatous ring is developed in the pericycle. The pith, which is composed of large thin-walled elements, is surrounded by a narrow small-celled perimedullary zone con- taining starch. The cork is formed rather late in the season in the outermost cortical layer as in other species of -laeagnus (cf. Moeller (\z082], px DL 7s Comparison of Structure of Hypocotyl and Eprcotyl. The hypocotyl is without the trichome structures so noticeable in the epicotyl; the stele is at first four-angled instead of cylin- drical; there are four phloem bundles and eight xylem bundles instead of a ring of phloem and six xylem bundles. At the end of the season the vas- cular tissue is alike in the two regions. but the pericyclic sclerenchyma of the hypocotyl is less abundant. Cork is. developed in the inner cortex of the hypocotyl and in the outermost layer of cortex in the epicotyl. MYRTACEZ. Eucalyptus globulus. — Structure of Hypocotyl. The epidermal cells, at first oblong,. radially elongated, become at length in Elaeagnus cross section nearly square. The umbellataé Cuticle, which is covered with eleva- tions, appears, when young, in cross. section minutely serrate. There is nohypoderma differentiated, but the outermost layer of the cortex is smaller-celled than the layers below. The cor- tical cells are large. They become flattened toward the end of the season by the growth of the internal tissues. The endodermis is composed of small cells containing starch 3. it at length becomes indistinguishable. Starch is for the most part absent from other tissues. Lysigenous secretion reservoirs: are found in the conjunctive tissue. Fic. 19. Lamaley : HYPOCOTYL AND EPICOTYL IN WCODY PLANTS. 123 The stele is four-sided. The general shape of the hypocotyl sometimes follows that of the stele (cf. Irmisch [1876]). There are originally four narrow curved phloem bundles and the same number of small xylem bundles. The phloem soon forms a closed ring; the xylem bundles increase in size, leaving for a time a cruciform pith; but eventually the xylem also forms a complete ring and the pith is cylindrical. Four small groups of sclerotic cells make their appearance in the pericycle about the time that the phloem ring is first formed. These groups eventually become somewhat broken up and numerous groups of fibers appear in the phloem ar- ranged in three or more interrupted circles. The pith is large-celled. It is smallin amount even from the first. No definite perimedullary zone was distinguished. Cork formation, according to Flot [1890], is cortical or peri- cyclic. Structure of Epicotyl. The cells of the epidermis are at first more nearly square in cross section than those of the hypocotyl. They at length be- come very much elongated in a tangential direction. The cortex is large-celled; the cells of the outer layer are rather small. No hypoderma is differentiated. Numerous lysigenous secretion sacs are present. The endodermis is thin-walled; the cells are small and con- tain starch. The stele is at first somewhat quadrangular and becomes at length elliptical, in cross section. In the youngest material ex- amined the phloem forms a closed ring. The number of xylem bundles is somewhat variable ; these are so disposed that the pith is generally at first somewhat cruciform. The pericycle becomes, at length, largely sclerenchymatous ; numerous interrupted rings of bast fibers begin to appear but are only slightly thickened the first year. The pith, at first cruciform, becomes somewhat quadrangular. There is a perimedullary zone (fide Flot [1893]). An inter- nal cambium produces a ring of phloem just outside the pith: This is mentioned by DeBary [1884]. A few sclerotic cells were noted at the inner limit of the internal phloem. According to Flot [1890] cork formation is sub-epidermal. 124 MINNESOTA BOTANICAL STUDIES. Comparison of Structure of [ypocotyl and Epicotyl. The hypocotyl is more nearly cylindrical than the epicoty]l ; its epidermis less flat, its stereom is better developed the first year. Stem internodes above the epicotyl are square. The primary structure of the hypocotyl is like that of Ze- coma. ‘There are four xylem bundles and four phloem bundles. In the youngest epicotyl examined the phloem forms a complete ring and there is a variable number of xylem bundles. No internal phloem was recognized in the hypocotyl] although it is quite dis- (i tinct in the epicotyl. The cork of the former region is pericyclic or cortical in origin, that of the latter subepi- dermal. BIGNONIACEZ. Tecoma radicans. Structure of Hypocotyl. The epidermal ceils are rectangular in outline when seen in cross section. From being originally radially elon- te gated they are, _at length, nearly ‘globulus Square. The cuticle in cross section appears minutely serrate. There are a few short, simple, epidermal hairs. The cortex is of loose parenchyma, generally about six layers in thickness. There is no differentiated hypoderma. The endodermal cells are smaller than the cells of the cortex. Originally they are irregularly hexagonal in outline, but toward the end of the year they become elliptical and have slightly thickened walls. Starch is entirely absent, except in the en- dodermal region until nearly the close of the first year, when it appears especially in the pith. The stele is originally very small and somewhat four-sided, containing four xylem bundles and four phloem bundles. The latter are next the pericycle. They alternate with the xylem bundles. These are presumably the four ‘‘ principal bundles ” found, according to Hovelacque [1888], in all Bignoniaceous stems. At quite an early stage the phloem and xylem form closed rings. Ramaley: WYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 125 About the time that this is apparent four small groups of sclerenchyma make their appearance in the pericycle just out- side the original xylem bundles. Each of these is composed of only six to ten cells with extremely thick walls. Later numer- ous isolated, lignified sclerenchymatous cells appear in the outer- most layer of the cortex; a few also are found in the phloem area. The pith is large-celled and thin-walled. The formation of a cambium layer in the small-celled perimedullary region begins some time before the close of the year. This will be further noticed in the description of the epicotyl. Cork formation takes place in the second cortical layer, 7. e., in the cell layer immediately below the sclerenchyma which is thus eventually lost. Structure of Epicotyl. The epidermis has a well-marked cuticle which, in cross sec- tion, appears minutely notched. The cells seen in cross sec- tion are about square, but become tangentially elongated toward the end of the season. There are occasional short epidermal hairs. The cells of the outermost layer of the cortex are somewhat smaller than those of deeper layers. These are considerably flattened. Although at first of about even thickness through- out, the cortex soon grows in thickness at four equidistant points giving the epicotyl a quadrangular prismatic shape. The endodermis, which in young stages is distinct, at length becomes indistinguishable owing to displacement and crowding of the cells caused by growth in the lower layers. Starch, though present in the endodermal region, is found only in very small amount in the cortex and pith until the close of the sea- son. Even then the cells are not closely packed with it. Even in the youngest stage examined, 7. e., second stage of our arbitrary division, the phloem and xylem form complete rings. The outermost cortical layer toward the end of the season be- comes largely sclerenchymatous, although here and there are cells with but slightly thickened walls. Certain cells of the pericycle, at first but slightly differentiated, form, at length, groups of very thick-walled cells. The pith is composed of large cells with thin, unlignified 126 MINNESOTA BOTANICAL STUDIES. walls. It is found to be practically in the center of the section, though Pedicino [1876] found that when the plant climbs the pith is eccentric. About the time that the seedling has developed one internode above the epicotyl a medullary cambium has begun to form in the epicotyl. This produces xylem without and phloem next the pith. Considerable masses of phloem may thus be formed. This peculiar cambium in Zecoma was noted by Sanio in 1864 and fully described later by De Bary [1884]. Young branches of the plant were studied by these investigators who did not examine seedlings. Cork formation, as previously described by Moeller [1882] for young branches, takes place in the second layer of the cor- tex. The cork cells are nearly square in cross section. Comparison of the Structure of Hypocotyl and Epicotyl. Although both hypocotyl and epicotyl are originally cylindri- cal, only the former remains so, the latter developing four thickened areas which make it somewhat quadrangular. The four small groups of scleren- chyma in the pericycle of the hypo- cotyl are represented in the other region by aconsiderable number of smaller groups forming an interrupted ring. The endodermis remains distinct in the hypocotyl for a longer time than in the epicotyl. The medullary cam- bium is formed later and is less active. Catalpa speciosa. Structure of EHypocotyl. The epidermis consists of small cells, square in cross section. Short, blunt epidermal hairs are rather numerous. No hypoderma is developed; all the cortical cells are thin-walled. The endodermis remains distinct for a long time. It consists of thin-walled cells which are but slightly smaller than the cells of the cortex. Starch is present, from the first, in the endoder- mis and toward the close of the season appears sparingly dis- tributed in medullary rays and cortex; it is apparently absent from the pith. Y Tecoma radicans BIG. 20. Ramatley - HYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 127 The stele is small; in cross section it is circular. There are at first six, seven or eight conjoint vascular bundles arranged in a circle. Eight is probably the original number, but fusions often take place between adjoining bundles. Complete zones of xylem and phloem are formed at an early stage. About this time four small groups of sclerencnyma appear in the pericycle ; they are equidistant. These eventually become somewhat divided, and other cells of the pericycle become sclerotic, so that a number of small groups of stereom are found in this area. The pith is small in amount. The cells are large, with thin, slightly lignified walls. There is a perimedullary zone of small cells containing starch. Cork formation takes place in the outermost layer of cortical cells. Structure of Epicotyl. The epidermal cells at first are oblong in cross section; the long axis is at right angles to the periphery of the section. Later the shape is more nearly square. There are many straight, blunt epidermal hairs. The first two or three cell layers of the cortex are collenchy- matous. The other layers are rather small-celled parenchyma. The endodermis, though at first distinct on account of the presence of starch in its cells, was not recognized in older ma- terial. Starch is absent from the other tissues in the early stages, but is at length found in the cortex, phloem, medullary rays and perimedullary zone. Toward the end of the first year a narrow, much interrupted ring of sclerenchyma appears at the outer edge of the phloem. The cells are small with very narrow lumen. The number of vascular bundles is somewhat variable. About twenty is the usual number. These soon unite to form zones of xylem and phloem. _ The pith is large, the cells thin-walled parenchyma. There is a perimedullary zone of small cells containing starch. Cork arises in the outermost hypodermal layer, as it does in the stem of Catalpa catalpa (cf. Moeller [1882], p. 184). Comparison of Structure of Hypocotyl and Epicotyl. The epidermis of the hypocotyl has fewer and shorter hairs than that of the epicotyl. The former region has no hypo- 128 MINNESOTA BOTANICAL STUDIES. derma, though in the epicoty] a distinct collenchymatous zone is developed. The endodermis of the hypocotyl remains distinct fora much longer time than that of the epicotyl; the stele has about eight vascular bundles, instead of twenty or more; sclerenchyma is first disposed in four groups instead of a considerable num- ber. The pith of the hypocotyl is much smaller in amount than that of the epicotyl. RUBIACEZ. Cephalanthus occidentalis. Structure of Hypocotyl. The cells of the epidermis are ob- Catala long or somewhat hexagonal in cross speciosa ~=section. About every fifth or sixth Fic. 22. cell is elongated radially and pointed, projecting somewhat beyond the gen- eral line of cells. These might be described as extremely short hairs. This characteristic feature continues for a consid- erable length of time. No distinct hypoderma is formed. The cells of the two or three outer layers of the cortex are rather thick-walled but not collenchymatous. The cortex is loose with numerous large in- tercellular spaces. The endodermis is large-celled. It remains distinct through the first year although the cells become at length very much flattened. They contain starch. Starch is later found in the various parenchymatous tissues. The stele is circular in cross section. There are originally four phloem bundles and an equal number of xylem bundles ; they are grouped in pairs. The xylem and phloem soon form closed zones, the xylem encroaching upon the pith which, at the close of the first season is almost entirely obliterated. The hypocotyl thus assumes a root-like structure—* rhizelle” of Van Tieghem [1891]. The cork is of epidermal origin. Ramaley: HYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 129 Structure of Epicotyl. The epidermal cells are rectangular in cross section. The radial is the long diameter at first but eventually the two diame- ters are nearly equal. Many of the cells are prolonged to form pointed hairs which are about three times as long as the ordinary cells of the epidermis. A narrow collenchymatous hypoderma is developed; this shades off gradually into the ordinary cortex, which is quite extensive. The endodermis is rather large-celled, the cells resembling those of the cortex but containing starch. The endodermis re- mains distinct throughout the first year. Starch, which is at first absent from the other tissues, becomes, at length, dis- tributed through all the parenchymatous elements. The stele, originally elliptical in cross section, follows the general shape of the epicotyl. Eventually the epicotyl be- comes cylindrical as does also the stele. In the youngest ma- terial examined, the phloem forms a complete zone surrounding a ring of about six xylem bundles, which soon fuse to form a closed ring. A few of the cells of the pericycle become sclerotic after a time. These are generally isolated; not aggregated in groups. The pith becomes quite small; it is surrounded by a well- developed small-celled perimedullary zone whose elements con- tain starch. The cork, like that of the hypo- cotyl, arises in the epidermis. Comparison of Structure of Hypo- colyl and Epicotyl. The hypocotyl differs from the epi- cotylin its shorter epidermal hairs, in the absence of a true hypoderma and in the much looser parenchyma of its cortex. The structure of the stele is also very different, the hypocotyl having originally four phloem bundles and Cephalanthus four xylem bundles, while in the epi- occidentalis cotyl the phloem, even in the youngest FIG. 23. 130 MINNESOTA BOTANICAL STUDIES. stage examined, forms a closed ring surrounding six xylem bundles. At the end of the first year the structure of the two regions is essentially the same except that the hypocotyl is without pith or differentiated hypoderma. GENERAL CONCLUSIONS. The following summary and conclusions are based on the facts shown in the foregoing pages. It is not intended to re- peat here all the points which are there given, but merely to bring together under appropriate headings the most important facts of structure of the hypocotyl and epicotyl in the plants studied. General Shape of flypocotyl and Epicotyl.—In cross sec- tion the hypocotyl is usually circular in outline, the epicoty] is, however, not infrequently hexagonal in outline and somewhat flattened. The hypocotyl has usually in early stages much the greater diameter. Comparison of the Epidermis of FHypocotyl and Epicotyl.— The epidermal cells of both regions when seen in cross section appear at first square or radially elongated. After a time, however, they became elongated tangentially, being stretched by the growth of the stelar tissues and not continuing to divide. In quite young stages some plants have in the epicotyledonary region, epidermal cells which, in cross section, appear tangen- tially elongated, viz.: U/mus americana, Liriodendron tulip- tfera, Butneria florida, Cercis canadensis. ‘'Trichome struc- tures are usually fewer and less complex in the hypocotyledon- ary region, ¢. 9., Ulmus americana, Broussonetia papyrifera, Butneria florida, Robinia pseudacacia, Ailanthus glandulosa, Catalpa speciosa, Cephalanthus occidentalis. In the following species the epicotyl has trichome structures but they are absent from the hypocotyl: Celtis occidentalis, Toxylon pomiferum, Gleditsia triacanthos, Ptelea trifoliata, Fhamnus purshiana, Eleagnus umbellata, Tecoma radicans. fly poderma in the Hypocotyl and Epicotyl.—Only one of the species examined has a definite collenchymatous hypoderma in both regions. This is Butnerza florida. ‘The following plants have a hypoderma in the epicotyl, but notin the hypocotyl: Ced¢t¢s Ramaley: WYPOCOTYL AND EPICOTYL IN WOODY PLANTS. Ve occidentalis, Broussonetia papyrifera, Liriodendron tulipifera, Menispermum canadense, Gleditsia triacanthos, Robinia pseu- dacacia, Ptelea trifoliata, Ailanthus glandulosa, Phamnus purshiana, Catalpa speciosa, Cephalanthus occidentalis. Comparison of Cortex of Hypocotyl and Epicotyl_—The cortical cells of the hypocotyl are nearly always much larger than those of the epicotyl. This is so commonly the case that various species need not here be specially mentioned; a good example is Parkinsonia aculeata. ‘The cortex of the hy- pocotyl is thicker than that of the epicotyl. Comparison of Endodermis and Pericycle in Hypocotyl and Epicoty!.—The endodermis in the hypocotyl is, as a rule, more distinct and persists longer than that of the epicotyl. Its cells contain starch. Pericycle is well developed in the hypocotyl, usually consisting of two or more layers of small cells. Typical Structure of the Stele of the Hypocotyl. ‘The stele is usually somewhat quadrangular. As a rule there are four phloem bundles and four xylem bundles. The phloem and xylem may be in contact or they may be separated by a small amount of undifferentiated parenchyma. In the latter case each phloem area is either directly outside of a xylem area (when the phloem may be spoken of as opposite the xylem), or the phloem bundles are removed from the xylem by greater or less angular distances (alternate arrangement). Using the foregoing terminology the arrangement of bundles may be de- scribed as opposite in the following: Lzréodendron tulipifcra, Menispermum canadense, Butneria florida, Cercis canadensis, Amorpha fruticosa, Ptelea trifoliata, Atlanthus glandulosa, Schinus molle, Berchemia racemosa, Rhamnus purshiana, Vitis cordifolia, Cephalanthus occidentalis. It is alternate in Fu- calyptus globulus and Tecoma radicans. Certain modifications of the more usual type just described would best be noted separately. There are four xylem bundles and eight phloem bundles in PRodénza pseudacacia. In Parkin- sonia aculeata, Gleditsta triacanthos and Eleagnus umbellata there are eight xylem bundles and four phloem bundles. The xylem bundles soon fuse together in pairs in the last two named species. In Celtis occidentalis and Toxylon pomiferum, the phloem forms two crescent-shaped areas while the arrangement of the xylem is normal. Unusual Structure of the Stele of the Hypocotyl.—In certain 132 MINNESOTA BOTANICAL STUDIES. species the stele of the hypocotyl does not have the typical structure just mentioned, the number and arrangement of vascu- lar bundles being different. Udmus americana has two xylem crescents and numerous small phloem bundles; Broussonetia papyrifera has a root-like structure; in Catalpa speciosa there are about eight vascular bundles. Primary Structure of the Stele of the E-picotyl.—tin shape the stele of the epicotyl is often originally somewhat hexagonal, though, as in the case of the hypocotyl, becoming at length cylindrical. As is well known there are usually from six to very many vascular bundles. Sometimes the phloem is com- pletely fused into a closed zone even in very young stages. Arrangement of Sclerenchyma tn the [ypocotyl. 'The scler- enchyma of the hypocotyl first appears as four masses in the pericycle in Zoxlyon pomiferum, Liriodendron tulipifera, Cer- cis canadensis, Gleditsia triacanthos, Robinia pseudacacia, Ail- anthus glandulosa, Vitis cordifolia, Eucalyptus globulus, Te- coma radicans and Catalpa speciosa. In the plants just named this original disposition of the sclerenchyma becomes altered either by the intercalation of parenchymatous elements in the areas of sclerenchyma or by the development of sclerenchyma at other points. In the following plants, however, there is practically no change in the sclerenchyma during the first year and the four original masses remain to the end of the season: Parkinsonia aculeata, Amorpha fruticosa, Ptelea trifoliata, Berchemia racemosa. Comparison of the Hypocotyl and Epicotyl with Reference to the Distribution of Sclerenchyma. Commonly the scleren- chyma in the two regions becomes, at the close of the first grow- ing season, equally well developed and similarly arranged. Ex- ceptions to this rule will now be noted. Sclerenchyma is absent from the hypocotyl of AZenzspermum canadense, Butneria flo- rida and Lthamnus purshiana although present in the epicotyl. In Ptelea trifoliata and Berchemia racemosa at the close of the first year there is a greater development of sclerenchyma in the epicotyl than in the hypocotyl. The reverse of this condition obtains in Celtzs occidentalis and Eucalyptus globulus. Only scattered sclerenchyma in small amount was recognized in either region in year-old material of Broussonetia papyrifera; in Cephalanthus occidentalis a few only of the pericyclic cells of the epicotyl] become sclerotic. Ramaley : HYPOCOTYL AND EPICOTYL IN WOODY PLANTS. 133 Cork Formation in Hypocotyl and Epicoty/. The cork cam- bium is developed in the layer of cells next below the epi- dermis in both hypocotyl and epicotyl of Celtzs occidentalis, Liriodendron tulipifera, Butneria florida, Ptelea trifoliata, Ailanthus glandulosa, Catalpa speciosa. In Cephalanthus oc- cidentalis it is of epidermal origin in both regions. In the following species cork formation is sub-epidermal in the epi- cotyl but the cork is produced in deeper cell layers of the hypo- cotyl: Ulmus americana, Toxylon pomiferum, Broussonetia papyrifera, Lehamnus purshiana, Eleagnus umbellata, Huca- lyptus globulus. In Cercts canadensis, Gleditsta triacanthos, Amorpha fruticosa, Robinia pseudacacia, Vitis cordifolia and Tecoma radicans cork formation in the epicotyl is cortical, while in the hypocotyl it takes place in some cases in the same cell layer, in other cases in deeper layers. Details are given in the previous descriptions for the separate species. Pith and Perimedullary Zone of HHypocotyl and Epicotyl. The pith of the hypocotyl is smaller than that of the epicotyl, sometimes it becomes nearly obliterated, e. @., Cephalanthus oc- cidentalis. The perimedullary zone is sometimes not distin- guished in the hypocotyl though present in the epicotyl, e. &., Parkinsonia aculeata, Gleditsia triacanthos, Eucalyptus globu- lus. The opposite condition is found in Cerczs canadensis and Berchemia racemosa. More often where a perimedullary zone is recognized it is equally developed in both hypocotyl and epi- cotyl. Structure of Hypocotyl and Epicotyl at the close of the first years growth.—Owing to secondary changes the two regions, though at first quite dissimilar in structure, may come to be very much alike. The xylem and phloem always form closed rings ; the endodermis often becomes indistinguishable; the cells of the cortex become flattened. The differences of pith, perime- dullary zone and sclerenchyma have already been given. Condensed Summary. Although secondary changes may cause a great resemblance in the structure of hypocotyl and epicotyl, the two regions are, in their primary structure, essentially dissimilar. The epidermis of the hypocotyl is more often without trichome structures, the cortex is thicker and composed of larger cells, 134 MINNESOTA BOTANICAL STUDIES. the endodermis is more distinct and persists for a greater length of time, the pith is smaller, sclerenchyma is often less well-de- veloped and differently arranged and a hypoderma, so common in the epicotyl, is nearly always absent. Cork formation in the hypocotyl begins either in the same cell-layer that it does in the epicotyl or in a deeper layer, never in a more superficial one. Starch is, as a rule, distributed in the same way in both regions. It is usually present in the endodermis in the early stages but does not appear in other tissues until the plant has developed foliage leaves, in considerable number. As to the structure of the stele it may be said that in the hypocotyl there are usually four primary vascular bundles. The exact disposition of the phloem and xylem elements is subject to some variation. Occasionally there are more than four bundles. In the epicotyl the vascular bundles are from six to eight or very many. The hypocotyl does not have a root-like structure. - EXPLANATION OF PLATES. Plate V. Drawings of cross sections to show the primary stelar structure of the hypocotyl of U/mus americana, Celtis occidentalis, Toxylon pomiferum and Broussonetia papyrifera. Plate VI. Drawings of cross sections to show the primary stelar structure of the hypocotyl of JZenzspermum canadense, Butnerta florida, Amorpha fruticosa, Robinia pseudacacia and Ptelea tri- foltata. Plate VII. Drawings of cross sections to show the primary stelar structure of the hypocotyl of Azlanthus glandulosa, Schinus molle, Berchemia racemosa, Eucalyptus globulus, Catalpa speciosa and Cephalanthus occidentalis. Plate VIII. Photographs of cross sections to show the primary structure of the hypocotyl. 1. Ulmus americana, 2. Liriodendron tulipifera, 3. Parkinsonia aculeata, 4. Cercts canadensis, 5. Berchemia racemosa, 6. Vitis cordifolia,'7. Eucalyptus globulus, 8. Tecoma radicans. The drawings were all outlined with the aid of a camera lucida. The magnification used was about five hundred diameters. For pub- lication the drawings have been reduced to one-half their original size. The magnification used in making the photographs was from fifty to eighty diameters. They have been slightly reduced. All drawings and photographs were made by the author from his own preparations. MOE. 11. MINNESOTA BO Ulnus americana Toxylon pomiferum NICAL STUDIES. PART TE Broussonetia papyrifera EV. MINNESOTA B( Mii; It. o ° ge .) ios fas oS 05,9 0 S lo 9.0.° Ragre i 2 CO wea Sy UY ig ie ene, eae qe DQ reteae e500 t) ene oe 21@, <@ Sos ae Menispermum canadense Amorpha AES . iY 28 a florida Butneri 12 HELIO’ PAR ut ANICAL STUDIES. Robinia pseudacacia Ptelea trifoliata BE VI. ’ WOTs. IT. MINNESOTA BO’ ee e. STUDIES. Jaye stat JUL eres Seite X ER somes: gessO) A Cephalanthus occidentalis wv ~ NOE. LL. MINNESOTA BC Parkinsonia aculeata Berchemia racemosa Cercio canadensis Ulmus americana | PLAJE THE HELIOT) Pip ANICAL STUDIES. PAE: @eteec. 9 83 a Eucalyptus globulus Liriodendron tulipifera Tecoma radicans Vitis cordifolia 4 am VIII. INTING CO BOSTON Ramaley - HYPOCOTYL AND EPICOTYL IN WOODY PLANTS. Too BIBLIOGRAPHY OF ARTICLES TO WHICH REFERENCE IS MADE IN THE FOREGOING PAPER. Dangeard, P. A. Le mode d’union de la tige et de la racine chez les angiospermes. Comptes rendus. 2e Sem. 107: 287. 1888. Dangeard, P. A. Rech. sur la mode d’union de la tige et de la racine chez les dicotylédones. Le Botaniste. 1: 75. 1889. De Bary, A. Compar. Anat. of the Vegetative Organs of the Phanerogams and Ferns. English translation. 1884. Flot, Léon. Sur la region tigellaire des arbres. Comptes rendus. rer Sem. 108: 306. 1889. Flot, Léon. Rech. sur la structure comparée de la tige des arbres. Rev. gén. de Bot. 2: 17-32, 66-77, 122-136. 1890. Flot, Léon. Rech. sur la zone périmédullaire de la tige. Ann. Se. Nat. Bok VIE 18237. 1893. Gérard, R. Rech. sur la structure de l’axe au-dessous des feuilles séminales chez les dicotylédones. _ Comptes rendus go: 1295. 1880. Gérard, R. Rech. sur le passage de la racine a la tige. Ann. Sc. Nate bot. Vio rr: 279. 1631. Goldsmith, Sophie. Beitrige zur Entwicklungsgeschichte der Fi- brovasalmassen im Stengel und in der Hauptwurzel der Dicotyledonen. Inaug. Diss. Ziirich, pp. 48. 1876. Herail. Rech. sur |’ anat. comp. de la tige des dicotylédones. Amniancc. Nat. Bot. VIL. 2: 203: ' 1885. Hovelacque, Maurice. Characteres anatomique genereaux de la tige des Bignoniacées. Bull. Soc. d’Etudes scientif. de Paris, rrme année. 1888. Irmisch, Th. Einige Beobachtungen an Eucalyptus globulus Lab. Zeitschr. f. ges. Naturwiss. 48:—1876. (Just. Bot. Jahresbericht 4: 440. 1878.) Klebs. Beitrage zur Morph. und Biol. der Keimung. Pfeffer’s Untersuchungen aus dem Bot. Inst. zu Tiibingen. 1: 536. 1885. Moeller, J. Anatomie der Baumrinden. 1882. Pedicino, N. A. Studii sulla struttura e sulla maniera di accres- cersi di alcuni fusti di piante dicotiledoni. Annuar. della R. Scuola Super. d’Agricoltura di Portici. 1876. (Just. Bot. Jahresbericht, De. | Os Ar. 1Ld0o.) Sanio, C. Notiz tiber Verdickung des Holzkérpers auf der Mark- seite bei Zecoma radicans. Bot. Zeit. 22: 61. 1864. Trécul. Des vaisseaux propres dans les Terébinthinées. Comptes Fendus 65207. S67. Troschel, J. Untersuchungen tiber das Mestom im Holze der dicotylen Laubbiume. Verhandl. des Bot. Ver. der Provinz Bran- denburg, 21: 78. 1879. 136 MINNESOTA BOTANICAL STUDIES. Van Tieghem, Ph. Sur les canaux sécréteurs des Liquidambarées et des Simarubacées. Bull. Soc. Bot. France II. 6: 247. 1884. Van Tieghem, Ph. Sur la limite de la tige et de la racine dans Vhypocotyle des Phanérogames. Morot Journ. de Bot. 5: 425. 1891. Williams, J. Lloyd. The sieve tubes of Calycanthus occidentalis. Ann. of Bot. 8: 367. 1894. Woronin, M. Ueber d. Bau d. Stammes von Calycanthus. Bot. Zeit. 18; 177. ToGo. m= CONTRIBUTION TO THE LIFE-HISTORY OF RUMEX. Bruce FInx«. Introduction.—The preliminary work leading to the present paper was nearly all done in the botanical laboratories of the University of Minnesota during the summers of 1896 and 1897. This work included a study of the macrosporangia and their con- tents in Bursa bursa-pastoris (L.) Britton, Szlene antirrhina L., Polygonum erectum L., Rumex acetosella L., Reumex salice- folius Weinm., and Rumex verticillatus L. At the close of the season of 1897, I had about an equal num- ber of good preparations of the last two species and had learned that /tumex verticillatus is a much more favorable plant for study than /eumex salictfolius because of the larger size of the structures to be investigated. Consequently, during the latter part of August, 1897, after previous study of the former plant had enabled me to select methods which seemed best adapted to my purpose, a large number of flowers and buds, selected to represent various stages of development of the gametophyte, were preserved for future study. During the summer of 1898 over three thousand macrosporangia were sectioned, and the drawings presented with this paper were reproduced from the preparations that gave the best results. /tumex saliczfolius was also further studied in 1898, and the series of slides of this plant is nearly as complete as that for /ewmex verticillatus. As the two plants gave very similar results, except for size of structures, I have not thought it necessary to multiply figures by giving a full series for both plants. The figures then are, in the main, drawn from preparations from the latter plant; those from the former being introduced only when the equivalent phases of development were not found in the other plant, or were poorly exhibited in the preparations. The plants of the genus studied are not well adapted to the 138 MINNESOTA BOTANICAL STUDIES. study of nuclear mechanism and phenomena because of the small size of the cells and contained nuclei, and I have conse- quently confined myself to other phases of the subject. Nor have I found anything in the study that would give additional evidence as to the nature of various structures within the em- bryo sac so that my work resolves itself into a description of the gametophyte and some comparative studies. Under the subject stated I shall, for the sake of relationships, begin with the archesporium, which is the last term in the sporophytic generation and also include stages immediately fol- lowing the establishment of the sporophyte. So far as I know this is the first work done on the female gametophyte of /twmex, or of any plant within the Polygona- cee, except Polygonum divaricatum, which has been investi- gated by Strasburger. I am under obligations to Professor Conway MacMillan for helpful suggestions as to technique and interpretation of struc- ture and for access to the literature of the subject. Origin of the Macrospore.—At about the usual stage in the development of the macrosporangium, an axial hypodermal cell at the summit of the nucellus begins to enlarge and soon contains a larger nucleus and denser cytoplasm than the sur- rounding cells (Fig. 1). This cell constitutes the archespor- ium, and in all instances examined, only one cell showed this archesporial nature. The archesporium in plants may develop directly into the macrospore; it may itself become a sporogen- ous cell (mother cell) and divide into a number of potential macrospores ; or more commonly it divides first into a tapetum and a sporogenous cell, each of which may divide, forming a cellular tissue. In /tumex the last order of development is followed. After increasing considerably in size (comp. Figs. 1 and 2), the archesporium divides by a periclinal wall into the inner sporogenous cell (the mother cell of the macrospore), and the outer hypodermal tapetum ( Fig. 2). This apparently pro- tective tapetum proceeds next to divide, sometimes by a peri- clinal wall ( Fig. 4), but no doubt much more commonly by an anticlinal (Fig. 3). In no instance did I observe more than four tapetal cells derived from the primary tapetum and often only three, one of the two derived from the primary tapetum apparently failing to divide. Indeed, sometimes only one tape- tal cell could be distinguished at a period of development which Fink: CONTRIBUTION TO THE LIFE-HISTORY OF RUMEX. 139 led to the suspicion that the tapetum may sometimes fail to di- vide. Fig. 5 shows four tapetal cells and the sporogenous cell below somewhat more elongated than usual with its nucleus un- usually near its upper end. That this elongated cell is the mother cell of the macrospore instead of the macrospore itself is proved by the outline of the inner seed coat, which shows its early development as compared with its more advanced condi- tion before the macrospore is produced (Fig. 10). Fig. 6 rep- resents a single tapetum lying above a dividing mother cell, and this may be the original tapetum which has failed to divide, though a cell lying almost directly below it, and hence not shown in the figure, may have been a second tapetal. Fig. 3 shows a typical mother cell apparently about ready to divide, and showing two tapetals, neither of which would be likely to divide again. After the formation of the protective cap of tapetal cells and the enlargement of the mother cell of the macrospore, the latter divides as usual, among the Archechlamydez at least, into a row of four potential macrospores. It-is here that our subject proper really opens since this mother cell, in which the reduction of chromosomes takes place, stands between the sporophytic and gametophytic generations, connected morphologically with the former and physiologically with the latter. As the mother cell divides the nucleus lies longitudinally at or a little above the center of the cell (Fig. 6). As elsewhere in these studies, the number of chromosomes could not be made out, but, in all probability through failure to get a complete view of individual chromosomes, there really seemed to be twenty-four in this nu- cleus. The two cells resulting from this division were not seen, but from the position of the dividing nucleus in several instances observed, it may be assumed that the division is into two cells of approximately equal size as observed by Strasburger* in Po- lygonum divaricatum. This dividing nucleus of the mother cell of the macrospore was about as large as that of the macro- spore itself (Fig. 9) and, being like the latter a nucleus of a large well-fed and consequently somewhat inactive cell, was ap- parently a long time in dividing. The reduction of chromosomes is supposed to be a process involving more time than is com- monly occupied in mitotic division of nuclei; and this process doubtless also added further to the time occupied by this mother * Strasburger, E. Die Angiospermen und die Gymnospermen, p. 5, 1879. 140 MINNESOTA BOTANICAL STUDIES. cell nucleus in dividing, so that it was seen dividing much more frequently than the equally large and apparently as well-fed nucleus of the macrospore. As stated above the two cells resulting from the division of the macrospore mother cell were not seen in the resting condi- tion, though the four potential macrospores resulting from their division were frequently found. The two cells were seen once with their nuclei dividing, but the nuclei were indistinct because of improper staining. The two-celled condition, which was not seen, is doubtless a very transient phase, the upper of the two cells almost immediately cutting off a small cell from its lower end, and the lower of the two likewise dividing at once into two cells, a large one below and a small one above ( Fig. 7 ). This four-celled stage seems to be constant and of quite long duration as it was frequently seen as represented in Fig. 7 or as in Fig. ro. In the latter the lower more successful cell which is to become the fertile macrospore has begun to absorb the other three potential macrospores for its own nourishment. This may be seen by observing the decrease in amount of cyto- plasm contained in each of the three cells as compared with the same three in Fig. 7 and the swelling of the softened periclinal cell walls between any two of them and between the lowest one and the absorbing cell below. The next observed develop- mental condition was that in which the three cells were all ab- sorbed except possibly a refractive cytoplasmic cap at the sum- mit of the absorbing cell, which more probably represents a nearly absorbed tapetal cell (Fig. 8). This brings us to the macrospore as shown in the figure. Germination of the Macrospore.—When first formed the macrospore has very nearly the shape of the four cells replaced and shows the nucleus at the center surrounded by cytoplasm, while the upper and lower ends are each occupied by a large vacuole. Though I cannot account for this apparent poverty in cytoplasm at this time, the condition seems to be typical. As the macrospore increases in size by the absorption of tapetal cells and those cells of the macrosporangium which surround its upper lateral wall, cytoplasm increases in amount (Fig.9). In the figure the nucleus is lying in its usual longitudinal direction about the middle of the macrospore while dividing. Before the nucleus divides, the macrospore increases considerably in size by the absorption and pressing upon surrounding tissues (comp. Fink: CONTRIBUTION TO THE LIFE-HISTORY OF RUMEX. 141 Figs. 8 andg). The cavity of the macrospore, which I shall now designate by the usual name of embryo sac, continues to increase in size as it approaches the condition shown in Fig. tr by the continual absorption and pressure upon surrounding cells. The relative position of cytoplasm and vacuoles in Fig. g is hardly normal, the nucleus of the macrospore more com- monly lying along the central longitudinal axis of the spore. In another preparation showing the dividing nucleus of the mac- rospore, the nucleus was in this more usual position, and no vacuole was seen. Fig. 11 shows the two nuclei derived from the nucleus of the macrospore and two tapetals and two other sporangial cells nearly absorbed. Of these two nuclei it is quite common to find the lower one larger, probably as a result of better nourishment, and, apparently consequently giving rise in division to a larger number of chromosomes. The condition as to chromosomes could not be studied, but I noticed relative sizes carefully. As instances of difference in size of the two nuclei, the researches of Sargant,* Mottiert and -Guignard{ may be cited. I examined a number of the embryo sacs showing the two nuclei, and it would seem that the lower nucleus becomes very slightly larger than the upper (Figs. 11 and 12). ‘The slight difference may not be constant, and indeed in the closely related Polygonum divaricatum Strasburger§ shows the upper nucleus larger thanthe lower. This stage of develop- ment was quite frequently observed, but the next, in which two nuclei appear in each end of the sac, was only seen twice (Figs. 13 and14). The first of the two figures shows the probable posi- tion of the two nuclei during division as a persistent spindle was seen between the anterior pair. Then the anterior pair seems to result from the division of a nucleus lying transversely in the anterior end of the sac and the posterior pair from a posterior nucleus lying longitudinally in the sac. This position of the two nuclei while dividing is the common one in plants so far as I can ascertain. Fig. 14 represents a later stage, in which the *Sargant, Ethel. The Formation of the Sexual Nuclei in Leléum martagon. I. Odgenesis. Annals of Bot. 10: 464. S. 1896. +Mottier, David M. Ueber das Verhalten der Kerne bei der Entwicklung des Embryosack und die Vorgiinge bei der Befruchtung. Jahrbiicher fiir wissen- schaftliche Botanik 31: 132. 1897. tGuignard, L. Nouvelles Etudes sur le Fecundation. Ann. des Sci. naturelles Botanique VII. 14: 187. 1891. §Strasburger, E. 1. c. Fig. 15. 142 MINNESOTA BOTANICAL STUDIES. nuclei have changed position somewhat. A difference in size has frequently been observed here also, in some other plants the lower pair being larger. I may cite Guignard* again and also Mottier.t However, the last named investigator only states that the nuclei forming the lower pair are much larger just before dividing, distinctly stating} that at an earlier period the four are all of equal size. I did not observe this phase of development often enough to be certain that the size is the same at all times in /twmex ; but it seems from the instances observed that, though coming from the nuclei differing somewhat in size, the four are so nearly of the same size at all times that any difference would be difficult to detect. During the change to the four-nucleate con- dition the sac increases somewhat in size, as may be seen by comparing Figs. 13 and 14 with Fig. 12. The development from the four-nucleate to the eight-nucleate condition must be even more rapid than that from the two-nucle- ate to the four-nucleate phase, for I was neither able to find the two tetrads in position, nor the division of the four nuclei leading to its establishment. The nearest approach to it was observed. in Rumex salicifolius (Fig. 15), when the polar nuclei were ap- proaching. The condition represented in Fig. 15 is an espe- cially interesting step in the life-history of the gametophyte of Fumex because of departure from the usual conditions and espe- cially from the nearly related Polygonum. On examining the figures of Guignard, Vesque, Strasburger, Ward and others, I find that in fully five-sixths of their drawings they show cell walls about the three anterior cells before the polars have fused. Strasburger figures for Polygonum divaricatum § three nuclei enclosed in cellular membranes and one free nucleus in each tetrad even before the two polars begin to approach each other. My own observations on Polygonum erectum L. (Fig. 16), Bursa bursa-pastoris ( L.) Britton and Sz/ene antirrhina L. gave the same results so far as the anterior end of the sac is con- cerned, though the evanescent antipodals of Bursa were not satisfactorily studied in this respect. Methods which brought out these walls in the three plants named above should show them, if present in /twmex, yet in this genus I find all the an- *Guignard, L., l. c. 188. t+tMottier, David M., l. c. 137. }Mottier, David M., 1. c. 136. § Strasburger, E. 1. c. Fig. 17. Fink : CONTRIBUTION TO THE LIFE-HISTORY OF RUMEX. 143 terior three cells free while the polars are approaching each other in Aeumex salicefolius (Fig. 15). When the polars have met, or are fusing in this plant, the cellulose walls seem to begin to form, and a suggestion of such a structure may occasionally be seen (Figs. 17 and 20) about the nuclei of the synergide. In Rtumex verticillatus, even shortly after the definitive nucleus is formed (Fig. 18), I could only distinguish a suggestion of a beginning of formation of a membrane about the lower end of the egg while the sister nuclei which form the synergide were yet free. In Mumewx salicifolius the synergide form earlier than in /tumex verticillatus and are more regular in form in the former plant (compare Fig. 24 with Figs. 19 and 23). So far as investigation has proceeded then, cell walls appear about the three anterior nuclei (excluding the anterior polar) before the meeting of the polar gametes in Polygonum; while in /umex the walls appear after the meeting of the gametes, or even after their fusion to form the zygote constituting the definitive nucleus. The matter is one of some taxonomic interest, which can only be solved by a laborious study of all the species of the two genera with reference to this particular problem in developmental history. Regarding conditions presented in Figs. 15, 17 and 20 some further statements are necessary. In these stages I was able to note no difference in size of the three anterior nuclei, while at later periods the nucleus of the egg had increased in size so that it was larger than those of the synergide (Figs. 18 and 19). So far as I could ascertain, the nuclei of the three antipodals were smaller than the three just discussed at all periods. Fig. 17 represents the three antipodals as best seen in Rumen salici- folius, and here their smaller size can be seen distinctly. The antipodals are difficult of observation at all times because of their lying in the small cazecum-like prolongation of the embryo sac so that, except in very thin sections, other cellular struc- tures of the surrounding macrosporangium almost always par- tially or completely obscure them. In the sac of the same plant showing a slightly earlier stage of development (Fig. 15), only the deeply-stained nucleoli could be seen through the overlying tissues of the macrosporangium, neither the nuclear membrane nor the cellular wall about each of the three nuclei appearing. In this plant the three-celled antipodal area was found persisting in the latest phases studied after the establishment of the em- bryonic sporophyte (Fig. 33) and seems to be quite constant, 144 MINNESOTA BOTANICAL STUDIES. though in the sac represented in Fig. 24 I suspect that there were really no walls about the antipodal nuclei. However, here again overlying tissues may have obscured them. /tumex verticillatus gave less satisfaction in the study of the antipodal region. In only one instance were three antipodals seen con- stituting a three-celled mass of tissue (Fig. 21). This figure represents the posterior end of a mature sac. In other instances various conditions of cell-wall formation were shown in this re- gion before the establishment of the sporophyte (Figs. 23, 27 and 28). Sometimes more than three nuclei could be distin- guished within this antipodal area (Figs. 23 and 28). After fertilization these cells seem to disintegrate more or less in this plant, and no such typical structure was found persisting as is figured for /tumex salicifolius (Fig. 33). The degenerating condition of the antipodal region just after fecundation is shown in Fig. 27, which is the lower end of the sac presented in Fig. 26. Here I was only able to make out a highly refractive area with neither walls nor nuclei. As to the time when the cellular membranes appear around the antipodal nuclei, I am not able to state certainly because of the difficulty encountered in inves- tigation, but it seems, from the facts presented and other ob- served phenomena, that in /eumex salicrfolius they form earlier than those about the three nuclei in the anterior end of the sac. In Reumex verticillatus walls were not detected till the time of maturity of the embryo sac when the typical three celled condi- tion (Fig. 21) presented itself, or that of three or more nuclei. within a common wall (Fig. 23). The antipodal area in Aumex salicifolius seems to agree substantially with the third of the four types proposed by Coulter* of ‘* three comparatively per- manent cells not notable for size or activity and usually associ- ated with a sac decidedly narrowed at the antipodal end.” The antipodals of /tuwmex verticillatus are not so permanent, but doubtless should be classed here also as should those of Poly- gonum erectum. The size of the two polars in plants may differ as well as their place of fusion. Schaffner finds the upper one larger in Alisma plantagot and Sagittaria variabilis.t Also he finds in *Coulter, John M. Contribution to the Life-History of Ranunculus. Bot. Gaz. 25: 80-81. F. 1898. + Schaffner, John H. The Embryo Sac of Alcsma plantago. Bot. Gaz. 21: 126. Mr. 1896. + Schaffner, John H. Contribution to the Life History of Sagittarta varia- bilis. Bot. Gaz. 23: 255. Ap. 1897. Fink: CONTRIBUTION TO THE LIFE-HISTORY OF RUMEX. 145 the last plants that the two polars fuse in the lower part of the sac, the larger gamete strangely enough traveling further than the smaller and thus showing greater activity. Mottier * finds them of about the same size in Seneczo aureus and that they fuse in the anterior end of the sac just below the egg, the lower of the two equal-sized polars thus showing greater activity. In ftumex, and all of the Polygonacez so far as studied, the two polar nuclei are of about the same size (Figs. 15, 17 and 20) and fused at or near the center of the sac. During the passage from the four-nucleate to the eight-nucle- ate condition, the sac increases considerably in size (comp. Figs. 13 and 15), and the increase in size is even more noticeable as the sac matures (comp. Figs. 15 and 24). The Mature Embryo Sac.—The mature sac contains typically the usual seven nuclei, though in Aeumex verticillatus, as stated elsewhere, the number in the antipodal region may vary, pro- ducing a corresponding variation in the total number contained in the sac. In Aumex verticillatus the synergide are some- what irregular in form (Figs. 19 and 23), while in Aeumex salicifolius they are more regular in outline (Fig. 24). The egg usually lies in contact with the lower part of one synergid in both plants (Figs. 19, 23 and 24), and its nucleus is much larger than those of the synergide. All three nuclei are en- closed in definite walls at this time. The definitive nucleus is much the largest nucleus in the sac (Figs. 18, 23, 24, 25 and 28). In ARumex vertecillatus it usually approaches the egg after formation and lies close to it till the time for fecundation approaches (Fig. 23) when it commonly recedes somewhat (Figs. 25 and 26). In Aumewx salicifolius it usually lies at some distance from the egg in the mature sac (Fig. 24) though it was once seen nearer in an earlier stage (Fig. 22). In /tu- mex salictfolius it may be assumed that the antipodals are in the typical three-celled condition at maturity as they were observed in this condition both before and after as already stated (Figs. 17 and 33). As before noted the condition of the antipodals in Fig. 24 is doubtful. The condition of the antipodals in Aumex verticillatus at this time has also been explained above. In shape the mature sac in both plants differs quite widely from that of Polygonum divaricatum as shown in Strasburger’s * Mottier, David M. On the Embryo Sac of Senecio aureus. Bot. Gaz. 18: Fig. 1 and p. 248. Jl. 1893. 146 MINNESOTA BOTANICAL STUDIES. figure,* the largest part being near the anterior end and the posterior narrowed end being quite long (Figs. 17, 23, 24, etc.) while his figures show the narrowed posterior caecum dilating quite abruptly, making the posterior half of the mature sac quite as large as the anterior half. Polygonum erectum the mature sac is much more like Aumex than Polygonum divaricatum as figured by Strasburger. Fecundation, and the origin of the Sporophyte. During the period of fecundation and the establishment of the sporophyte, the embryo sac continues to increase in size rapidly, as may be seen by comparing Figs. 24 and 33 and also 23 and 30, making allowance for the greater reduction of Figs. 30 and 33. The first evidence of a poilen tube approaching or already present in the sac is the disappearance of one of the syner- gide (Fig. 25). When actually present the tube is an easy object to detect because it stains more deeply than surrounding tissues, hence in those instances in which one synergid was breaking down while no tube could be distinguished, I have concluded that the tube was just approaching the sac. In Fig. 25 the tube has not yet discharged, as it shows two nuclei and the end is intact, and it lies as usual beside the persistent syn- ergid. The definitive nucleus here occupies a position at some distance from the egg as it frequently does at this time. Here this nucleus is in a resting condition, and I find it so constantly up to this time. After the pollen tube has discharged the re- maining synergid disappears, and the definitive nucleus soon divides (Fig. 26).’ Though in some plants the definitive nu- cleus seems to divide sometimes before the entrance of the pol- len tube into the sac, its presence in the style probably furnish- ing the necessary stimulus, it does not divide till after the entrance of the tube in Rumex verticillatus and probably usually not till after the fecundation of the egg. In the discharge of the sperm nucleus the tip of the tube is ruptured as shown in Figs. 26, 29, 30 and 35. After the pollen tube has discharged a deeply stained spot may always be seen within the tube as shown in the same figures. This is the shrunken second and undischarged sperm nucleus. The fusion of the sexual nuclei was only seen once, and in that instance the egg was badly distorted. The tube in this case was closely applied to the egg though the figures given herewith seem to indicate that this *Strasburger, E. 1. c. Fig. 21. Fink: CONTRIBUTION TO THE LIFE-HISTORY OF RUMEX. 147 is not the usual relation during fusion of these nuclei. The conditions existing in Fig. 28 are somewhat of an enigma. The sac is surely larger than it could be previous to fecundation, but the two smaller nuclei in the anterior end look like those of the synergide. Also the definitive nucleus lies near the egg as I uniformly find it in /twmex verticillatus before fecundation. I thought for a time that I had here a pollen tube showing one sperm nucleus within and the other discharged and entering the egg. However, taking into account the appearance of the two smaller nuclei and the restful condition of the egg, I have concluded that the sac is one whose egg failed to be fertilized and in which one synergid has disintegrated leaving its nucleus while the other is rapidly dwindling. If this is true, the sac has gone on increasing in size the same as if the egg had been fecundated. The next stage observed after that already ex- plained (Fig. 26) is that represented in Fig. 30, in which the tube had discharged, the first two endosperm nuclei had divided and the spindles were persisting, and the egg was evidently preparing to divide. A little later phase was also found (Fig. 29), in which the tube was persisting and showed the broken end beautifully, the egg was dividing and three of a probable four endosperm nuclei were visible. The tube frequently persists in both plants till the sporophyte is well established (Fig. 35), and in one instance an undischarged tube was seen in the sac of Rumex verticillatus after the embryo was well established and four endosperm nuclei were dividing (Fig. 34). This tube, containing two sperm nuclei, is a second one which entered the sac after fecundation had been accomplished. An exhaustive study of the embryo will not be attempted ; but I shall state a few observed facts concerning its origin and early development, making no use of the terms suspensor and pro- embryo, but designating the structure from the beginning as the embryo. The first division then of the embryo occurs soon after the egg has elongated and secreted a cellulose wall about its base to attach it to the anterior end of the sac and is trans- verse (Fig. 29). The second and third divisions were also ‘transverse in several instances observed (Fig. 35), and the fourth was a longitudinal division of the distal cell of the em- bryo (Fig. 31). In an instance observed the third division was transverse and closely followed by a longitudinal dividing the second cell from the distal end (Fig. 32). During its early de- 148 MINNESOTA BOTANICAL STUDIES. velopment the embryo is not always attached centrally at the anterior end of the sac, nor does it often lie in an exactly longi- tudinal direction in the sac. It has already been stated that the definite nucleus in Aumex does not divide till after fecundation of the egg, an observation based upon examination of two or three hundred sacs. It is worthy of note that the endosperm nuclei observed while divid- ing in any given sac were all in the same mitotic phase (Figs. 30 and 34). The last figure shows only one of four nuclei ob- served dividing. ‘The endosperm nuclei were not so numerous, at the stages studied, in Aewmex verticillatus as in Ftumex salici- folius (Fig. 33); nor were they yet enclosed in cell walls in either plant. feelation of the Gametophyte to the Macrosporangium.—lI have studied carefully the position of the base of the sporogen- ous tissues and derived embryo sac with reference to lines con- necting the points of origin of the seed coats in order to as- certain how much of the enlargement of these structures is associated with a downward growth and consequent crowding of the tissues of the macrosporangium and how much is accom- plished by upward growth, keeping pace with the growth of the nucellus. The position of the base of sporogenous tissue, and later of the sac, with reference to these lines is not always quite the same at any particular stage of development; but by comparative study, safe conclusions have been secured. Be- tween the base of the archesporium and the lines connecting the supposed points of origin of the future seed coat are five or six cells. After the tapetum is cut off (Fig. 2), there are only three or four cells between the base of the mother cell of the macrospore and the lines. By the time of division of the tape- tum (Fig. 3) the base of the mother cell is within two or three cells of these lines. During this time the nucellus has increased in length very little, its increase in size being principally in width. Consequently, this fact, together with the relative posi- tion of the base of sporogenous tissue and the lines at various stages of development, indicates that the sporogenous tissue has grown downward in the nucellus. As no evidence of absorp- tion of cells was seen at this time, I conclude that this down- ward growth is accomplished by crowding downward and out- ward the subjacent layers of cells of the macrosporangium. However, the effects of the crowding were so distributed among Fink: CONTRIBUTION TO THE LIFE-HISTORY OF RUMEX. 149 several layers of cells that they were scarcely visible in any particular cell. After the mother cell has reached its full length, there is no further downward growth, the further increase in length of the sporogenous tissues and subsequently of the sac being accompanied by a proportionate elongation of the nucel- lus. By the time of the establishment of the macrospore, a thickening of the walls of cells in the chalazal region for the support of structures above has begun in a layer of cells ex- tending transversely between the points of origin of the inner seed coat. As the superimposed structures become heavier, the thickening extends to several layers of cells below the ones first thickened and gives rise to quite a mass of thick-walled tissue extending entirely across the chalazal region. As the macrospore matures and prepares to divide (Fig. 9), absorption of tapetals above and pressure on surrounding cells of the upper nucellus becomes evident and is apparent in all subsequent stages of development of the gametophyte. How- ever, the swelling of the cell walls of the upper nucellar tissue just beneath the epidermis as observed by Strasburger* in Poly- gonum divaricatum as a result of absorption I have not seen either in Atwmex or Polygonum. By the time development has proceeded to the condition rep- resented in Fig. 11, the subepidermal cells of the upper end of the nucellus have all been absorbed, and from this time on till the establishment of the conditions shown in Fig. 28, or possibly not later than those shown in Figs. 25 or 26, the increase in size is due, at least principally, to the absorption of cells of the nucellus surrounding the middle portion of the sac, which still continues to increase in size. During this time the sac is in- creasing in length, and since there has been no further sinking of its posterior end into subjacent tissues, as is shown by the fact that its lower end is still removed from the lines connecting the points of origin of the inner seed coat by two or three cells, as was the lower end of the mother cell, this increase in length is accompanied by an equal upward growth of the nucellus. Dividing nuclei were seen in the basal region of the nucellus, both in the epidermis and in the sub-epidermal cells, up to the latest stages studied, indicating that this basal portion of the nucellus is its chief region of growth at these stages. After the growing gametophyte has absorbed all the sub-epidermal tissues of the *Strasburger, E. |. c. Figs. 10 et seq. 150 MINNESOTA BOTANICAL STUDIES. upper nucellus, the sac does not cease to expand laterally, but presses the remaining epidermis of this portion of the nucellus outward as it still further increases in size. These epidermal cells contain cytoplasm and may divide even after the cells of the inner nucellus, or their cytoplasmic contents, at least, have been absorbed; but the increase in epidermal surface accom- panying the continued increase in size of the sac is doubtless due principally to increase in length of these upper epidermal cells and the division of those near the base of the nucellus. Fig. 28 shows certain of these conditions of the nucellus brought about by absorption of its tissues by the growing game- tophyte and by its own growth. All of the epidermal cells ex- cept those at the summit show elongation in the direction of upward growth of the nucellus. All of the epidermal cells ex- cept those at the summit are also well filled with cytoplasm, indicating activity. The lowest sub-epidermal cells of the nucellus shown in the figure are also well filled as were the cells of five or six layers lying between the lowest shown in the fig- ure and the area of thickened cells in the chalazal region. In passing upward from the base to the summit of the nucellus, we find greater and greater absorption of the cytoplasmic cell-con- tents and finally a partial breaking down of the cell walls and a beginning of the consequent collapsing of the cells. In the lower portion of the nucellus the absorption by the gametophyte has only affected the layers of cells near the sac, while in the upper portions all of the sub-epidermal cells are affected. The upper and older cells of the nucellus, where not too much disin- tegrated, also show an increase in size over those of younger nucelli, which accounts in part for the elongation of this organ as it keeps pace with the growth of the sac. Methods.—After trying one per cent. solution of osmic acid, one-half per cent. and one per cent. chromic acid, a saturated solution of corrosive sublimate in seventy per cent. alcohol and the last two plus a small addition of acetic acid to prevent shrinkage, the corrosive sublimate with acetic acid was found to give best results. The sections thus fixed also took best the stains used. After this method of fixing, the tissues were washed in seventy per cent. alcohol containing iodine, gradually transferred to ab- solute alcohol, imbedded in pa affine through xylol, stained and mounted in balsam in the usial way. Alcohol safranin fol- Fink > CONTRIBUTION TO THE LIFE-HISTORY OF RUMEX. I)51 lowed by methyl blue was found to be the best stain for stages before the establishment of the macrospore, and safranin or Del- afield’s hematoxylin gave best results from this period up to the fecundation of the egg, after which the hematoxylin proved best. The nucellus of Polygonum erecium is transparent enough to show the nuclei of various stages of development of the sac, ex- cept the antipodals, as well as the cytoplasm and vacuoles with oil immersion lens, without sectioning or any treatment what- ever. The results, however, were not reliable enough for my purpose, nor are they certain enough for use in instruction. The figures are all drawn to the same scale by using one-inch Leitz eye piece, one and one-half-inch Leitz objective and camera lucida. EXPLANATION OF PLATES. Notre.—All figures are of Rumex verticillatus unless otherwise indicated. The figures of the first two plates are reduced to one-half the original size of drawings, those of the last two to one-third. Plate IX. 1. Upper portion of nucellus showing the archesporium. 2. Later stage showing the primary tapetum cut off above and the mother cell of the embryo sac below. 3. The tapetum has divided into two tapetal cells and the mother cell has increased considerably in size. Stage between 2 and 3 showing the tapetum dividing. Four tapetal cells and the elongated mother cell. The mother cell dividing and one tapetal cell above. The four potential macrospores derived from the mother cell. The macrospore and a highly refractive cytoplasmic cap, repre- senting an almost completely absorbed cell, either a tapetal cell or the upper one of the potential macrospores. 9g. The nucleus of the macrospore dividing. 10. A somewhat older stage than 7 and showing the upper three potential macrospores partly absorbed by the lowest one, which is to become the macrospore. 11. Embryo sac containing two nuclei resulting from the division of the nucleus of the macrospore, the lower one being somewhat larger. The highly refractive remains of fer nearly absorbed cells of the nu- cellus are also shown. Te DANS 1h? MINNESOTA BOTANICAL STUDIES. Plate X. 12. Embryo sac of Rumex salicifolius showing the two nuclei as above and the lower one also slightly larger. 13. Embryo sac of Rumex salicifolius showing the four nuclei de- rived from division of two corresponding to those figured in 11 and 12, and also showing the persistent spindle in the anterior end of the sac. 14. Embryo sac of Rumex salictfolius showing the corresponding four nuclei in a somewhat older sac. 15. Embryo sac of Aumex salictéfolius showing eight nuclei derived from four corresponding to those figured in 13 and 14. The polar nuclei are approaching each other. The three anterior nuclei are of about equal size and not enclosed in walls. Only the nucleoli of the antipodals could be seen. 16. Embryo sac of Polygonum erectum showing the earlier forma- tion of cell walls about the anterior nuclei, one of which was lying be- low the other two. The polar nuclei are approaching each other, and the cellular structure in the antipodal area is distinct, one cell here also lying below the other two. 17. Embryo sac of Rumex salicifolius showing the egg yet free, cell walls forming about the synergidz, the polar nuclei fusing and- three distinct nucleated antipodal cells. 18. Embryo sac showing the sister nuclei which form the synergidz yet free and the wall forming about the egg after the definitive nucleus is formed. ; 19. Anterior end of a mature embryo sac showing the synergidz and the egg. 20. Embryo sac of Raumex salictfolius showing all of the nuclei of the mature sac, except the antipodals and at about the same stage as the corresponding nuclei in 17. 21. Posterior end of a mature embryo sac showing three antipodal cells. 22. Anterior end of the embryo sac of Rumex-salictfolius showing walls forming about the egg after the definitive nucleus is formed. 23. Mature embryo sac showing more than three nuclei in the anti- podal end. 24. Mature embryo sac of Rumex salictfolius, with the antipodal region perhaps not distinctly seen. Plate XT. 25. Anterior end of embryo sac showing one synergid degenerated, the pollen tube entering and showing two nuclei very indistinctly. The egg and definitive nucleus are also shown. 26. Anterior end of embryo sac showing a pollen tube discharged and containing an undischarged sperm-nucleus. The two synergidz VOL. TT: MINNESOTA BOTANICAL STUDIES. PART Ik PEATE! IX HELIOTYPE PRINTING CO. BOSTON. MINNESOTA BOTANICAL STUDIES. Ppa ware WA@UER 1M PLATE. xX. HELIOTYPE PRINTING CO. BOSTOR i p MINNESOTA BOTANICAL STUDIES. PART IW. VOL. If. PLATE XI. HELIOTYPE PRINTING CO, BOSTON VOL. II. MINNESOTA BOTANICAL STUDIES. PART teh PEATE: Xi HELIOTYPE PRINTING CO, BOSTON ey. ir PTT P > ne Ww me = . 4 ots ava fice et, } = 4 - i a . ; - 5 Fink : CONTRIBUTION TO THE LIFE-HISTORY OF RUMEX. 153 have degenerated, the fecundated egg is extending its cell wall up- ward to attach it to the anterior end of the sac and the definitive nu- cleus is dividing. 27. Posterior end of the sac whose anterior end is shown in 26. The antipodals are degenerating. 28. Embryo sac and surrounding tissue of the nucleus showing mutual relations of the two structures. 29. Anterior portion of embryo sac showing three of a probable four endosperm nuclei, pollen tube ruptured at the end and the egg di- viding to form the two-celled embryo. Plate XIE. 30. Anterior end of an embryo sac somewhat younger than that fig- ured in 29 and showing the discharged pollen tube, the egg apparently preparing to divide, and four endosperm nuclei with a spindle persist- ing between each pair. 31. Anterior end of embryo sac showing a five-celled embryo and three endosperm nuclei. 32. Young embryo in anterior end of embryo sac showing four newly formed nuclei and two persistent spindles. 33. Embryo sac of Rumex salic¢folius showing a somewhat more advanced embryo, a larger number of endosperm nuclei and the anti- podals still persisting. 34. Anterior end of embryo sac showing five-celled embryo, one of four observed dividing endosperm nuclei, and an undischarged pollen tube which entered the sac after the egg was fecundated. 35. Anterior end of embryo sac of Rumex salictfolius showing three-celled embryo, whose middle cell is dividing, and the persistent discharged pollen tube in the micropyle and extending into the sac. XI. OBSERVATIONS ON GIGARTINA. Mary E. Otson. The genus Gvgartina is of wide distribution, especially in temperate latitudes. It is found both in the Atlantic and Pa- cific oceans as far north as the coast of Greenland and as far south as Cape Horn. The material used in preparing this paper was collected in Puget Sound, at Channel Rocks, near Seattle, Washington, on August 3, 1897. It does not correspond exactly with any of the specific descriptions recorded. Indeed it has been with some reluctance that this plant has been included in Gzgartcna. The material used for study in the preparation of this paper had been preserved in 75 per cent. alcohol, with the exception of some dried material which was studied in determining the color, size, shape and other external characters. In the dried material the fronds appear thin and membrana- ceous and very brittle. The alcoholic material is leathery in texture and quite tough. ‘The older fronds show a considerable increase in thickness over the younger omes, and are strongly Calliblepharis-like in appearance. HaBpiIT AND EXTERNAL APPEARANCE. The fronds are reddish purple in color and occur at a depth of eight fathoms. Several or more fronds are generally found growing crowded together from united holdfasts. Such a group is seen in Fioltm, Pl. 13. The general outline of a frond whether branched or un- branched is typically cuneate. Fig. 3, Pl. 13 shows a good specimen of the unbranched type. When branched the general wedge-shaped outline is retained by a more or less regular, dichotomous form of branching, in which the branches spread and remain of considerable width. (Fig. 1, Pl. 13.) Olson: OBSERVATIONS ON GIGARTINA. 155 fToldfast.—The holdfast is disc-like, and varies in size accord- ing to the number of plants springing from it. In an isolated in- dividual (Fig. 3, Pl. 13) it is seen to be but little larger than the circumference of the base of the stipe. On the under side, by which it is attached to the substratum, it is seen to have a smooth surface, and two areas are clearly distinguishable in the alco- holic material used in this study; a central, nearly circular al- most translucent area, and an outer apparently denser portion (Fig. 3, Pl. 13). When viewed from the upper side the signi- ficance of these areas is understood. If a large holdfast be ex- amined, from which part of the fronds have been removed, the upper surface will be seen to be uneven and dotted with circu- lar pits bordered by very distinct rims (Fig. 6, Pl. 13). On running the point of a needle through one of the translucent areas of the under side it is found to correspond with the pit of the upper side. A still more interesting demonstration is to pull away one of the fronds still attached to the holdfast. If this be done carefully it will be seen that a characteristic pit remains to mark the point of attachment of the stipe. An old holdfast is found to be covered with these scars, which are very perfect markings of the outline of the stipe at the point of attachment. Stipe.—The stipe is a well-marked organ whose presence is more or less evident in all the plants, especially as they attain their mature size. Some of the members of the group in Fig. 1, Pl. 13, show that in the first stages the width of the frond varies but little from the base to the tip, so that a distinct stipe is scarcely distinguishable. Very soon, however, the upper por- tion of the frond begins to expand and a typical stipe becomes evident. Its outline just at the point of attachment to the hold- fast is circular, but above this it becomes slightly compressed in one diameter so that its cross section appears oval or elliptical. The transition trom stipe to lamina is so gradual that no distinct demarcation can be detected between them. The shorter diam- eter becomes still shorter, and the longer one increases to the width of the frond till all appearance of the stipe has van- ished and even the greater thickness at the center of the lamina merges so gradually into the thinner margins that it cannot be be said to be present as a midrib. Lamina.—The lamina is seen to attain its greatest width at some little distance from the tip. If it is branched the division is dichotomous, though the lobes are often unequal in size, and 156 MINNESOTA BOTANICAL STUDIES. occurs near the tip of the frond, so that there is no branching of the stipe. The margins present a more or less wavy, undula- ting outline. In fertile fronds the margins all along the upper part of the frond are prolonged into little leaf-like outgrowths bearing the cystocarps (Fig. 2, Pl. 13). These proliferations also occur on the surface of the frond and sometimes are scarely more than the stalk of the single cystocarps they bear. In many fertile fronds the surface is almost entirely covered with these outgrowths. Asa rule they occur much more densely on one side than the other. Frequently they are found as simple small leaflets bearing no cystocarps. ‘Their presence gives a look to the lamina quite suggestive of fruiting Callblepharis. ‘INTERNAL ANATOMY. Floldfast.—A section of the holdfast shows a very distinctive structure unlike anything seen elsewhere in the plant. The ap- pearance of a section is seen in Fig. 9, Pl.) 1g; ishowmeuine outline of the pit or scar and the depression from which the stipe was removed. ‘The cellular structure varies but little from the general type except in the transition zone from the holdfast to the stipe. Along the upper surface the cells are covered by a gelatinous envelope or cuticle of considerable thickness often 15 mic. deep (Fig. 10a, Pl. 13). This, of course, in one of the scars extends only to the border of the pit as indicated in Fig. g, Pl.13. Some sections also show a similar layer on the lower side at joints (Fig. 9 b, Pl. 9), while the remainder of the lower surface shows a rough irregular margin of cells (Fig. 9 c, Pl. 13). The possible explanation suggests itself that the cuticle, when occurring on the lower side, appears at joints which, through some unevenness of the substratum, are not closely ap- pressed to it and hence are left exposed as it were. The tissue of the holdfast is characterized by cells having a quadrilateral outline as seen in zone ‘‘ b” in the right-hand part of Fig. ro, Pl. 13. This general type becomes greatly modi- fied in various regions. In the transition zone ‘‘c” (Fig. 10, Pl. 13), from the holdfast to the stipe, the angular outline of the cells disappears and they become more rounded and smaller in size. In all regions cells of irregular outline are frequently met with scattered among the cells of typical quadrilateral out- line. In general the cells are arranged in approximately regular Olson : OBSERVATIONS ON GIGARTINA. alia 7/ rows extending vertically through the holdfast. This is more apparent outside the region of the pit than within. In passing from the holdfast proper to the stipe the character of the cells is seen to change, the outline becomes rounded, the cells smaller and the arrangement very irregular (Fig. 10 c, Pl. 13). On the upper side of this zone the cells become elongated and soon merge into-the structure of the stipe (Fig. 10 d, Pl. 13). At the side, in the region where the surface of the holdfast passes to the surface of the stipe, an interesting curvature of the rows of cells of the holdfast is noticed (Fig. 10 e, Pl. 13). The upper cells of the holdfast gradually become smaller and merge into the outer cells of the stipe, so that at the periphery there is not so marked a change in the character of the cells as in the center. The central elongated cells of this stipe extend farther into the holdfast in the center than at the periphery of the stipe region. This is to be expected from the outline of the pits from which the stipe has been removed, and an examination of these pits under high power reveals the fact that none of the elongated stipe cells are present in the scar, showing that the separation zone when a stipe is pulled from a holdfast is at the region shown in Fig. 10, Pl. 13 at c, and curves upward at the sides, thereby forming the pit-like scar. In many sections of stipe and holdfast the elongated cells are seen to extend much farther down than in the section of Fig. 10, Pl. 13, so that only one or two layers of holdfast cells lie between them and the lower surface. In a few sections an interesting development of outgrowths on the lower surface was noticed. These occur below the stipe region and show the same cellular structure as the holdfast proper. They are apparently rhizoid-like growths (Fig. 10, Ps. 03). The sections used were placed in an alcoholic solution of fuchsin for a few moments, then washed with alcohol and mounted in glycerine jelly. The stain failed to bring out any cell contents and to all appearances the cells are empty. Stipe and Lamina.—The structure of the stipe and lamina is very similar, the chief difference being that in the former the cells are of somewhat smaller diameter. The upright portion of the plant may be divided, anatomic- ally, into two fairly distinct portions: the pith, consisting of the larger, apparently empty cells and the cortex, of smaller 158 MINNESOTA BOTANICAL STUDIES. cells containing the chromatophores. Because of this dif- ference in the cells the sections for the structure of the stipe and lamina show the outlines of the cell wall in the pith, but in nearly all the sections examined the cell walls of the cortex could not be distinguished, and it is the outline of the cell con- tents that is represented in such sections. The inner portion or pith consists of elongated, cylindrical cells, united into loosely interwoven filaments, extending prin- cipally in the direction of the long axis of the frond. The union between cells is so irregular that often the filamentous arrangement is scarcely recognizable. Surrounding this pith region, which is of compressed cylindrical contour in the stipe, is the cortex, which consists of much smaller cells, arranged in radiating rows, more or less regular, perpendicular to the sur face of the frond. Pith.—The transverse sections of both stipe and lamina show the cross-sectional outline of the pith cells to be more or less circular with considerable space between the cells (Figs. 12 and 13, Pl. 13). In the lamina portions of a filament are often found running through the section. The outlines of the pith cells in both lamina and stipe show an area of larger cells just within the cortex, passing inward to a central portion of smaller diameter (Fig. 11, Pl. 13). Within the pith itself there is con- siderable variation in the size of the cells, showing that smaller filaments anastomose with the larger ones. The average size of the cells is from 100 to 170 mic. long by 17 to 33 mic. wide. The most interesting feature of the pith is the presence of protoplasmic pits connecting the cells. These occur not only in the end walls, but also in the lateral walls, as seen in Fig. 11, Pl. 12. By these connections, as well as by lateral pressure in some cases, the cylindrical outline of the cells becomes variously modified. Communication of adjacent cells of the pith region is thus completely established. The significance of this will be more clearly seen as the physiological importance of this area is dis- cussed. These pits were first discovered by staining with hama- toxylin. A more careful trial of different stains showed that these pits always take the stain more deeply than any other part, either cell contents or cell wall. Both methyl violet and fuchsin produced good results. An alcoholic solution of the stain was used. Olson : OBSERVATIONS ON GIGARTINA. 159 A study of these pits showed them to be of the form of small plates or rings, apparently one in each of the two adjacent cells (Figs. 11 and 13, Pl. 9). When seen edgewise they appear as two small plates narrower than the width of the cell wall, so that the inner line of the cell wall appears to curve out to meet them. Often the entire outline of one ring may be seen and only part of the other, which apparently lies beneath it. Again the sec- tion will lie so as to show both rings. Some sections, especially with the methyl violet, showed a faint outline of cell contents just within the wall and in all cases extending close up to the rings or pits. Careful observations of this sort led to the opinion that these connecting pits wust be of the nature of the protoplasmic cell contents rather than the cell wall. Schmitz confirms this opinion. Unfortunately the writer did not have access to Schmitz’s original article, but in George Murray’s /ntroduction to the study of Seaweeds, 1895, in the chapter on Rhodophycez (which he states is based upon Schmitz’s papers) the following is found (p. 201): ‘* The plates stand in direct connection with the protoplasm lining the cell wall and are, in fact, so coherent with it that they may be regarded as transformed or rather differentiated protoplasm lo- cally covering the pit. However it is probable that a thin layer of protoplasm covers them in turn.” All observations have gone to show that there is an intimate protoplasmic connection between the contents of neighboring cells. Zimmermann’s Botanical Microtechnique was consulted as to re-agents for testing these rings. The use of sulphuric acid and a mixture of iodine and potas- sium iodide is recommended for cellulose walls giving a blue color. This was used, but neither the cell walls nor the pits showed any trace of blue staining. Cuprammonia was also tried, but with no success. Some interesting results were ob- tained, however, with the use of sulphuric acid. The sections were first stained and then treated with the acid. Although the acid at once destroyed the original color it was found better re- sults were obtained than without first staining. A trial was then made as to the strength of acid which would give most satisfac- tory results, and it was found that treating sections prepared as before described, with a 50 to 60 per cent. solution of the acid produces at first no apparent effect beyond a slight swelling of the cell wall. The sections were left mounted in the acid, and 160 MINNESOTA BOTANICAL STUDIES. after twenty-four hours re-examined. It was then found that the cell walls were all dissolved and only the rings remained (Fig. 17, Pl. 14). This leads to the conclusion that the rings are not of the same composition as the cell wall. The pith of the cystocarpic proliferations of the frond shows a marked difference from that of the vegetative portion (Fig. 18, Pl. 14). The cells have become very irregular in outline and are so anastomosed and interwoven as to form a network which becomes more and more dense in passing from the stalk to the pericarp proper. . Here, as well as in the cortex, the cell contents havea dense granular appearance, and the cell walls appear only very faintly, if at all. In most sections stained as in the vegetative part they cannot be distinguished. This is true also of the protoplasmic pits, though it is evident there is close communication throughout. From one or two unusually clear sections it was ascertained, however, that the rings are present, but are very small. The cells in this region measure from 25 to 37 mic. long by 2.5 to 7 mic. wide. Cortex.—The transition from the pith to the cortex is some- what abrupt. In longitudinal sections the pith cells are seen to decrease in length until in the four or five outermost rows the outline of the cells is spherical or slightly oblong. In the transi- tion zone, or the inner part of the cortex, they measure from 5 to 12 mic. along either diameter. In the outermost layers of smallest cells, measuring from 2.5 to 5 mic. in diameter, the cell contents are very dense and the cells are apparently imbedded in a gelatinous matrix from which it is impossible to distinguish their walls. In one section, how- ever, the writer was able to make out faint outlines of the walls, but it is difficult to represent them and maintain the proportional thickness of the wall (Fig. 11, Pl. 13). The cortex cells are seen to lie in communication also, but only along the radial lines of the thallus. The cells are so small no rings can be distinguished, but protoplasmic threads are seen running from cell to cell (Fig. 13, Pl. 13). There are no lateral protoplasmic connections between cells. A surface view of the thallus shows a somewhat regular arrangement of the end cells of these radial rows. They appear as a rule in groups of two or occasionally three, surrounded by the gelatin- ous matrix (Fig. 14, Pl. 14). If the sections be placed in water this swells rapidly, as do also the cell walls. The walls often Olson: OBSERVATIONS ON GIGARTINA. 161 increase to three or four times their width, as seen in alcoholic material, and a stratification of the walls becomes evident (Fig. EOn ae a4). REPRODUCTIVE ORGANS. Cystocarps.—The material studied was too far advanced to show antheridia or the development of the cystocarp. The cystocarps are found scattered very abundantly along the margin and over the surfaces of the fertile fronds. As a rule they are much more abundant upon one surface than the other. They are borne in leaf-like proliferations of the frond and are usually more or less distinctly stalked, though often they are nearly sessile. Fig. 4 shows one of these leaf-like outgrowths bearing no less than nine cystocarps. Generally the number is smaller, from two to four (Fig. 5, Pl. 13). The larger leaflets, with a larger number of cystocarps, are usually found along the margin. In form the cystocarps are subglobose with a marked indenta- tion at the apex which seems to indicate a distinct carpostome, but in the large number of sections observed no opening could be detected. The nearest approach to it was seen in the section represented in Fig. 19, Pl. ro, but even here there is no sign of a true pore or even of a rupture, so that evidently the cystocarp is closed. From the uniform closure of the cystocarp it is diffi- cult to include the plant in question with Gzgartzna. The struc- ture of the cystocarp regions of the thallus shows the same two general areas described for the stipe and lamina with the modi- fication of the inner region or pith described above. The spores are developed within the central region, the cortex and outer part of the pith forming a true pericarp. Except at the apex both areas surround the central mass of spores. Here it is covered only by the cortex. A peculiar structure observed was one in which the surface of the thallus was only slightly raised to indicate its location, and numerous: long filaments were seen with their tips protruding slightly from the surface (Fig. 18, Pl. 14). The section was stained with fuchsin and the clear filaments were sharply dis- tinguishable from the other cells with their granular contents. They measured 150 mic. in length. There was some little doubt as to what should be the interpreta- tion of the section represented in Fig. 18, Pl.14. The pith cells 162 MINNESOTA BOTANICAL STUDIES. have the appearance characteristic of the cystocarp region. A comparison with figures in which trichogynes are represented shows but slight similarity in appearance. The pericarp consists of the two layers found in the lamina and stipe. The outer small cells containing chromatophores pass somewhat abruptly to the cells of the interior (Fig. 21, Pl. 14), which are elongated and connected to form a more or less dense net-work. In most cases the transverse connecting cells are more numerous than in Fig. 21, so that the pericarp presents the reticulated appearance of the tissue in Fig. 19. Very frequently, however, the cells show much lateral crowding in the pericarp. The cystocarp is compound and the spores are aggregated into distinct groups (Fig. 19, Pl. 14). This is clearly seen in all but the oldest cystocarps and even here a carefully cut sec- tion shows it. These groups are separated from each other by large, empty cells, with smaller cells of the same character ex- tending between them (Fig. 20, Pl. 14). This is brought out very clearly by staining the section with iodine and then wash- ing in water. The carpospores are more or less oval in shape, often some- what angular. They measure from 12 to 15 mic. along one diameter by 10 to 12 mic. along the other. The normal cystocarps measure from 1 to 2 mm. in diameter, but it was noticed that frequently some were met with from two to three times as large as the ordinary ones, measuring 2.5 to 3.5 mm. On examination it was discovered that the fronds of another small alga were always found upon these large cysto- carps. Several specimens were studied and it was found that there was evidently more than one species infecting them. The largest one discovered is represented in Fig. 7a, Pl. 13. It ap- pears to emerge directly from the apical depression of the cys- tocarp. 120-121. 1894.) Lunds Univers. Arskr. 27: 41. 2 Di gaGgo[or. Wille, N. Beitrag zur Entwicklungsgeschichte der physiologischen Gewebesysteme bei einigen Florideen. Nova Acta Acad. Leop.- Carol. 52: 49-100. fl. 3-8 1887. (Bot. Jahresb. 16: 754. 1891.) Osterhout, W. J. V. A Simple Freezing Device. Bot. Gaz. 21: 195-201. 1896. Schmitz, F. Die Gattung Actinococcus Kiitz. Flora. 77: 367—- ATG, Pl 7o. 1Og2. Olson: OBSERVATIONS ON GIGARTINA. 167 Moore, S. Le M. Studies in Vegetable Biology.—I. Observations on the Continuity of Protoplasm. Journ. Linn. Soc. 21: 595-621. fi. 79-27. 1886. EXPLANATION OF FIGURES IN PLATES XIII. anp XIV. All drawings were made from material preserved in alcohol. Fig. 1. Group of fronds springing from a common holdfast. One- half natural size. Fig. 2. Upper portion of fertile frond showing marginal and sur- face proliferations with cystocarps produced upon them. One-half natural size. Fig. 3. Young frond showing typical shape and single holdfast. One-half natural size. Fig. 4. Small leaflet with cystocarps. x 4.5. Fig. 5. Four cystocarps on a smail stalk-like proliferation. x 4.5. Fig. 6. Surface view of holdfast showing scars. x 4.5. Fig. 7. Cystocarps infected by a parasite. x5. Fig. 8. Surface view of nemathecia. x5. Fig. 9. Section through a scar of the holdfast: a@, layer of cuticle on upper surface; 4, region of the lower surface covered with cuticle ; c, rough broken lower surface. x 5. Fig. to. Section through the holdfast with rhizoid-like outgrowth from the lower surface. A portion of the tissue is omitted in the cen- ter, as it issimply a continuation of that represented on either side. Toward the left the drawing stops in the stipe region. On the right the transition from the stipe to the holdfast is shown: a, cuticle of upper surface; 4, region showing cellular structure characteristic of the holdfast; c, cellular structure marking transition from tissue of holdfast to that of stipe; d, beginning of tissue of the stipe; e, curva- ture of cells at juncture of stipe and holdfast. x 300. Fig. 11. Longitudinal section of the lamina: a, cortex; 64, outer region of pith showing larger pith cells; c, inner region of pith with smaller cells. x 300. Fig. 12. Cross section of stipe: @, cortex; 6, larger celled pith; c, smaller celled pith. x 300. a Fig. 13. Cross section of the lamina showing many pith cells con- nected by pits and cortex cells connected by protoplasmic threads. X 300. Fig. 14. Surface view of thallus. x 300. Fig. 15. Outline of cross section of stipe showing cortex and pith BReAS., XX A-5. Fig. 16. Pith cells of lamina stained with methyl violet and mounted in glycerine. Cell wall is much swollen and one shows stratification. x 300. 168 MINNESOTA BOTANICAL STUDIES. Fig. 17. Effect of sulphuric acid on cell wall and pits; @, pith cells treated for ten to fifteen minutes in 50 per cent. solution; 4, pits as seen twenty-four hours after treating the section with the acid. The walls have been entirely dissolved. x 300. Fig. 18. Cluster of filaments. x 300. Fig. 19. Longitudinal section through a cystocarp. x 85. Fig. 20. Group of spores from cystocarp with thin-walled cells separating them. x 300. Fig. 21. Longitudinal section through the pericarp: a, cortex; 5, pith; c, spores. x 300. Fig. 22. Longitudinal section through an infested cystocarp. The upper left-hand portion shows the manner in which the parasite pene- trates the host. x 56. Fig. 23. Portion of the same region enlarged showing to the left the tissues of the host pericarp and to the right the long filamentous cells of the parasite. x 300. Fig. 24. Section through frond producing nemathecia. On the upper surface is a mature nemathecium, on the lower surface two two younger ones: a, gelatinous cuticle; 4. pore-like break in cuticle; c, cortex cells; d, cells from which tetraspores are produced; e, para- site; f, pith filaments. x 300. WOOL. (IT. MINNESOTA G2 E> Yow © CC. Bry OO|bio|Ood0 12 © © J) HO oie © hn, © Oreos 332099 e359 o8 00! en O50 ary iy H co 4 3 ° a i Go. ry $9959 90; os eee f ITANICAL STUDIES. . PART ( ae} : ee 14 16 i> ute = ate °o eer 8 a2 es Pe HELIO! eA aie \NICAL STUDIES. S he O°, a EB XIV. NTING CO, Boston > A 7 ee oe 7 ° = wn : cn a { : be | { z F : 6 7 i ¥ é * Sit te 7" f Ee ‘: : r J e eae " » 4 t? % " . x . x wn" ' a . re < t te . a Son ivte ‘ = q re XII. SEED DISSEMINATION AND DISTRIBUTION OF RAZOUMOFSKYA ROBUSTA (Engelm.) Kuntze.* D. T. MacDouca. The branches of the bull pine (P:nus fonderosa scopulorum), of the southwestern United States, offer suitable conditions of nourishment for the growth of /tazoumofskya robusta, a parasite belonging to the Loranthacez. Some of the members of this family, such as the mistletoe (Phoradendron flavescens), which live on deciduous trees in temperate latitudes, are furnished with a fair amount of chlorophyll. These forms are able to carry on more or less food-formation during the warmer portions of the season in which the deciduous hosts lack leaves. /tazoumofskya, however, fastens on an evergreen conifer, and hence has no such need or use for chlorophyll. Itis, therefore, furnished with this substance in minute quantity only, and its leaves are reduced to mere bracts. It is dicecious, and the aérial shoots of both kinds may appear in close contiguity on the same branch of the host or be separated some distance. The shoots start up from the submerged rhizomes in the latter part of April or early in May, the flowers maturing in June and the seeds in August. After the dispersal of the seeds the aérial portion of the plant dies away, leaving only the haustorial rhizomes buried in the tissues of the host plant. With the opening of the next season shoots are produced as before. The submerged portion of the parasite penetrates the branches of the host long distances longitudinally, and where aérial shoots are given off the tissues of the host show abnormal structures, the branches undergoing enlargement, while the development of the nearest buds is variously checked and altered. The dis- tortion is magnified with age, and old trees exhibit the most grotesque malformations. The writer has seen trees a meter in height infected, and the size of the older branches bearing the *An abstract of this paper was read before the Botanical Club of the A.A.A.S. at Boston, August 25, 18908. 170 MINNESOTA BOTANICAL STUDIES. parasite is such as to justify the statement that they may live ten to twenty years after the parasite has fastened upon them. A common type of structure resulting from the attachment of the parasite to the pine consists of an old branch bent downward, from the infected tip of which numbers of smaller branches stand erect, forming a coarse ‘‘ witch’s broom” (See Plate XVI.). Perhaps the most interesting facts in connnection with the history of /teazoumofskya, are those which concern the distribu- tion of the seeds. The single-seeded berries are borne on short stalks curved semi-circularly, from which they are easily de- tached when ripe. The berry is joined to the stalk by a scission layer, which is ruptured by the slightest touch or may be burst away by the action of forces set up in the berry, which also expel the seed. The shooting of the seeds from the berry has been known for many years, and a note of the fact has found place in American text-books of systematic botany, but it has failed of wider recognition. Engler and Prantl remark con- cerning the seeds of the Loranthacee : ‘‘ The stickiness enables some seeds falling from branch to branch to become attached ; on the other hand, birds crush the fruits and discard the seed, which is surrounded by a viscid layer.” (Naturlichen Pflan- zenfamilien. Theil III.) Kerner says: ‘* The dissemination of the European mistletoe is effected as in all Loranthacez through the agency of birds, thrushes in particular, which feed upon the berries. and deposit the undigested seeds with their ex- crement upon the branches of trees.” (Nat. Hist. of Plants, I: 205, 1894.) Keeble, the most recent observer who has pub- lished upon the Loranthacee, says: ‘‘ The berry-like fruits of the Loranths are technically speaking indehiscent; yet owing partly to growth of the embryo, partly to weakening of the fruit wall in some species, this latter becomes ruptured on the ripening of the fruits, e. g., Loranthus neelgherrensis L. cuneatus; in others a very slight pressure is sufficient to cause the complete extrusion of the seed, sometimes basally, some- times apically. In all cases the seed slips out, but in Vescum ortentale Willd., a gentle pressure will cause the fruit wall to crack and the seed to be jerked out.” (Observations on the Loranthacee of Ceylon, Trans. Linn. Soc. Lond. 2nd Ser. Bot:5 et: 3,8) 076.2090.) In view of the above statements it is safe to conclude that Razoumofskya is the only Loranth furnished with a mechanism | MacDougal: DISTRIBUTION OF RAZOUMOFSKYA ROBUSTA. 171 for the expulsion of the seeds from the berries without the in- tervention or coéperation of outside factors. The expulsory mechanism is best seen in a longitudinal section of the berry. The base of the berry is joined to the stalk by a scission layer several cells in thickness. The outer coat is firm and smooth, and is composed of an epidermal layer with the outer wall ex- tremely heavy and cuticularized. Beneath the epidermis is a mass of parenchymatous tissues, the outer portion of which is slightly palisaded and containing chlorophyll, the inner layer showing only starches and sugars. Immediately internal is the fibrovascular framework which fuses at the apex in a solid mass of mechanical tissue. Lying inside the fibrovascular strands, and continuous with the parenchymatous tissue external to it is also a mass of similar thin-walled elements of ovoid or cylindri- cal form rich in carbohydrates. These cells have their axes at right angles to the surface of the berry. The second layer in- ternal to the fibrovascular tissue is the expulsory layer, consist- ing of very long thin-walled cylindrical tubes with their axes parallel to the long axis of the berry at the apex of the seed or variously inclined from this position according to the location, but all so arranged that their longitudinal expansion would tend to force the seed out of the mouth of the sac formed by the berry. Immediately coating the seed is a layer of globoid cells with thick mucilaginous contents. The seed has the form of a modern rifle bullet, conical at the basal end and truncate at the apical end, with a general cylindrical outline. The scission layer appears to cut into the mucilaginous layer or at least very nearly so in the mature berry. During the ripening period the contents of the expulsory layer undergo such chemical changes as to give the contents a very high isotonic coefficient. The consequent osmotic attraction of water into this layer sets up a turgescence which could not be measured, but which probably amounted to many atmospheres. ‘The steady increase of the turgidity of the expulsory layer brings the tension to the breaking strain of the scission layer, and its sudden and complete rupture permits the full force of the pressure to act upon the seed, send- ing it to a distance of two or three meters. The entire arrange- ment is that of a mortar cannon. The muzzle of the gun is sealed by the stalk, and the charge amounts to several atmospheres, which is allowed to act upon the seed when the muzzle is freed. The firing of this unique 172 MINNESOTA BOTANICAL STUDIES. gun may result from the overcoming of the resistance of the restraining layer at the muzzle, or this event may be precipi- tated by any force from the outside which would result in the disturbance of the scission layer. One may stand under a pine tree on a quiet morning and hear the sharp click accompanying the expulsion of the seed from the berries at irregular intervals. If the branches are jarred or shaken, however, the irregular ex- plosions give way to fusillades by which nearly all of the berries on a plant will be set in action at once. The expulsion of the seed occurs as soon as the berry has broken loose from the stalk, and as these berries were originally in all positions the seeds are sent out in all directions. The mucilage adhering to the seed causes its attachment to the branches or other bodies it may strike. In this manner dissemination is effected throughout a cylindrical space about seven meters in diame- ter and extending downward to the ground. The only localities which offer suitable condi- tions for the germination and growth of the seeds, however, are the tips of branches or the shoots of young trees underneath. It is to be seen that no animals are to be found in the habitat of the parasite which would in ordi- Dre. & nary usage carry the seeds to these locations. Seed of azoumof. The only part, therefore, that animals play in skya adhering to the dissemination of the seeds would be in BER causing the discharge of the berries, a matter of no direct value, since they are capable of quite as efficient action independently. The berry of /tazoumofskya is, there- fore, to be classed as a sling fruit, and is probably the only one of this class from the United States which has been de- scribed, though many doubtless exist. A second point of interest in this plant consists of a fact bear- ing upon its local distribution. During the course of some re- cent field work in northern Arizona the writer found that Razoumofskya was most successful in its attacks on the pine trees along the rims of cafons or along the brows of hills or margins of mesas. A study of the meteorological conditions shows that this method of distribution has a direct connection with the vertical movements of the air. As the air resting on lowlands in cafions or valleys is warmed MacDougal: DISTRIBUTION OF RAZOUMOFSKYA ROBUSTA. 173 by radiation during the time of exposure to the sun’s rays, it rises and expands. During the ascent some heat is converted into the work necessary in expansion, causing a cooling of one degree Fahrenheit for every one hundred and eighty-eight feet of elevation. The decrease in temperature lowers the dew point or increases the relative humidity, a matter of very great im- portance to germinating seeds and transpiring leaves. Aeazou- mofskya is especially abundant, precisely at the places where the effect of the ascending humid currents of air is greatest, along the margins of hills and mesas and the rims of cafions. This is very noticeable along the Grand Cajion of the Colorado river, in the Coconino Forest reserve, where the air rising more than a vertical kilometer from the river bed pours across the pine-covered mega at a much lower temperature and very much nearer the dew point than the body of air which it replaces. In its rise it has lost heat at the normal adiabatic rate to the amount of about twenty-five degrees F., and has undergone a great variation with respect tothe dew point. Asa consequence of the increased humidity favorable to germination, the pines near the rim of the cafion are most thickly infested with the parasite over a belt one to four or five kilometers in width run- ning parallel to the margin. One may walk through the forest and note the decreasing abundance of /rtazoumofskya as the dis- tance from the cafion increases. In recapitulation of the facts adduced in this note it is to be said that the berries of /teazoumofskya are to be classed as sling fruits, the only one from North America hitherto described, and that this genus is the only one of the Loranthacee furnished with means of seed-disseminaticn independent of gravity and animals. The writer also believes that he is justified in an- nouncing the discovery of the influence of vertical air-currents upon the distribution of plants, and that this factor must be taken into account in the consideration of the boundaries of zones in mountainous regions or those with irregular topography. EXPLANATION OF PLATEs. Plate XV. D. Staminate plants of Razoumofskya. B. Pistillate plants with mature berries. The distortion of the branch of the host is plainly shown. Plate XVI. A. Prxus ponderosa dying from the effects of the para- site, photograph of a specimen growing on the extreme edge of the rim of the Grand Cajion of the Colorado, June, 1898. C. Specimen of Prxus ponderosa showing drooping of branches attacked by Razoumofskya. vy ta Spa mitone 4% ath nt & ae hPL < Rieite Wop sea. VA =e is 2 ae 7 , © —[iF - é 4 M's - Z * ve : x re ‘ Nh, ah Sa a (tis VOL. II. MINNESOTA BOTANICAL STUDIES. PAIR OXeVE. THE HELIOTYPE PRINTING CO. BOSTOR PART le WHOM Ee MINNESOTA BOTANICAL STUDIES: evenlega bt PE AW Ey evil: THE HEL'OTYPE PRINTING CO BOSTON XIII. OBSERVATIONS ON CONSTANTINEA. E. M. FREEMAN. flistory and Literature.—The earliest mention of the red seaweed now classified under Constantinea is found in Gmelin’s* flistoria Fucorum published in 1768, in which he describes Fucus rosa-marina from the material collected by G. W. Steller during the years 1742-1745 at Kamtschatka. The description is as follows: ‘‘Peculiare sistet hec planta fuci specimen, cujus exemplum aliud in omni reliqua fucorum historia non oc- currit. Caulis teres est, carnosus, penne anserine crasitie, ramis sibi similibus, quibus, tanquam totidem pendiculis, adpli- cantur verticillatim folia petaloidea terna vel plura, rotunda, concava, circulo in centro notata, pulchre expansa, plerumque sissa, ramo per illa penetrante, exeunte, et pollicis dimidii inter- vallo nova fronde priori simili, prolifico, tertia nunnunquam pari ratione accedente. Petala convoluta pulchre representant flores polypetalos, ut Rosam, anemonen, cet. Substantia tota gelatinoso-membranacea, aqua dissoluenda, pellucida. Color e rubro flavescens. Magnitudo semipedalis. Locus. Circa Lapatka inter spongias ad Kamtschatcam occurrit.” Such terms as ‘‘ petala convoluta,” ‘* flores polypetalos,” etc., show what a profound impression the superficial resemblance of the described plant to a rose had made upon the author. In the years 1826-1829 the Russian vessel Seniavin, Fr. Liitke, Captain by the order of Czar Nicolaus I., sailed through Russian waters and collected a large amount of valuable algo- logical material. The results were published in 1840 by Pos- tels and Ruprecht in their ///ustrationes Algarum in itinere, etc. The authors in their preface to this work state that the collections of H. Mertens and the plates of Alex. Postels form the basis of the entire work. The genus Constantinea is here described, founded upon Gmelin’s /ucus rosa-marina, and three species are recognized, Constantinea rosa-marina, C. sitchensis and C. reniformis. *Gmelin, S.G. Hist. Fuc. 102. 27.5, f.2, 2¢. 1768. 176 MINNESOTA BOTANICAL STUDIES. According to the descriptions the first two species of Constan- tinea differ in the length of stipe between annuli, the mode of branching of the stipe and in the nature of the edge of the frond. The following are also noted: Constantinea rosa-marina is the smaller (one-half foot or less); branched even to the base ; termi- nal frond two inches in diameter, laciniate (laciniz 3-6), rarely remaining entire; 2-4 laminz under the terminal one laciniate in a similar manner. C. sztchensis has solitary terminal fronds at the apices of the branches; the fronds are 4-6 inches in diameter, entire but laciniate when older; young fronds are often 8 lines in diameter. In these supplementary descriptions all differences are com- promised except the following: length of the ‘‘ internode” of the stipe; the number of fronds on each branch; the method of branching of the stipe and the difference in size. The tetra- spores of C. sztchensts alone are described. The ‘‘ gongyli rotundi” as Kiitzing* has since pointed out are but ordinary cells of the intermediate layers of the frond. Postels and Ru- precht also mention Cozstantinea rentformis, arare Mediterran- ean plant supposedly of this genus. In 1843 Zanardinit described the C. reniformis of Postels and Ruprecht under the name of Veurocaulon foliosum from material collected on the shores of Dalmatia in the Adriatic. In the same year Kiitzing{ called attention to the incorrect view of Postels and Ruprecht concerning the ‘‘ gongyli rotundi” and to the great similarity in vegetative structure but great difference in outward appearance and tetraspore formation between Con- stantinea and Huhymenia. He describes the tetraspores and states that the cystocarps are unknown. His statements are evidently based solely on Postels, and Ruprecht’s observations. In Speczes Algarum, 1849,§ the two arctic species of Postels and Ruprecht are described under (Veurocaulon as JV. rosa- marina and JV. sttchensis. Two years later J. Agardh || accepts the Postels and Ruprecht generic name of Constantznea and adds to the previous descrip- tion of the vegetative parts and tetraspores that of the cysto- * Kiitzing. Phyc. Gen. 400. 1843. tZanardini. Saggio class. 49. 1843. Se CUILZIN On le C= § Kiitzing. Spec. Algar, 744. 1849. || Agardh, J. Spec. Gen. et Ord. Algar. 2: 295. 1851. Freeman: OBSERVATIONS ON CONSTANTINEA. LT carps. Since no new observations nor collections of the arctic species are cited, his generic description of the cystocarp is probably based upon Constantinea rentformis, the Mediter- ranean species. ‘The cystocarps are described as ‘‘ kalidia in media fronde numerosa, clausa, disruptione partis ambientis demum liberata, nucleolis pluribus composita; nucleoli intra periderma gelatinosum hyalinum gemmidia, nullo ordine dis- posita foventes.” The zonate division of the tetraspores is noted. The collections of C. renzformis cited are: In the Ad- riatic sea on the shores of Dalmatia (Meneghini! and Zanar- dini!) and in the Mediterranean sea at Cette (Salzman!) and at Marseilles (Solier!). Nothing new is added concerning C. sttchensts and C. rosa-marina, but C. renzformis is fully dis- cussed. The latter had been collected also by Mertens and de- scribed by him under the name of Aalymenza reniformis. In 1822 it was described by Agardh* under the name of Haly- menia rentformis, and Postels and Ruprecht + describe it as a third species of Constantinea—C. rentformis. In 1822 J. Agardh{ stated that the tetraspores of C. renzformzs had not been found. Harvey’s description of Constant:nea in 1858 § is based upon the observations and literature cited above. The similarity in structure and the difference in external form and in position of tetraspores is noted. The branching of the stipe is described as at first irregular but later dichotomous; the dichotomy, how- ever, is often lost in the abortion of one branch. In 1862 in a notice of a collection of alge made by Dr. David Lyall at Vancouver Island in the years 1859-1861, by W. H. Harvey|| specimens of Constantinea sttchensts with torn laminz which were probably six to eight inches in diameter when perfect were reported ‘‘ adrift on the beach at Victoria harbor.” And Harvey observes that ‘‘ perhaps this is only a luxuriant state of Constantinea rosa-marina.” In Kiitzing’s work of 18674] the genus is again described un- der Weurocaulon and WV. foliosum and JV. rosa-marina are mentioned. The work of Postels and Ruprecht is not cited. *Agardh. Spec. Alg.201. 1822. t+ Postels and Ruprecht. Ill. Alg.17. 1840. tAgardh, J. l.c. § Harvey, W. H. Nereis Boreali-Americana 2: 173. 1853. || Harvey, W. H. Journ. Proc. Linn. Soc. Bot. 6: 172. 1862. ( Kiitzing. Tab. Phyc. 17: 24. p/. 83. 1867. 178 MINNESOTA BOTANICAL STUDIES. Mention is again made of the genus by J. Agardh in 1876.* He had seen specimens of C. vosa-marina from the Museum of St. Petersburg and alsoa plant from Californian shores which he referred to C. sztchensis; but on account of inability to satisfy himself as to the structure of the fruiting bodies he based his descriptions upon those of Postels and Ruprecht. In 1885 Constantinea thiebauti was described by Bornett from a single specimen collected at Majunga onthe north coast of Madagascar. Bornet very properly calls attention to the re- markable range which this genus, with the addition of his new species would enjoy: C. rosa-marina and C. setchensts in arctic seas, C’. renzformzis in the deep waters of the Mediterranean and C.. thiebautz in the tropical waters of the Indian ocean. In 1891 there appeared in the Botanical Magazine of Tokyo ‘*Remarks on some alge from Hokkaido”} in which Con- stantinea sitchensis Post. and Rupr. (?) is mentioned, but the position of the described plants in the genus Constantinea is ad- mitted to be doubtful. In ‘‘ Die Natiirlichen Pflanzenfamilien” § the genus under- goes rearrangement. C. reniformis is restored to the genus Veurocaulon of Zanardini, where probably should also be placed Bornet’s C. thzebautz. The reproductive bodies, as well as the vegetative structure of C. rosa-marina and JV. foliosum, had been studied personally by Schmitz.|| The classification of Schmitz and Hauptfleisch includes under Constantinea the species C’. sttchensts and C. rosa-marina and under /Veurocaulon the single species JV. foliosum. I have no certain knowledge as to whether Schmitz had or had not in possession any material of C’. sttchensis. Collection.—In August, 1897, and again in the summer of 1898 collections of a species of Constantinea, reported as C. sitchensis, were made by Miss Josephine E. Tilden, and it is upon this material that the following observations are based: On August 3, 1897, specimens of C. sttchensis were found * Agardh, J. Spec. Alg. 3: 225. 1876. tBornet. Alg. de Mad. Bull. Soc. Bot. de France 32:18. f. 7. 2. 1885. ¢Okamura, R. Remarks on some alge fron Hokkaido. Bot. Mag. Tokyo. 5: 333-336. 18091. § Schmitz and Hauptfleisch. Nat. Pflanz. I. Teil. Abt. 2: 517, 519, 520, 525. 1897. | Schmitz. Sys. Ubers. der bisher bekannten Gattungen der Florideen. Flora 72: 436. 1889. Lar A Freeman: OBSERVATIONS ON CONSTANTINEA. 179 growing upon holdfasts of /Vereocystis litkeana in 8 fathoms of water at Channel Rocks, near Seattle, Washington. Dur- ing the summer of 1898 large collections were made at many of the stations where Dr. Lyall collected in 1859-1861. Ceol- lections were made at the following places: (1) Fairhaven, Washington. May 25. Washed up on the beach. (2) Near Minnesota reef, San Juan island, Washington. June 5. At- tached to stones on a flat, sandy beach. ‘This and the three following were found just below lowest tide. (3) Near Friday Harbor, San Juan island, Washington. June 5. Attached to rocks on rocky, steep beach. (4) Oak Bay (a suburb of Vic- toria), British Columbia. July 1. On a sandy beach. (5) Esquimalt, British Columbia. July 2. Attached to rocks. The first and last two collections contained abundant tetraspore material. Preservation.—The material collected in 1897 was killed and preserved in 80% alcohol. Owing to the small amount of this material and to the better condition of that collected in 1898, all of the following drawings except Fig. 1 have been made from the 1898 material. The larger part of it was killed and preserved in a 2 per cent. formalin solution in sea water. In this the color was very well preserved. The firmness of the tissues, however, suffered considerably more in the formalin- solution material than did that of the alcohol material of 1897. The formalin material was still sufficiently firm to admit of very satisfactory manipulation. Still other plants were preserved in camphor water, and some in I per cent. chromic acid solution. The camphor material lost its color almost as completely as the alcoholic. It preserved, however, a great firmness, which ren- dered the tissues excellent for section cutting, and especially for hand sections. The gelatinous cell walls, however, were so cleared that they were not as easily defined as in the formalin material. The chromic acid collections were in a poor state of preservation; the tissues were very soft, the cell walls almost invisible and the contents usually, at least partially, disorganized. Methods.—Various methods were employed in cutting the tis- sues. A part of the material was transferred directly from water to 20 per cent. glycerine, thence to a gum arabic solution upon an Osterhout freezing chamber.* Material was some- times placed directly from the sea water on the freezing chamber *Osterhout, W. J. V. Bot. Gaz. 21: 195. 1896. 180 MINNESOTA BOTANICAL STUDIES. with results almost, if not equal, to those obtained when it was passed first into 20 per cent. glycerine. When the sections were removed from the knife they were placed in 20 per cent. glycer- ine, and from this into 50 per cent. and then into absolute glycer- ine; or from 20 per cent. glycerine to water and then through the alcohols to an alcohol stain or directly into a water stain. The freezing device mentioned above proved very satisfactory in many respects. The tissues can be frozen in a minute’s time, are held firmly in place, and the gum arabic, when of the proper consistency, is an excellent imbedding medium. The difficulty experienced with this method of cutting such delicate tissues as are found in many of the seaweeds lies in the handling of sec- tions after they are cut. Especially is this true when it is desir- able to stain the sections and when they must be transferred through several per cents. of alcohol and glycerine. With such tissues as are found in the frond of Constantinea, where they part with great ease, the difficulty is augmented, and it was found almost impossible to preserve thin sections whole. An attempt was made to obviate this difficulty by mounting the sec- tions directly from the microtome knife into glycerine jelly at a temperature sufficient to keep the jelly semi-fluid. This was in part an improvement, but necessitated the mounting of many worthless sections. When sections unstained were mounted in glycerine or glycerine jelly, the great transparency of the swollen cell walls added a new difficulty. Material was also passed into paraffin and stained on the slide and mounted in Canada balsam by the usual methods. Ex- treme care was found necessary, on account of the delicacy of the tissues, to make the stages from one fluid to another by very gradual changes. I was unable to prevent a partial shrinkage of the cell contents. Sections by this method were otherwise quite satisfactory, having the advantage of use in serial work. Staining is necessary in this method since the sections become almost invisible in Canada balsam. Sections cut freehand with a razor or in a hand microtome with the material imbedded in pith have furnished most of the material from which the accompanying plates are drawn. Suf- ficiently thin sections were obtainable in this way with the great advantage of certainty as to the normal condition of the tissues and of speed in preparation. A large number of such sections can be cut and preserved in 2 per cent. formalin for a long time and are ready for use at any moment. Freeman: OBSERVATIONS ON CONSTANTINEA. 181 A number of staining fluids were used, section staining, either on or off the slide proving more satisfactory than stain- ing in toto. Aniline stains were used almost exclusively. The following were employed with at least some degree of success: Aniline blue: Stains the gelatinous wall of the cells pale blue, the chromatophores of the cortical cells deep blue and the contents of the tetraspores and of the paraphyses light blue. ° The central part of the pyrenoids of the endophytic alga which is usually present in these collections takes on a light blue. The best results were obtained with a 4o or 60 per cent. alcoholic solution, acting from 24 to 36 hours. For sections on the slide 5 to 10 minutes in a strong 60 per cent. alcoholic solution was usually sufficient. My best staining results were obtained from aniline blue. Methylen blue: Stains the cell walls, especially the outer portions, which become quite clearly defined. Sections were stained in a strong 95 per cent. alcoholic solution from 5 to 10 minutes. Fuchsin: Best results were obtained from sections left in a ro per cent. alcoholic solution 36 hours. The gelatinous walls were stained light red while the protoplasm of the paraphyses and chromatophores of the cortical cells took up a deep carmine red. Delafield’s hematoxylin: Same strength and time as fuchsin. The granular contents of the cells are stained a reddish purple. Several other stains were used, but without success. In a weak solution of iodine in potassium iodide the granules of the cells of the middle and intermediate layers assume at first a yel- lowish-brown tinge which finally deepens to an amethyst purple. Stained in a strong solution for a few moments these areas take on a dense violet color. Gross Anatomy.—In general form Constantinea presents sev- eral interesting peculiarities. (Figs. 1-6.) The plant is differen- tiated into a cylindrical stipe and peltate frond atits summit. It is of a purple-reddish color, stands upright in the water below lowest low tide mark, and the texture is quite firm and brittle. It is somewhat gregarious in habit. The stipe is ‘terete, branched, ringed and the apex of each branch expanded into an orbicular peltate lamina; stipe 1-4 mm. in diameter, 2-8 cm. in length; lamina 2 cm.—3 dcm. in diameter.” (Tilden, Am. Alg. no. 203. 1897.) The laminae have numerous minute 182 MINNESOTA BOTANICAL STUDIES. dark brownish spots on their upper surfaces. The older fronds are irregularly perforated with circular holes and are more or less torn at the edges (Fig. 3). When young the fronds are en- tire (Fig. 2). Asthe plant continues its growth the stem pushes up through the center of the frond, forming a new growing point which elongates into an internode and soon forms another frond at its summit. The old frond then falls off, leaving an annulate marking on the stipe (Figs. 5, 6). There is often, espe- cially in the youngest portions of the oldest plants, a formation of two growing points at the center of each lamina giving rise to a dichotomy of the stipe (Fig. 5). Irregularity in the develop- ment of these growing points gives rise to an irregular system of branching (Fig. 5). Growing points are not confined to the bases of laminz but may be found occasionally arising from other portions of the stipe. There are often a group of small branches formed upon the holdfast (Fig. 1). The latter is a disc-like body arising as an expanded portion of the stipe at its base and usu- ally concave below (Fig. 22). Upon the lower surface of the fronds especially the larger ones, can be seen distinct radial striations caused by small ridges running from the region near the stipe toward the edge of the frond (Fig. 4). These mark the course of bundles of elongated narrow filaments (see below). The nemathecia which bear the tetraspores are found only on the lower surface of the frond. They occur more abundantly on the larger fronds and are often so numerous as to almost completely cover the under surface. They form small wart-like bodies of whitish color and gelatin- ous consistency. Minute Anatomy.—As is well known the thallus of the red seaweed is a group of dichotomously branching filaments whose fused branches form tissue-like areas. In Constantinea this method of branching can readily be seen in the frond but par- ticularly so in the cortical area of the stipe (Fig. 18). Frond.—A cross section of the frond shows three areas of cells: (@) a central layer of loosely woven filaments; (4) an in- termediate layer of large approximately spherical cells stuffed with starch granules (Fig. 7); (c) a cortical layer of pseudo-pa- renchymatous cells. a. There are in the central area filaments of large cells, usually slightly elongated, often stretching across the frond and perpendicular to the frond surface (Fig. 11). These cells ap- Freeman: OBSERVATIONS ON CONSTANTINEA. 183 proach the large cells of the intermediate layer in form and size, transitional stages between them being abundant (Figs. 8, 9, 10). Their average size is 55 mic. x 12 mic., but they may attain 75 x16mic. A cross section of the lower part of the frond, tangential to its orbicular outline, shows in the central area at more or less regular intervals corresponding to the external stri- ation on the lower surface of the frond, a number of bundles of greatly elongated cells, woven into strengthening bundles (Figs. 7, 8, 9). In the lower region the general course of these bundles is radial in the frond. Divergence from this course is found in the upper part so that a tangential cross section of the frond shows some of these bundles in longitudinal or oblique section (Fig. 9). These filaments are articulated; the cells of- ten attain a length of 325 mic. They average about 8 mic. in breadth. Thin sections of the frond often contain clean-cut cir- cular spaces where the bundles of elongated cells have been pulled out, showing the compactness of the bundles and indicat- ing for it a strengthening function. The bundles vary in size; the striations on the lower surface of the frond mark only the largest bundles. 6. The typical cells of the intermediate area are spherical and average 46 mic. in diameter. They are packed with Flor- idian starch granules which turn brownish with a weak KI so- lution of iodine and finally purple or violet in a strong solution. Those cells toward the surface of the frond contain ordinarily a few chromatophores, usually in the end near the surface. The cells of the intermediate layer shade off towards the surface into the cortical cells. c. The cortical cells are characterized by a compact pseudo- parenchymatous grouping in which the long diameters of the but slightly elongated cells are perpendicular to the frond sur- face. Many of the cells are approximately cylindrical or pris- matic. The external layer is of a sufficient regularity in struc- ture and form to recall forcibly an epidermal layer. In these the outer wall is rounded. The cells of the cortical layer con- tain but a small amount of starch, and this is found in the cells adjoining the intermediate layer. Chromatophores are, how- ever, abundant and more numerous toward the frond surface. Almost all of them are to be found in the first three or four rows of cells from the surface (Fig. 10), but some are found in still deeper layers. They occupy, in the great majority of 184 MINNESOTA BOTANICAL STUDIES. cases, a position in the peripheral end of the cell. Under the microscope they appear finely rose-colored pink with perhaps a purple tinge. They are irregular in outline, apparently in most cases assuming such a shape as will allow them to occupy the peripheral end of the cell to best advantage. The epidermal cells average about 13 x5 mic. and the chromatophores about 10 X2 mic. Stipe.—(Figs. 14-21). A cross section of the stipe shows a similarity in structure to that of the frond. The same areas are present with modifications, however (Fig. 14). The cortical area is composed of cells more elongated than those in the frond and is a larger number of cells deep. Internally these pass into the large cells of the intermediate area (Fig. 19). Here there is a noticeable difference from the condition in the frond. A large number of the elongated central cells find their way into this area and a cross section of the stipe shows them in cross, oblique and even longitudinal section between the large starch-containing cells of the intermediate area which stand out very clearly in large radial filaments (Fig. 21). The central area in the stipe 1s a very compact area and is made up of a large number of thin filaments interwoven in a very com- plicated manner (Figs. 20, 21). These cells correspond to those of the elongated filament bundles in the frond. There are in the stipe, moreover, a small number of larger cells also elon- gated and corresponding to the cells of similar shape and posi- tion in the frond. Incross section the cut-off ends predominate (Fig. 20), while in longitudinal section the cut-off ends are few; the longitudinal view of the filaments is the predominant one and the intricate weaving (Fig. 21) is very plainly seen. A Jongitudinal section through the annulate portion shows the ab- sence of the cortical and intermediate layers in the region of the annulus indicating the continuity of these areas in the frond and stipe (Fig. 15). A longitudinal section through the growing point shows but two areas in the growing region. ‘There is no distinct interme- diate area although a number of large cells may be present (Fig. 16). The two areas are the cortical in which the fila- ments are all parallel, perpendicular to the stipe surface and pseudo-parenchymatous in character, and the central, which is as before a mass of densely woven elongated filaments. The end of the intermediate area of the frond can readily be seen in such a section (Fig. 16). Freeman: OBSERVATIONS ON CONSTANTINEA. 185 FHloldfast.—The holdfast, 7. ¢., the expanded portion of the stipe at its base, presents a similar anatomical structure to that of the stipe with the exception of one modification. The lower cortical and intermediate areas which are in contact with the surface to which the plant is moored are changed into a yellow- ish brown disorganized mass which probably serves as a cement- ing substance in attaching the plant to the rocks. Indications of the former cell structure can be seen in occasional cavities and in the arrangement of these cavities. The area adjoining the cement layer is composed largely of elongated filaments (Fig. 24). Protoplasmic Connections.—In freshly cut material the proto- plasmic connections between cells, is very plainly to be seen in many of the starch cells of the intermediate area. They are also easily seen in the larger cells of the central area, as well as in the small cells of the same area. The cortical area of the stipe furnishes particularly good views of this continuity (Figs. ES, 10). Reproductive Tract (Figs. 11, 12).—I have been unable to find, in the material at hand, and it is considerable in amount, any trace of cystocarp development. The following description of the occurrence and structure of the cystocarp of the genus is translated from Schmitz and Hauptfleisch,* the former of whom has made a personal study of the cystocarp of C. rosa- marina (Schmitz, 1. c.): The carpogonial branches and aux- iliary cell branches are distributed in the fertile portions of the frond in large numbers in the loosened inner portion of the inner cortex of the upper side of the leaf, together with numer- ous vermiform sterile cellular threads. Cystocarps distributed in large numbers on the fertile fronds, comprising a broad zone along the edge of the frond on its upper surface, imbedded in the much loosened inner cortex of the upper surface of the frond, swelling out into an arch the superposed outer cortex which is punctured by pores. The nucleus of the form of a mulberry and pierced by single strands of sterile tissue. Goni- molobes separated only at first, later confluent. Kiitzing ¢ states that the ‘‘ gongyli rotundi” described by Postels and Ruprecht as two kinds of fruiting bodies in Constan- tinea rosa-marina are ordinary cells of the cubcortical layer. * Schmitz and Hauptfleisch. In Engl. and Prantl. Nat. Pflanz. I. Teil. Abth. 27 520,521. 1897: ft Phyc. Gen. 400. 1843. 186 MINNESOTA BOTANICAL STUDIES. Tetraspores are known only in C. s¢tchensts. ‘They are ob- long and zonate, lodged in nemathecia. The nemathecia are found exclusively on the lower surface of the frond more abun- dant near the outer half. They are in the form of delicate, slightly-raised ‘* wart-like,” often confluent bodies of a whitish color. They average 3.5-4 mm. in diameter and often become so numerous that they completely cover a very large part of the lower surface of the frond. ‘The nemathecia are covered by the gelatinous layer on the surface of the paraphyses. The para- physes are elongated, narrow, peripheral cells (Figs. 11, 12). The tetraspores arise as club-shaped elongations of peripheral cells between the paraphyses and are divided zonately into four chambers. The tetraspores are numerous in each nemathecium. Their average size is 108 x 22 mic. Endophyte.—In a large majority of sections and upon all material examined, are present peculiar green approximately spherical bodies imbedded in the cortical tissue of the frond. They are endophytic algz and probably the Chlorochytrium tn- clusum of Kjellman. In general they are pear-shaped with their small end toward the surface and the cell wall at that end thickened. They are greenish in color and contain a number of conspicuous pyrenoids, the central areas of which stain very readily, having a particular affinity for aniline blue. The proto- plasm is denser toward the small end where the cell wall is also thick. It is with this end that the endophyte breaks through the cortical tissues of the nurse plant. Ihave as yet been un- able to detect any zodspore formation. This interesting little endophyte will receive a more complete discussion in a subse- quent paper. Conclustons.—The material upon which these observations are based was distributed by Miss Tilden as Constantinea sttchensis Post. and Ruprecht. A careful comparison of it with the plates and descriptions of Postels and Ruprecht shows however that the plant under observation might as well perhaps be placed under Constantinea rosa-marina. ‘The material agrees in almost every particular with C. rosa-marina hav- ing, however, single terminal fronds and an occasional evi- dent dichotomy of branching. The differences enumerated above in the descriptions of the two species can hardly be con- sidered of specific importance. The length of the internodes may vary considerably. ‘The greater part of the material under Freeman: OBSERVATIONS ON CONSTANTINEA. 187 observation contained long internodes. This material was col- lected in late summer. The number of fronds on a branch is also given specific value by Postels and Ruprecht. It is prob- ably of important significance that xo tetraspores were found by these authors upon C. vosa-marina but that large numbers were found upon C. sztchenszs. Figure 6 represents a young frond having no tetraspores but with a succession of fronds similar to those of C.. rosa-marina, while almost all of the re- maining material had solitary terminal fronds crowded with tetraspores. The material collected in May, 1897, contains tetraspores. The dichotomy of the branching of C. sefchens7s is a com- paratively late development in the growth of the stipe and is not seen in the older parts. C. setchenszs is further described as larger in all parts than C. rosa-marina, though actual meas- urements given do not accord with this. Zhese facts suggest the probability that the C. sitchensis of Postels and uprecht ts the late summer stage of C. rosa-marina. This supposition ex- plains satisfactorily the absence of tetraspores in the one and of dichotomy of the stipe in the other, the comparative lengths of ‘* internodes” and the difference in sizes of the two plants. The observations and impressions of subsequent writers add additional weight to this view. In recounting the founding of the genus by Postels and Ruprecht upon the /ucus rosa-marina of Gmelin, Agardh (1. c. 1851) observes of C. sztchenszs ‘* novo consimili adjecta specie” (p. 294) and of C. rosa-marina ‘‘ pre- cedente (sitchensis) videtur proxima, cum nulla alia confun- denda” (p. 296). Of the material collected by David Lyall at Vancouver island ‘‘adrift on the beach at Victoria harbor” and reported as C. sttchensts, although corresponding in size to C. seéchenszs Har- vey (I. c. 1862) remarks ‘‘ perhaps this is only a luxuriant state of C. rosa-marina.” sae From these facts it would seem, therefore, highly probable that C. sztchens?s and C. rosa-marina are but different forms of the same plant, and since the work of Schmitz has removed C. rentformis to the genus /Veurocaulon where also it is probable that C. ¢hzebauté should be classified, that Constantinea is a monotypic genus, with Constantinea rosa-marina as the only species. 188 MINNESOTA BOTANICAL STUDIES. DESCRIPTION OF PLATES. Fig. 1. Young plant with group of young branches on the holdfast. xy. Fig. 2. Portion of the plant showing the entire edge and form of young fronds (from dried material). x ¥%. Fig. 3. An old frond showing lacerated border and perforations (from dried material). x Y%. Fig. 4. A frond showing striations on the lower surface (dried specimen). x. Fig. 5. Plant with fronds almost entirely cut away showing the growing points, the annulations of the stipe and the dichotomous branching. a@. annulations. ¢.f. growing point. f. frond cut off. xy. Fig. 6. Small branch showing a rapid succession of lamine. x¥%. Fig. 7. Diagram of a tangential cross section of the frond. c. cortical area. zzt. intermediate. e. bundles of enlarged filaments. Z. loosely woven cells of central area. Fig. 8. Cross section of a frond showing bundle of elongated cells in central area in transverse section. Letters as above. Drawn with camera lucida. x 250. Fig. 9. Cross section of a frond showing a longitudinal section of a part of a bundle of elongated central cells. Drawn with camera Plucida., 250. Fig. 10. Cross section of a frond stained for a minute in a strong solution of Tin KI. ch. chromatophores. s¢. starch grains. Drawn with camera lucida. x 345. Fig. 11. Cross section of a frond through a nemathecium. cezz. central layer. . paraphyses. ¢. tetraspores. Cells are drawn only in outline. Contents have been omitted. Camera lucida. x83. Fig. 12. Cross section of a frond through a nemathecium showing tetraspores. Drawn from a glycerine mount in which the gelatinous walls became almost entirely obliterated. The walls are, therefore, omitted except around the tetraspores. x 250. Fig. 13. Cells from central area showing protoplasmic connec- tions. Camera lucida. x 250. Fig. 14. Diagram of cross section of stipe in internode. Letters as in frond. Fig. 15. Diagram of longitudinal section of a stipe through a node. Fig. 16. Diagram of a longitudinal section of the stipe through a growing point. Fig. 17. Peripheral cells from cross section of stipe showing the striations in the outer gelatinous covering, x 345. Tig. 18. Filament from a cross section of the stipe in the cortical WOL. II. Loo Coal PART ANICAL STUDIES. XVII. NG CO, BOSTON “eu MINNESOTA BOTANICAL STUDIES. PART Gi VOL. I. SLD) = @ mae = om SERGI, x BIEDOT EDzioy Gy ——— ———" Cf: i SE PLATE XVIII HEUGTYPE PRINTING CO, BOSTON Freeman: OBSERVATIONS ON CONSTANTINEA. 189 area showing the dichotomy of the branching. Camera lucida for an outline. x 250. Fig. 19. Cross section of stipe including the cortical and the begin- ning of intermediate layer. The cut off ends of elongated central fila- ments are seen in the intermediate area. Camera lucida. x 130. Fig. 20. Cross section of stipe in the central area; shows the pre- dominance of elongated filaments. x250. Outlined with camera lucida. Fig. 21. Longitudinal section of a stipe at the inner edge of the cortex; shows the complication of the elongated filaments. Outlined with camera lucida. x 250. Fig. 22. External view of holdfast seen from below. x. Fig. 23. Diagram of cross section of a holdfast. mm. layer of disor- ganized mass on lower surface by which the holdfast adheres. Fig. 24. Lower portion of the cross section of the holdfast showing the attaching layer m. Camera lucida. x 250. Fig. 25. Diagram of a cross section of a frond showing the distri- bution of an endophytic alga (probably Chlorochytrium). Larger number on the upper surface. ezd. endophyte. Figs. 26 and 27. Cross section of a frond; detail drawing of endo- phyte. w. endophyte wall. fd. pyrenoid. ~. cell of nurse plant in outline. Camera lucida. x 250. BIBLIOGRAPHY. 1. Gmelin, S.G. Historia Fucorum. 1768. 2. Postels, A., and Ruprecht, F. Illustrationes Algarum in itinere circa orbem jussu Imperatoris Nicolai. I. 18, £7. 49, f. 88. 1840. 3. Zanardini, G. Saggio di classificazione naturale delle Ficee con nuovi Studi sopra l’Androsace degli Antichi. 49. 1843. 4. Kitzing, F. T. Phycologia Generalis. 400. 1843. 5. Kiitzing, F. T. Species Algarum. 744. 1849. 6. Agardh, J. Species, Genera etOrdines Algarum. 2: 295. 1851. 7. Harvey, W. H. Nereis Boreali-Americana 1: 21; 2: 160, 173. 1553. = 8. Harvey, W. H. Notice of a Collection of Alga made on the Northwest Coast of North America, chiefly at Vancouver’s Island, by David Lyall, Esq., M.D., R.N., in the years 1859-61. Journ. Linn. Soc. Bot. 6: 172. 1862. 9g. Kiitzing, F. T. Tabule Phycologice 17: 24. pl. 837. 1867. to. Zanardini, G. Iconographia Phycologica Adriatica 2: p/. 78. 1860-76. 11. Agardh, J. Species, Genera et Ordines’Algarum. 3: 225. 1876. 190 MINNESOTA BOTANICAL STUDIES. 12. Bornet, E. Algues de Madagascar recolteés par M. Ch. Thie- baut. “Bull: Soc. Bot. de France: 92:16. 72 7, Za) ease 13. Hauck, F. Die Meeresalgen, in Rabenhorst’s Krypt. Fl. 2: T4b. | (1885: 14. Schmitz, F. Systematische Ubersicht der bisher bekannten Gattungen der Florideen. Flora 72: 435-456. 1889. 15. Okamura, R. Remarks on some alge from Hokkaido. Bot. Mag. Tokyo 5: 333-336. 1891. 16. Schmitz, F., and Hauptfleisch, P. Die natiirlichen Pflanzen- familien. JI. Teil. Abt.2. 517,519, 520, 525. stooge mi. EXTENSION OF PLANT RANGES IN THE UPPER MINNESOTA VALLEY. L. R. Mover. The following notes refer to plants that have been collected in the upper Minnesota valley since the publication of Professor MacMillan’s Metasperme of the Minnesota Valley. Duplicates have been deposited in the Herbarium at the State University. Thlaspi arvense L. This old world crucifer has become well established in the railroad yards at Montevideo and is spreading rapidly. Conringia orientalis (L.) Dumorr. This plant, first collected in wheat fields in Chippewa County in 1894, is spreading very rapidly and seems likely to become one of the worst ‘‘ mustards” with which the farmer has to con- tend. Sisymbrium altissimum L. This plant, too, is spreading very rapidly along the railroad tracks at Montevideo, and seems likely to become a very trouble- some weed. Peucedanum nudicaule (Pursn) Nutr. This plant is found to be quite plentiful in the upper Minne- sota valley on thin gravelly land near ledges of igneous rock. It is one of the earliest spring flowers. Potentilla hippiana Lrenum. This western Potentclla is occasionally found on ledges of gneiss rock near Montevideo. Helianthus annuus L. Among the recent arrivals at Montevideo, an immigrant from the West, is the annual sunflower. It is traveling along the railroads. 192 MINNESOTA BOTANICAL STUDIES. Grindelia squarrosa (PursH) DuNAL. This is another western plant that seems to be traveling east. It has but recently become well established in the railroad yards at Montevideo. Lactuca scariola L. Another Old World weed that has but recently arrived at Montevideo is this species. It is spreading with great rapidity. Senecio vulgaris L. Recently arrived at Montevideo, this plant is becoming com- mon as a weed in gardens and waste grounds. Poa pseudopratensis Scris. & Ryp. A Poa collected at Clara City, in Chippewa County, has been identified by Professor Lamson-Scribner as this species. Fraxinus lanceolata Borcx. A study of this species based on a large collection of ma- terial from the prairie portion of the State, some of which was sent east for comparison, makes it probable that all of our ash trees are referable to this species. /. Americana appears to be absent from western Minnesota. Cactus viviparus Nutt. This cactus is found quite abundantly on granite ledges in the ancient valley of the ‘*‘ Warren,” about two miles southeast of Ortonville. Its bright red flowers are strikingly beautiful. Astragalus flexuosus (Hoox.) Doue. This species is quite plentiful near the railroad yards at Or- tonville. SV. List OF HEPATICA: COLLECTED ALONG THE INTERNATIONAL BOUNDARY JBY J VERO ZINGER, 1897. ALEXANDER W. EVANS. . Aplozia autumnalis (DC.) Herc. F,N. . Bazzania trilobata (L.) S. F. Gray. F,R, U. . Blepharostoma trichophyllum (L.) Dumorr. R. . Cephalozia catenulata (Husen.) Spruce. F. . C. media Linps.. P. . Frullania Eboracensis Gotrscue. Ff. . Jungermannia barbata Scures. F, G, P. . J. quinquedentata Wes. P,R. . J. ventricosa Dicks. S. . Lejeunea serpyllifolia (Dicxs.) Lis. U. . Lepidozia reptans (L.) Dumort. P,R,S, U. . Plagiochila asplenioides (L.) Dumorr. N, P, U. . Porella platyphylla (L.) Linps. F, U. aitiidiamciiare Cl.) NEES = © oi. S..U) . Radula complanata (L.) Dumort. C, F, U. 16. Scapania glaucocephala (Tayu.) Aust. F. C = Camp IV., on the Prairie Portage, shore of Basswood Lake, near the rapids from Sucker Lake. OO OonNT ANN BW bd H =x Ss HS SS FS AS im ih &® N AH F = Fall Lake, near the foot of Kawasatchong Falls, seven miles north of Ely. G = between Gunflint Lake and Grand Portage. N = stream flowing from North Lake into Little Gunflint Lake. P = Grand Portage Island. R = Pipestone Rapids, on Basswood Lake. S = Safety Island. U = United States Peninsula. XVI. OBSERVATIONS ON CHLOROCHYTRIUM. E. M. FREEMAN. In 1850 Mettenius found numerous green cells in the thallus of Polyzdes lumbricalis which resembled closely what are now Classified as the Endospherez of the Protococcacee. He interpreted them, as spore-mother cells of the red seaweed upon which they were found. Thuret fourteen years later observed these same structures and interpreted them as parasitic zodspores which on germination produce the bushy thallus of Cladophora lanosa. Cohn in 1865 was able to confirm the observations of Mettenius and of Thuret, except as to the germ- ination of the endophyte into Cladophora lanosa. The condition of endophytism was considered at that time as indicative of parasitism. Hence new interest was aroused in the investigation of these lower forms when Rees and Schwen- dener at about the same time (1871) advanced independently the theory that the Co//ema type of lichen is to be derived from a discomycetous fungus, the mycelium of which has pene- trated the mucilage of a /Vostoc completely surrounding the latter. Reinke’s observations on /Vostoc in the stems of Gun- nera scabra and the work of Milde and Janczewski on /Vostocs in liverworts demonstrated the occurrence of Protococcus-like algal forms in the plant tissues of higher plants. Cohn in 1872 suggested that the presence of the Nostoc filaments in Gunnera and Anthoceros is perhaps to be explained as an acci- dental entry of the movable /Voséoc filaments into the tissues of the nurse plant, their continued growth in this new sheltered position and their subsequent imprisonment by the growth of the surrounding tissues of the nurse plant. In contrast to this form of endophytism Cohn describes the new genus Chloro- chytriuwm, which he considers to be a true parasite in certain species of Lemna. The zodspores, very numerous on the sur- face of the host, send out a germination tube between two epi- 195 196 MINNESOTA BOTANICAL STUDIES. dermal cells. The membrane of this tube becomes thickened by subsequent layers, the tube swells with the absorption of the chlorophyll and protoplasm and the intercellular endophyte results, with a cellulose button protruding from the point where the germinating tube entered. The endophyte then becomes pyriform and almost opaque on account of the density of the chlorophyll. By free cell formation large cells are formed in the endophyte and these finally break into a large number of zoospores which are expelled through the cellulose protuber- ance from the nurse-plant epidermis. The endophyte is, there- fore, an independent organism closely related to Wydrocytium (Charactum A. Br.) on the one hand and to Syuchytrium on the other. With the Eu-Synchytrium group its cell form and the formation of zoédspores by a preliminary division into seg- ments, corresponding to the zodsporangia of Syuxchytréum, agree, but it differs in the presence of chlorophyll and of a germination tube and in its intercellular position. Upon these observations Cohn founded the genus Chlorochytrium and de-. scribed it as follows: Planta endophytica viridis unicellaris, globosa ovoidea vel irregulariter curvata bi, tri, multiloba dense conferta plasmate Viridi, primum in segmenta majora diviso dein secedente in zoosporas immersas pyriformes virides processibus tubulosis ex- tus emissas. Chlorochytrium lemne upon which the genus is based is then described. Cohn pronounces Chlorochytrium a true parasite. That no deleterious effects upon the host are visible is paralleled in Per- onospora and Syuchytrium. In its intercellular position it re- sembles the Uredinez. Two years after Cohn’s observations were published Kny described a new species of Chlorochytrium endophytic upon Ceratophyllum demersum. It differs from Chlorochytrium lemne in size and in the absence of a cellulose button. In 1877 Wright established a third species of Chlorochy- trium, C’. cohni Wright. ‘The zodspores impinging on the fronds of several species of marine alge quickly assuming a figure-of-eight form, the lower sphere growing into the frond and rapidly assuming com- paratively large dimensions, the upper sphere remaining as a tube-like neck portion to the larger mass. On the cell arriving Freeman: OBSERVATIONS ON CHLOROCHYTRIUM. 197 at an adult stage, the whole of the green protoplasmic contents divides into a number of from 10-30 nearly circular zodspores, which escape through the neck-shaped portion. ‘‘ Living in the thallus of various species of Schzzonema, Polysiphonia, etc.; also on the Infusoria found at Howth.” Wright states further that there are two kinds of zodéspores, large and small, the latter being the more numerous. Szymanski in 1878 described oe knyanum apparently identical with the plant mentioned by Kny four years before as inhabiting the tissues of Ceratophyllum demersum. ‘This species was found on Lemna minor and possessed a cellulose button which did not protrude farther than twice the thickness of its outer wall above the epidermis of its nurse plant. Klebs published the results of his investigations on C. /emne in 1881. Chlorochytrium in the younger vegetative stages contains a light green chlorophyll-bearing protoplasm with iso- lated starch grains surrounded by a cell sap vacuole (see below, pyrenoids). Inthe later stages the grains increase in number, the mesh-work of green bands becomes smaller, the chlorophyll darker until almost opaque and the protoplasm finally becomes coarsely granular. After a resting period of a week or more the zoéspores are formed by successive bipartitions of the cell contents, at first by perpendicular, later by radially disposed walls. ; The number of divisions is not known. Liberation of the zoOspores is accomplished by absorption of water resulting in the splitting of the Chlorochytrium wall and of the superposed Lemna tissues. The conjugation of the biciliate zoo6gametes into larger quadriciliate zodzygotes was observed, a fact which may throw light on the macro- and microzoéspores of Wright’s species. Klebs observes that Cohn’s account of the liberation of the zoOspores is without observational foundation and doubts its ac- curacy. He also calls in question the appropriateness of placing Wright’s species in the genus Chlorochytrium and further sug- gests the probability that C. knyanum is but the asexual form of C. /emne since no copulation had been observed between the zoéspores. C. pallidum Kiebs and many similar forms are probably mere ‘* place varieties” of C. knyanum. Klebs points out with much truth that no proof has been given of the much averred parasitism of Chlorochytrium by Cohn 198 MINNESOTA BOTANICAL STUDIES. and other previous investigators. On the other hand C&loro- chytrium has well developed chlorophyll and lives near the sur- face where abundant light is available. The requisite inorganic matter may gain access to the cells by the constant or at least periodical submersion in water. Chlorochytrium lemne penetrates dead as well as living leaves and culture methods demonstrate an entire lack of de- pendence of the endophyte upon a host plant. In many endo- phytes zodspores can be developed on culture slides for months. No proof has as yet been adduced for any injury of the host beyond the results of the mechanical pressure exerted. The explanation then of endophytism is to be found not in parasitism but in the mechanical protection of position, which the inter- cellular spaces of the host offered; hence the appropriateness of the term ‘‘ Raum Parasiten.” It is, of course, possible for parasitism to develop from such a condition and this develop- ment seems to be in evident progress in such a nearly related form as Phyllobium dimorphum and also perhaps in JVostoc lichenoides. In the systematic relations of Chlorochytrium and the nearly related genera, Klebs briefly points out the inter- mediate position of Chlorochytrium and Endosphera, between the Protococcacee and the Chytridez, the-isolated position of Scotinosphera and the, probable affinities of PAyMobtum on the one hand with Chlorochytrtum and on the other with Bo- trydium. « Schaarschmidt, 1881, found zodspores of Chlorochytrium in a Desmid culture in which the zoédspores subsequently devel- oped, confirming Kleb’s view on the parasitism of the endo- phyte. In 1883 Kjellman described the following species: Chlorochytrium tnclusum Wjellman.—‘‘In the vegetative stage spherical or subspherical, entirely included within the nurse plant, with the formation of the zoédspores becoming slightly elongated, short-conical, flask-shaped, ovoid or ellip- soidal, finally bare at the pointed apex, which penetrates the cor- tical layer of the nurse plant and emitting the zoéspores through an ostiole which has been formed.” This species is endophytic upon Sarcophyllis arctica, mostly near the surface but some- times in the middle of the host. It averages 80-100 mic.,-has yellowish-green contents and a cell-wall which is thin and of equal thickness. The chromatophore is thin and is spread along the wall. With the elongation at the formation of zoospores Freeman: OBSERVATIONS ON CHLOROCHYTRIUM. 199 the membrane thickens towards the outer surface especially, and a cone-shaped growth of cellulose is formed. The proto- plasm then takes on a more intense yellow green and divides into numerous zodspores. Numerous bulgings of the plant cell are produced probably by the growth of the surrounding tissues of the host. An ostiole is formed at the apex of the cellulose out-growth by which the zodspores escape. Those cells found in the center of the host may attain as great a diameter as 275 mic. The even thickness of the wall of these cells suggests that they may be resting stages. Kjellman refers this plant to the genus Chlorochytrium, but hesitatingly on account of his lack of knowledge about the further development. He found C. znclusum in all of the Sarcophyli’s edulis material which he has examined. ‘Zodspores are most abundantly produced in winter, but are also found in summer. The endophyte occurs in greatest abundance and most strongly developed in Sarco- phylts arctica. Its range is apparently coextensive with that of Sarcophyllis arctica; 7. e., throughout the arctic region, except in the North Atlantic, most abundant in the eastern part of the Siberian sea. Three more new species were described by Schroeter in 1883. Chlorochytrium rubyum with red contents and occupying the air spaces of the leaves and stems of Peplis portula and of AZentha aquatica; Chlorochytrium viride, in the leaves of Rumex obtus- tfolius; and C’. Jetum, a spherical cell with yellow contents which become green in water, found in Lychnis flos-cucul?. The investigations of Moore on Chlorochytrium lemne pub- lished in the following year brought to light no new facts of importance. Moore held that the nearest affinities of Chlorochy- trzum lie with Protomyces. In 1887 Hieronymus described C. archertanum in punctured cells of Sphagnum leaves, and characterized by a greatly devel- oped cellulose button. Zodspores are formed regularly but do not copulate. P. Hariot, 1889, collected C. czclusum on species of Gzgartiena at Cape Horn, supposedly identical with Kjell- man’s C. znclusum. In his Conspectus of Endophytic Alga, Mobius, 1891, men- tions in addition to those species enumerated above, C. dermato- colax which was described by Reinke and found on species of Polysiphonia and Sphacelarta, and in his opinion should be classified under the genus C/lorocystzs, since it is marine, is in- tracellular and emits zodspores singly. 200 MINNESOTA BOTANICAL STUDIES. C. schmitzit was described (1893) by Rosenvinge from Greenland material on Cruorza arctica. The cell is without a cellulose button or papilla, is more or less attenuate at the base and has a single chromatophore with sometimes two pyre- noids. Zodspores were not observed. Collections, Methods, etc.—Upon the Constantinea material which was collected by Miss J. E. Tilden at different points along Puget sound and was distributed as Constantinea sitch- ensts Post. and Rupr., were found a large number of endo- phytic unicellular, chlorophyllaceous alge, referred to the genus Chlorochytrium of Cohn. The endophyte was found in abundance on all of the Constant:nea collected. One collection was made in August, 1897, near Seattle, and in the summer of 1898 (May 25-July 2), five were made at as many points farther up the Sound. Most of the material used in the following in- vestigations was fixed and preserved in a 2 per cent. sea-water solution of formalin. The green color of the endophyte was well preserved. ‘The lower ends of the cells have in very many cases an irregular outline which may possibly be in part due to shrinkage but is caused for the most part by pressure of the surrounding Constantinea cells. In all of this endophyte formalin material and in the dried material as well, though not so markedly, brown bodies were found jutting out between the chromatophore and the cell wall and assuming various forms (see below). These bodies were undoubtedly due to a chloro- phyllan reaction, the hypochlorin reaction of Pringsheim. The formalin solution when tested was found to give a slightly acid reaction. Sections of Constantinea cut freehand between elder pith furnished most of the material for study. ‘They were mounted either in the two per cent. formalin sea-water solution, in glyc- erine, or in glycerine jelly. Material carried through the usual paraffin method stained and mounted in Canada balsam has also furnished useful sections. The abundance of the endophyte makes it easy to get favorable surface and sectional views of it. General Habit and Structure.—The endophytes on Constan- tinea sitchensis occur on both the upper and lower surfaces of the fronds. I have been unable to find any on the stipe. They are most abundant on the older fronds and especially towards the peripheral portion. From some young fronds they are almost altogether wanting. They occur in greatest numbers Freeman: OBSERVATIONS ON CHLOROCHYTRIUM. 201 on the lower surface while rather few are found on the upper side. The following figures are taken from a medium-sized frond in the peripheral region: on the under side 140-160 (sometimes as many as 230) in one square mm.; on the upper surface 60-65 in the same area. Many areas of 4 square mm. on the upper surface contained no endophytes. The endophytic cells are found almost exclusively in the tis- sues just beneath the pseudoepidermis of the nurse plant, with the slightly pointed end just at or just below the surface. They occur in rare cases in the central part of the frond completely enclosed. The pointed ead protrudes from the tissues of the nurse plant in but few cases and then not moxz than for a distance equal to one-half the thickness of its outer wall. The cells not infrequently occur between the paraphyses of the nemathecia of Constantinea where they usually penetrate to but not into the tissues beneath. The predominant form assumed by the endophyte is pear- shaped with the smaller end directed toward the surface of the nurse plant. The cells are often ovoidal and even ellipsoidal. In the central portion they assume a spheroidal form. In the paraphyses they become elongated or assume a figure-of-eight form similar to that described by Wright for Chlorochytrium cohniz. ‘The inner ends of the cells are marked more or less by the bulgings undoubtedly caused by the pressure of the adjacent cells of the nurse plant. The cells average 85-115 mic. in length and 4o-85 mic. in breadth but often attain 143 x 100 mic. ‘The wall in some cases is 28 mic. thick at the outer surface and 8 mic. thick around the remainder of the cell, but usually is less than one-half of these dimensions. The lamellation of the cell wall can clearly be seen in many sections (especially glycerine mounts) and is due probably to the apposition of successive layers of cellulose. Chlorophyll occurs in the form of a single yellowish-green plate in which are included a large number of fine refringent gran- ules. This chromatophore extends around the entire wall of the cell and contains a varying number of very conspicuous pyrenoids, which are flattened spherical in form, 5—1I mic. in diameter, and jut out into the cavity of the cell. As many as thirty-nine have been found in a single cell and at least one pyrenoid can be seen soon after the cell begins to penetrate the tissues of the nurse plant. The pyrenoids show a clear central 202 MINNESOTA BOTANICAL STUDIES. portion probably proteid. In sections stained heavily with aniline blue the central portion appears blue. Around the clear center are arranged 5-10 plates of starch which stain brown with both a KI and an alcohol solution of iodine. By careful washing of material stained in an alcoholic solution of iodine and with the aid of a +; oil immersion lens a distinct violet tinge is discernible in the plates. The protoplasmic contents of the cell are usually most dense in the pointed part. Between the chromatophore and the cell wall are found numerous rust-brown to black (in a few cases copper-colored) bodies of different form and size. In some places they occur in diffuse patches the limits of which are often indefinable, in others as five-pointed rosettes. Again they may appear filiform, partially and usually irregularly coiled or forming a delicate and loose network. I have interpreted these bodies as products of the action of the dilutely acid formalin solution and as identical with the hypochlorin of Pringsheim. His plates agree closely with much of the material at hand. In accordance with - Pringsheim’s account of the chemical reactions of hypochlorin, these brown bodies are wanting in those sections which have been carried through the alcohols in the method for paraffin embedding. A large amount of material has been examined but in no case has even a trace of the production of zodspores or gametes been found. The stages in the penetration of the nurse plant, consisting in the elongation of the at first spheroidal cell, the subsequent withdrawal of the protoplasm into the inner end and the increase in size of the latter to form the mature pyriform cell, have been observed, but nothing to indicate the formation of zoospores. Conclustons.—It is therefore upon the basis of vegetative characters that the endophyte described above is provisionally placed in the genus Chlorochytrium under C. inclusum Kjell- man. Upon examination of the Chlorochytrium inclusum found upon Dilsea (Sarcophylis) distributed in Phycotheca Boreah- Americana (Fasc. XI., no. 514) this is seen to possess a thicker cell wall than the material on Constantinea sttchensis, is almost spheroidal, larger, has denser dark green contents, con- tains no pyrenoids (or very inconspicuous, if present at all) and resembles a resting stage. The time of collection, December, moreover, strengthens this last supposition. The material under Frreeman: OBSERVATIONS ON CHLOROCHYTRIUM. 203 observation was collected, on the other hand, in summer, May to July. The light yellow-green color, the absence of repro- ductive bodies and the abundance of small cells point strongly, I think, to the conclusion that this endophyte is but a young stage of Chlorochytrium tnclusum WKjellman. A similar dif- ference in the vegetative and resting stages of C. /emn@ are recorded in Klebs’ observations (f/. 39, f. 2 and g). The form and habit of the endophyte upon Constantinea accord best with Kjellman‘s description of C. zzclusum; no mention, however, is made by Kjellman of pyrenoids, which are so conspicuous in the Constantinea material. It is suggested by De-Toni and also by Miss Whitting that Kjellman’s species may possibly belong to Ch/lorocystzs, a genus established by Reinhard in 1885. Chlorocysts is described with but one pyrenoid while the endophyte on Constantinea contains many. C. schmitziz, described by Rosenvinge, is but imperfectly known. I have seen in the sections studied a number of cases where the endophyte in Constant:nea has assumed approxi- mately the same irregular or obovoidal form shown in the figures of Rosenvinge. The only remaining marine species of Chlorochytrium, C. dermatocolax, lives in the outer mem- brane of Polyszphonza and, according to Mébius, belongs prob- ably to Chlorocystis. 4 The great similarity in vegetative structure to that described by Kjellman for C. znclusum would indicate that the Cons¢tan- tinea endophyte described above is a midsummer stage of C. tnclusum and I would provisionally place it in that species awaiting further information on the life history and develop- ment. BIBLIOGRAPHY. Cohn, F. Ueber parasitische Algen. Remarks before the ‘*‘ Wand- erversammlung der Schlesischen Gesellschaft fiir vaterlandischer Cul- tur. Botanische Section. 1872. Beit. zur Biologie der Pflanzen. Bey Os Pl..2:- 1975, Kny, L. Ueber eine griine parasitische Alge. Sitz. Gesellch. na- tiirf. Freunde zu Berlin. 1874. Wright, P. Ona new species of parasitic green alga belonging to the genus Chlorochytrium of Cohn. Trans. Roy. Irish Acad. 26: 1877. Szymanski, F. Ueber einige parasitische Algen. Inaug. dissert- der Univ. Breslau. 1878. Kirchner, O. Die Algen Schlesiens. Breslau. 1878. Klebs, G. Beitrage zur Kenntniss niederer Algenformen Bot. Centralb. 39: 16-21. fl. 37, g. 1881. 204 MINNESOTA BOTANICAL STUDIES. Hariot, P. Algues recuillies par la Mission scientifique du Cap Horn. Paris. 1882-83. Reinke, J. Algenflora der westliche Ostsee deutschen Antheils. Sep.- Abdr. VI. Ber. Commiss. Untersuch. deutschen Meere, in Kiel. 1889. De-Toni, J. B. Sylloge Algarum. 1: 635-637. 1889. Mébius, M. Conspectus Algarum endophytarum. Notarisia. 6: 1282. 1891. Rosenvinge, M. L. Kolderup. Les Algues Marines du Groenland. Ann. Sci. Nat. Bot. 19: 169. f. 56. 1894. Whitting, F. G. On Chlorocystis sarcophyc?. Phyc. Mem. Pt. 2. 41-45. 1893. Wille, N. Die natiirlichen Pflanzenfamilien. I. Abt. 2. 66. f,. 42. “1897; Schaarschmidt, G. A Chlorochytrium Erdelylen. Magyar N6- venyt. Lapok. 5: 37. 8181. Kjellman, F. R. Algz of the Arctic sea. 320. pl. 37. f. 8-17. 1883. Schréter, J. Neue Beitrage zur Algenkunde Schlesiens. 61. Jahresb. Schles. Gesellch. Cult. 178-189. 1883. Moore, S. Le M. Remarks on some endophytic Alge. Journ. of Bot. 22: 136-138. 1884. Lagerheim, G. On Chlorochytrium cohnit Wright och ders fér- hallande till narstande arter. Sv. V. A. Oefvers. 7: 91-97. pi. 2. (Bot. Jahres: 12>: 271." 18386;) Hieronymus, G. Ueber einige Algen des Riesengebirges. Jahresb. Schles. Gesellch. Vaterl. Cultur. 293-297. 1887. Tilden, J. E. American Algae. Cent. III. 1898. DEscRIPTION OF PLATE XIX. 1. Diagram of a cross section of a frond of Constantinea sttchen- szs showing the endophytic Chlorochytrium cells on the upper (less numerous) and lower surface. e. endophyte. f. pseudoepidermis. z. intermediate area. c. central area of the frond. 2. Surface view of the frond of Constantimea (lower surface) showing the pseudoepidermis (#) with cavities (e) through which the endophytes have penetrated. x 57. 3,4, 5,6. Stages in the penetration of the nurse plant. s. pyre- noid. x 288. 7. Chlorochytrium cell witha not infrequent form. Shows fila- mentous form of hypochlorin. x 288. 8. Cell showing elongated form of endophyte found amongst the paraphyses of the nurse plant. x 288. 9. Large Chlorochytrium cell of typical form. x 288. 10. Detail showing the position of a Chlorochytréum cell in the cross section of the Constantinea frond. Lettersasinno.1. x 288. 11. The outer end of a Chlorochytrium cell showing the lamella- tion of the cell wall. x 288. w VOL. II. MINNESOTA BOTANICAL STUDIES. PART III. PLATE XIX. THE HELIOTYPE PRINTING CO., BOSTON. XVII. OBSERVATIONS ON RHODYMENIA. FrepeEeric K. BuTTERs. History and Literature. —The genus /thodymenia was founded by Greville in 1830. As originally constituted it con- tained species which have since been referred to Graczlarza, Gigartina, Kallymenta, Calliblepharis and other genera. Agardh (3), p. 376, states that in his Mediterranean Alge (1) he transferred many species to the genera Gracilaria, Gigan- tina and / allymenta, but united Calliblepharts with /thody- menta under the latter name. In 1849 Kiitzing in his Species Algarum, p. 778, united the species of the genera /-hodymenza and Graczlarva together with some other species under. the generic name Sfherococcus. Agardh (2), p. 15, (3) p- 375, revised the genus /thodymenza and gave it substantially its present limits and generic characters. The generic description as given by De-Toni and Levi, p. Ig, is as follows: Frond plane, membranaceous, dichotomous or pal- mate, proliferate from the margin or surface, composed of two layers ; interior cells oblong, cortical minute, rounded ; cystocarps scattered throughout the frond, each within an hemispherical pericarp opening by a carpostome, composed of cells, the outer radiating the inner concentric ; cystocarp with a simple rounded or somewhat lobed nucleus; nucleus naked within the pericarp, on a basal placenta with paniculate-branched placental fila- ments sustaining the lobes; young fertile lobes disposed radially composed of articulated filaments, older obconico-rotund, bear- ing numerous protospores; tetraspores often collected into sori, cruciately divided; antheridia produced in superficial sori com- posed of minute hyaline cells in a single vertical series. Greville spelled the name of this genus APkodomenia. Mon- tagne, p. 44, in 1839, employed this spelling in a list of Bra- silian cryptogams, but in a footnote states that, in conform- ity to its etymology it should be spelled Ahodymenia. J. 205 206 MINNESOTA BOTANICAL STUDIES. Agardh (3) adopted the latter spelling and it has since been in general use, although Athodomenza and, according to Ardis- sone, A’hodhymenia and Rodhymenza also have been occasion- ally employed. Fhodymenia pertusa was first described and figured by Pos- tels and Ruprecht, p. 20, p/. 36, under the name Porphyra pertusa. Kiitzing, p. 693, describes it under this name. He introduces a question, however, in respect to its generic de- termination. J. G. Agardh (3), p. 376, points out that the spe- cies should be classed as a Rhodymenza and not as Porphyra. He describes it as Ahodymenia pertusa (Post. and Rupr.) and places it, together with Ahodymenia palmata (Linn. Sp. 2: 1630) and PRhodymenia peruviana (J. Ag. Mscr.) in the sec- tion Palmate characterized by ‘‘ tetraspores occurring throu gh- out the surface of the frond, scattered or collected into sori.” In his Epicrisis (Agardh, J. G. (4), p. 379), he assigns it the same position. Agardh (3) gives the habitat of -Rhodymenia pertusa as ‘in the Arctic Ocean near Kamtschatka (Mus. Petropolitani!) ; and near Greenland (Wormskjéld!).” Kjellman reports it it from the northwest coast of Spitzbergen and the west coast of Greenland. It was first reported from the northwest coast of America by Harvey (1) who found it among the algae collected by Captain Wilkes’ exploring expedition. Harvey gives its locality as the Straits of St. Juan de Fuca. He compared Wilkes’ specimens with an authentic specimen furnished him by Dr. Ruprecht and found them identical in species, though Wilkes’ specimens were considerably larger than Ruprechts’s. Harvey, p. 171, also reports the species as collected by Dr. Lyall in 1859-61, his specimens being ‘‘ cast ashore on Point Roberts, and on rocks at low water, Fuca Strait.” Cystocarps were present on both sets of specimens mentioned by Harvey. In commenting on the specific name, Harvey (1) p. 148, states that to him the perforations of the thallus appear to be due to casualties. In 1893 Carruthers, p. 80, examined one of the specimens mentioned by Harvey as collected by Dr. Lyall. He found that in that specimen the cystocarps occur all over the much perforated segments; the majority are very young and project but little from the surface of the thallus. The structure of the thallus is that typical of the genus. He says ‘‘ The cystocarp Butters: OBSERVATIONS ON RHODYMENIA. 207 projects on one side of the thallus and possesses a fairly thick fruit wall of five or six cells in thickness arranged irregularly inwards. At the bottom of the fruit cavity is to be found the placenta formed of numerous small cells closely packed to- gether and lying on the medullary layer of large cells. ‘¢ From the placenta the gonimoblast of more or less irregular shape, is borne on an elongated style cell and spreads upward into the empty fruit cavity. ‘*This gonimoblast is composed of numerous pear-shaped lobes which lie close together. The spores of these lobes be- come ripe nearly at the same time. ‘¢ The ostiole is generally quite in the center of the projecting fruit wall, and is similar to the ostioles of the kindred species.” Collection and Preservation.—All the material at hand was collected by Miss Josephine E. Tilden at Port Orchard, Kitsap County, Washington, on August 2, 1897. The specimens were dredged in water 4-6 fathoms deep. A small portion of the _ material was killed and preserved in 80 per cent. alcohol. The larger part of the material was dried. By soaking, this dried material so far regained its original form that the anatomy could be well studied although the cell contents were largely disor- ganized. All the observations upon the stipe and the prolifera- tions were made upon this dried material; observations upon other points were made mainly upon the alcoholic material. Methods.—The dried material was soaked in water until it regained its natural consistency. Various methods were em- ployed in cutting the tissues. Much of the material was cut upon the Osterhout freezing chamber. (Osterhout, W. J. V., p- 195.) The alcoholic material was first passed into water —preferably through about three intermediate grades of dilute alcohol. When the alcohol was completely removed the mate- rial was in some cases infiltrated with gelatine solution and then mounted in a drop of gum arabic solution upon the freezing chamber; in other cases it was embedded directly in the gum arabic. On account of the firm nature of most of the tissues, this method of mounting directly in gum arabic proved quite as successful as that in which the tissues were first infiltrated with gelatine. The sections, as soon as they were removed from the knife, were passed into 20 per cent. glycerine. Those which were to be stained were transferred from this glycerine to the staining 208 MINNESOTA BOTANICAL STUDIES. solution, thence, after washing, back to the glycerine solution. The glycerine solution was allowed to concentrate by evapora- tion, and the sections were thence mounted into glycerine jelly. Portions of the frond were also embedded in pith and cut free- hand with the razor. Some of the material was embedded in parafline, according to the usual methods and cut on the microtome. The sections thus obtained proved, in some instances, very successful, espe- cially in the case of such firm tissues as the stipe and the vege- tative portions of the lamina. Several staining fluids were used. Both section-staining and staining zz toto were employed. The former method proved most successful. The following stains were found useful. Aniline blue: Sections were placed for 5-10 minutes in a saturated solution of aniline blue (spiritlés) in 50 per cent. alco- hol. The walls were stained a deep blue, gelatinous structures and cell contents a light blue. This proved the most useful stain for clearly defining anatomical details. Hoffman’s violet: Sections were stained with a saturated so- lution of Hoffman’s violet in concentrated sulphuric acid. As soon as the section had taken a brown stain they were placed in water and the acid washed out. ‘The protoplasm took a blue stain, the walls were unstained. This method proved useful in staining the protoplasmic connections between the cells. Fuchsin: Dilute alcoholic solution of fuchsin stained the walls light red, the protoplasmic contents a deeper red. Iodine in potassium iodide: This stained the floridian starch a dark yellow-brown, the other cell contents a light yellow- brown. It proved very useful in staining the protoplasmic con- nections between the cells, especially in the gonimoblast fila- ments. Gross anatomy (Fig. 1).—The plant consists of a broad, flat, membranaceous, more or less subdivided, blood-red lamina borne upon a short stipe which is sensibly continuous with the lamina. The stipe is expanded below into a small holdfast. The lamina is sometimes nearly entire in outline, ovate, or broadly lanceo- late, sometimes very deeply lobed, or divided almost to the base into 2-4 lobes which may be widely divergent. The lamina is 17-40 cm. long and 8-17 cm. wide. The two faces of the frond are in all respects similar. The lamina is perforated by numerous holes ranging from .5 mm. to 10 cm. in length and Butters: OBSERVATIONS ON RHODYMENIA. 209 from .5 mm. to 2 cm. in width. The smallest are almost circu- lar in outline. Those somewhat larger (up to about 5 mm. in diameter) are usually somewhat oval, being elongated in the direction of the long axis of the frond. The larger perforations are of irregular elongated outline. Sometimes these perfora- tions are exceedingly numerous, as many as four or five per square cm. being not uncommon. In some specimens numerous proliferations are borne on the faces and edges of the lamina and also on the stipe. These are cylindrical bodies, about 1 mm. in diameter, seldom more than 25 mm. long, often flattened somewhat towards the apex and often branched once or twice or deeply lobed at the apex. In two specimens the upper portions of the laminz present a peculiar mottled appearance which was at first supposed to be due to the presence of tetraspores. It appears that this is not the cause of the phenomenon noted. This subject will be dis- cussed further under the description of the cystocarp. The entire stipe was present in only one of the specimens at hand. In this it was 3.5 cm. long, about I mm. in greatest diameter, flattened somewhat, parallel to the flat surface of the lamina, passing insensibly into the lamina above, and expand- ing abruptly below to form the holdfast, which is a small, thin, irregular disk, about 5 mm. in diameter. All of the specimens have cystocarps scattered irregularly over the entire surface of both sides of the lamina. These are bodies I-1.5 mm. in diameter, nearly hemispherical, or pro- truding slightly at the apex, and are extremely numerous, as many as fifteen being often found on one square cm. of the lamina. Minute anatomy: Lamtna.—The lamina consists of pseudo- parenchymatous tissue of which two principal areas may be dis- tinguished in the cross-section (a), a central layer of large- celled; and (b) a cortical layer of small-celled tissue (Fig. 2). (a) The cells of the central area are large, generally some- what flattened parallel to the surface of the frond, isodiametrical in tangential section (Fig. 3). The cells vary greatly in size, the average being 73.5 x 105 mic., while cells occur as small as 6x14 mic., and as large as 100 x 200 mic. The largest cells are situated near the central portion of this area, and from them the size of the cells decreases quite regularly towards the more superficial portions. The more superficial cells of this area 210 MINNESOTA BOTANICAL STUDIES. differ considerably in other respects also, from the central cells. The outer cells are more flattened than the central ones, they are more densely protoplasmic and are filled with grains of floridian starch, while but little starch occurs in the central cells. In this more superficial portion of the central area the proto- plasmic connections between the cells of the filaments of which the tissue is composed can be easily made out. They are plainly visible in both stained and unstained preparations, and in both tangential and cross sections. In accordance with the less dense protoplasmic contents of the central cells, the con- necting strands are less evident among them, but when the walls of this area are stained, numerous pits are shown penetrating the walls of the central cells (Fig. 3). These pits appear to be of irregular distribution and often more than one are to be ob- served between the same pair of cells. They can be best ob- served in a tangential section. (6) The cortical area consists of small cells almost spherical or with the longest diameter perpendicular to the surface of the frond, of quite uniform size (averaging 5.7 x 8.5 mic.), arranged in 1-3 layers, either in filaments perpendicular to the surface of the frond or somewhat irregularly. In a surface view of the lamina they appear entirely irregular in arrangement (Fig. 4). The cells are densely protoplasmic and contain chromatophores. Stipe.—The general structure of the stipe (Fig. 5) is similar to that of the lamina, but there are numerous special modifications of the several areas. The cells of the central area are elongated somewhat in the direction of the axis of the frond and are some- what compressed parallel to the compression of the stipe. They are of more uniform size than were those of the lamina, and they were otherwise more nearly uniform than those of the lamina. Their average size is 57 x86x143 mic. The cortical ° area of the stipe is much thicker than that of thelamina. Itis 3-8 cells deep. The cells are larger than the corresponding cells of the lamina (average 14 x 23 mic.) and are conspicuously ar- ranged in filaments running perpendicular to the surface of the stipe. Proliferations.—The structure of the proliferation is similar to that of the main frond. The central cells are somewhat elongated in the direction of the axis, and, in general, are more numerous and smaller than the corresponding ones of the main frond. All the cells except those of the cortical area contain considerable floridian starch. Butters: OBSERVATIONS ON RHODYMENIA. Ae Reproductive tract.—Cystocarp (Fig. 7-12). The cystocarp projects on one side of the thallus. The pericarp is composed of thickened cortical tissue, which, in the mature cystocarp, is 8-40 cells thick. The outer walls of the pericarp are small and resemble those of the cortex of the vegetative part of the frond. The inner cells are large (14 x 29 mic.), flattened paral- lel to the surface of the cystocarp and show numerous irregu- lar protoplasmic connections. The cystocarp opens by a carpo- stome situated at the apex of the pericarp. The sporogenous tissue is in the form of an irregularly lobed mass, borne on a basal placenta, and partially filling the cavity of the cystocarp. The space between the spore mass and the pericarp is filled with gelatine. The placenta is a mass of small- celled tissue containing numerous intercellular spaces, which rests upon the large-celled central tissue of the lamina. The gonimoblast filaments (Fig. 9-11) branch repeatedly. They consist of irregular elongated or rounded, often club-shaped cells, with dense, finely granular protoplasm and very trans- parent walls. They contain no starch. They are connected in filaments by very broad protoplasmic connections surrounded midway by a callous-like ring. The upper cells of the fila- ments are smaller and more rounded than the lower. The structure of the filaments was best shown in preparations made by pressing out the contents of a mature cystocarp upon a mi- croscopic slide, staining lightly with iodine in potassium iodide, and then pressing out with a cover glass. The spores themselves are irregular, ovoid, thin-walled cells, densely packed with floridian starch. Their average size is 21.5 x 34.5 mic., but they vary considerably inthis respect. The number of spores produced in each cystocarp is very great, 20,000-—30,- ooo being notuncommon. ‘The lowest lobes of the sporogenous mass appear to be in all cases sterile. They form small masses of compact tissue consisting for the greater part of cells having about one-half the diameter of mature spores and containing but little starch. A few long cells like those of the goni- moblast filaments also occur in this region. In many cystocarps branched filaments of cells rise from the vegetative tissue at the base of the cystocarp. The cells of these filaments (Fig. 8) exhibit peculiar lateral outgrowths which appear to fuse with the adjacent cells or with similar outgrowths from them, forming a peculiar loose, irregularly PAW MINNESOTA BOTANICAL STUDIES. connected tissue similar to that which forms the inner portion of the cystocarp wall. These filaments are not present in all cys- tocarps. They appear to result from the tearing of the sub- cortical tissue in the formation of the cavity of the cystocarp. In the young cystocarp (Fig. 12) the thickened wall is al- ready present and shows its permanent division into two layers. The cells of the outer layer are arranged in filaments perpen- dicular to the surface, those of the subcortical layer are ar- ranged in oblique rows converging towards the apex of the pericarp. The placental area and spore mass are represented by a few connected cells with very dense contents. The cavity of the cystocarp is, at this stage, comparatively small. The carpostome is already developed even in very young cystocarps. It appears to be formed by the tearing apart of the cells to- gether with the destruction of some of the cells. As was men- tioned in the account of the gross anatomy of the lamina, in two specimens the apical portions of the frond present a pecu- liar mottled appearance. Cross sections of these areas show that in places there are slight protuberances from the surface of the thallus associated with an unusual development of cortical and, in some cases, also of the subcortical cells. The cortical cells are rather narrower than elsewhere, and more elongated perpendicularly to the surface of the frond. The cortical layer is also a greater number of cells deep than elsewhere, and in some cases there is also increase in the number of the smaller central cells situated immediately beneath the cortex.. These areas in some cases involve only a few cells, in others they are .5 mm. in diameter. The structure of the larger protuberances agrees essentially with that of the young cystocarp described above, except that in no cases could any cavity be discovered in them. ‘They appear to me to be very young stages of cysto- carp development, but in no case could positive evidence of their nature be discovered. No indubitable cases of tetraspore formation were seen, but in some cross sections taken through the upper part of the lamina, some of the cortical cells are peculiarly divided pro- ducing somewhat the appearance of tetraspore formation (Fig. 6). Except for their peculiar arrangement, these cells appear in all respects similar to the ordinary cortical cells. Whether they are tetraspores or not, could not be determined. Their method of division is cruciate or somewhat irregular approach- ing the tetrahedral arrangement. Butters: OBSERVATIONS ON RHODYMENIA. Die BIBLIOGRAPHY. Greville, R. K. Alg. 1830. Montagne, C. Crypt. Brasil. in Ann. Sci. Nat. Bot. II. 12: 42- 55. 1839. Postels A. and Ruprecht, F. J. Illust. Algar. 20. A/. 36. 1840. Kitzing, F. T. Spec. Algar. 693, 778-784. 1849. (1) Agardh, J.G. Alg. Med. et Adriat. 1842. (2) Agardh, J.G. Alg. Liebm. (3) Agardh, J. G. Spec. Gen. et Ord. Algar. 2: 374-383. 1852. (4) Agardh, J. G. Epicr. syst. Florid. 324, 329. 1876. iagivey, VV. Fi. Ner. Bor.-Am. 2: 147, 148. 1852, Harvey, W. H. Coll. Alge Northwest Coast of North America. in Journ. Linn. Soc. Bot. 6: 157-177. 1862. Rjcuman, Bok. Alp. Arctic sea. 150, 157. 1683. Ardissone, F., Phiyc. Med. 1: 212-215. 1883. De-Toni, J. B. and Levi, D. Schem. Gen. Florid. Illustr. Nota- risia. 3: appendix I-XXV._ 1888. Carruthers, J. B. On the cystocarps of species of Callophyllis and Rhodymenia. Jour. Linn. Soc. Bot. 29: 77-86. 1893. Osterhout, W. J. V.) “A simple freezine device. Bot. Gaz. 21: Ig5-201. 1596. DESCRIPTION OF PLATE XX. 1. Mature plant showing perforations, proliferations and cysto- carps. xd. 2. Cross section of frond. Med. medullary area, cor. cortical Bheds XLT. 3. Longitudinal section of central area of frond, showing pits in the walls. x 132. 5. Surface view of frond, showing irregular arrangement of surface cells. Stained with fuchsin. x 335. 5. Cross section of stipe. Drawn with camera lucida. x 4o. 6. Vetraspores.? “x 335. 7. Cross section of mature cystocarp (not cut through carpostome) showing placentation and general structure of the spore mass. The upper part of the spore mass is somewhat scattered. Only the spores have their contents filled in. x39. 8. Peculiarly branched and interwoven filaments from the base of a cystocarp. xX 237. 9g. Gonimoblast filaments stained one minute in iodine in potassium iodide. x 237. 10, 11. Gonimoblast filaments, contents not drawn. x 237. 12. Young cystocarp showing carpostome and early stage in the development of spore mass. Contents have been omitted except from sporogenous cells. Drawn with Camera lucida. x 230. Rae ie es r : ae velit 4.) 1) ‘ Le Ree a pe, ds : } } > 4 <= ar ante | A } , PPE ane 7 ; , ; bt . 4 oe a be ait 7 ia of fyi te ‘ Dae tae reve ee iy ane : ate nn % - A ee Ik Pw ay afree ‘ a4 Sy } ’ a , (| Cyaan akan. = ee a ee wh as LMA Aj ae . ~ ; eo ; : ME ; 5 i ; ‘ 3 i ' P 53 ‘ Y & ( i] ' = ANT Arie usu? { a eT F bey 4 : AS - ; = - . ' Fi 4 4 Laer Atay Jou4 ~: , i 2 roe hey é Pore Seleeer. pene it . Rey A on oly Ate TY OP ALDER a 1k . nihey r nia t ; he - io ny : i , MINNESOTA VOL Il. 0 | of 38 \ ho AD } Q 00 id ° y 00 0g sal | old! TK) f) FR IN / eo *0 Y, () o id \1 i THE HELIOTYP! TANICAL STUDIES. XX. i LK BE ‘TING CO., BOSTON. i PART III. XVIII. CONTRIBUTIONS TO A KNOWLEDGE OF THE LICHENS OF MINNESOTA.—IV. LICHENS OF THE LAKE SUPERIOR REGION. BruceE FInx. CONSIDERATIONS OF DISTRIBUTION AND HABITAT. The area treated in this paper includes essentially the counties of Cook and Lake, comprising about 5,000 square miles of land. It lies to the northwest of lake Superior, bordering on the lake for about 150 miles and on the province of Ontario, Canada, for about 125 miles, thus forming the extreme northeastern portion of the State of Minnesota. It was supposed, before studying it, that the region, because of its position, would furnish many lichen species new to the State and to the interior of North America, and the investigation has fulfilled expectations. Besides its geographical position, certain physical features have produced diversity of lichen species as will be shown later. The lichens of the region have never been studied previously. Tuckerman, in his Synopsis,* mentions collections from the north shore of lake Superior by John Macoun and L. Agassiz. The collections by Agassiz were made in 1848 and published f from 1850 to 1852. He traversed the north shore from Sault Saint Marie to Fort William. The collecting by Macoun was done in July, 1869, along the north shore in Canada, and in 1884 around lake Nipigon. A publication may soon be ex- pected from the latter collector, giving a complete list of the Canadian lichens, and this should add much of interest con- cerning some species listed in this paper. Of the two collec- tors, Agassiz, at Fort William, came within about 50 miles, while Macoun probably came within 150 miles of certain points * Tuckerman, E., Synopsis of the North American Lichens, Parts I and II, 1882 and 1888. + Agassiz, L., Lake Superior, its Physical Character, Vegetation and Animals compared with those of other and similar Regions, pp. 170-174. Boston, 1850. 215 216 MINNESOTA BOTANICAL STUDIES. reached by me along the international boundary or the shore of the lake. Several species of the lichens collected by Dr. C. C. Parry in 1848 and published in 1895 * show northern range, but care- ful investigation | indicates that they were collected south of lake Superior. Thus it appears that the collections listed herein are the first made on the north shore of lake Superior in Minnesota, or along the international boundary for more than 100 miles west of the lake. The collecting was done by the writer, assisted by Mr. A. S. Skinner, during the latter part of June and the whole month of July, 1897. We were fortunate enough to be associated with Dr. A. H. Elftman, who wished to traverse the region for geo- logical study and whose thorough knowledge of the territory covered, alone made it possible for us to find the best collecting stations in this for the most part uninhabited region, and thus to accomplish good results in a comparatively short time. We reached Grand Portage island on the north shore of lake Superior, June 16th, and began collecting at once. The plan was to study the lichen flora of the international boundary and the north shore of lake Superior within the boundaries of Minnesota and to reach some of the inland portions of the two counties. On the whole trip we sought localities as collecting stations offering the greatest differences as to elevation, rock formations, arboreal flora, soil, moisture, etc. Beginning at Grand Portage island we traveled westward by Pigeon river and the chain of lakes along the international boundary to Gunflint, thence south by a series of lakes to Poplar river and down the river to Lutsen, on the shore of lake Superior. From here we proceeded down the lake to Tofte, Beaver Bay and Two Har- bors. We next went directly to Ely and thence eastward to Snowbank lake. We made as thorough a study as possible of the lichen flora of the Grand Portage area, and then stopped for study whenever and wherever we found enough of difference in * Fink, B., Lichens collected by Dr. C. C. Parry in Wisconsin and Minne- sota in 1848. Proc. Iowa Acad. Sci. 2: 137. 1895. + Parry, C. C., Systematic Catalogue of Plants of Wisconsin aud Minnesota, made in connection with the geological survey of the Northwest during the season of 1848. In Owen, D. D., Report of a geological survey of Wisconsin, Iowa, Minnesota and incidentally of a portion of Nebraska Territory. Appen- dix, article V. 606-622. 1852, mentions Cladonia rangiferina (\.) Hoffm. and Gyrophora muhlenbergii Ach. from Falls of St. Croix. Fink: LICHENS OF THE LAKE SUPERIOR REGION. PANY physical environment to lead to the conclusion that time would be profitably employed. The writer was relieved of camp duty as much as possible, so that there was some time for collecting each day, even when traveling. However, the few collections thus made are nearly all recorded in the list with those of the nearest well studied locality. The principal collecting stations are given below, with elevation and time spent in collecting for each. Since many of these stations are in uninhabited and little known regions, I have given the township and range of each one. I. Grand Portage and Grand Portage island, 9 days, elevation Goato 1305 feet, Tog N-; RGB. II. English portage, 3 hours, elevation 1339 feet, T. 64 N., R. 4.E. ; III. South Fowl lake, 3 hours, elevation 1436-1450 feet, T. 64 N., R. 4 E. IV. Moose lake, 2 hours, elevation 1492 feet, T. 65 N., R. arn. Moelkose lake. daypelevation, 1526 feet, i. 65.IN.,k. 1 We VI. Paulson iron mines, 2 days, elevation 1825 to 2000 feet, W265 N.;. R. 4, W. VII. Gunflint, 1 day, elevation 1547 to 1650 feet, T. 65 N., R. ai: VIII. Winchell lake, 2 days, elevation 1910 to 2230 feet, T. 64.N., Ra 2 W. IX. Brule lake, 5 hours, elevation 2084 feet, T. 63 N., R. 3 W. X. Tofte, 3 days, elevation 927 to 1529 feet, T. 59 N., R. 4 W. XI. Beaver Bay, 2 days, elevation 602 to 1250 feet, T. 55 N., R.8 W. XII. Great Palisades, 6 hours, elevation 602 to 1200 feet, T. 56 NZ Roy Ww. XIII. Two Harbors, 2 hours, elevation 692 feet, T. 52 N., R. Em VV: XIV. Prairie portage, 1 day, elevation 1300 feet, T. 64 N.,R. g W. XV. Iron Mountain lake, 1 day, elevation 1342 feet, R. 64N., R. 8 W. XVI. Snowbank lake, 4 days, elevation 1424 feet, T. 64 N., Reig WwW. XVII. Disappointment lake, 1 day, elevation 1449-1850 feet, E64 Ni Se W |) On ae ‘ is © XVIII. Moose lake, 1 day, elevation, 1339 feet, T. 64 N., R. 9 W. XIX. Wind lake, 1 day, elevation 1359 feet, T. 64 N., R. 9 W. XX. Ella Hall lake, 3 hours, elevation 1306 feet, T. 64 N., R. 10 W. XXI. Fall lake, 3 hours, elevation 1313 feet, T. 63 N., R. rr W. Of the stations given above, numbers XIV to XIX inclusive have been designated in the list of species as the Snowbank lake area, XX and XXI as Ely, VIII and IX as the Misquah hills, and VI and VII as Gunflint. All other areas include each but asingle station. The quantity-collecting being largely done when we left Grand Portage, we were able to move rapidly, as only plants not found in this first area needed to be collected in bulk. For illustration of distribution, the collections were made as complete as possible at Grand Portage, Gunflint, in the Misquah hills, at Tofte, at Beaver Bay and in the Snowbank _ lake area. There is an appreciable difference between the lichen flora of Grand Portage island and that of the mainland two miles across the bay. The island reaches an elevation of only 125 feet above lake Superior while Mt. Josephine on the mainland reaches an altitude of about 800 feet above the lake. The Keweenawan series of rocks, which appears on the island, is wanting on the portion of the adjacent mainland explored, while the Animikie series is found in both places. However, I could not ascertain that difference in petrographical construction in any noticeable way determines the floral differences either here or elsewhere in the territory studied. Passing by the Gunflint area for the pres- ent, I may say that the Misquah hills were regarded as especi- ally important, since they contain the highest areas in the state, and were as carefully studied as our time would permit. Carl- ton peak at Tofte, and the Palisades were points of special in- terest. Two Harbors was of interest as it is the most southern point reached in the survey, and Ely was, also, as it is the most western. However, the Snowbank lake area somewhat further east was much more thoroughly studied than Ely. On the whole all of the two counties was studied thoroughly enough to know that practically all of the lichens generally dis- tributed over the area were secured as well as many more which 218 MINNESOTA BOTANICAL STUDIES. Fink: LICHENS OF THE LAKE SUPERIOR REGION. 219 as yet show only local occurrence. Examination of the route will show that we covered all of the international boundary be- tween Ely and Grand Portage, except about 20 miles in a straight line from the most eastern portion of the Snowbank lake area to Gunflint. Thus the boundary was well studied. The line of travel through the Misquah hills from Gunflint to Lutsen gave a fair view of the interior of the region as well as the highest area in the state. A day spent along the shore at Lutsen failed to furnish any species not found at Grand Port- age. Consequently, as we were to stop at Tofte, only 10 miles distant, no record was made of the species found at the former place. Tofte, Beaver Bay and the Palisades gave a good view of the lake shore and higher elevations near by in the Sawteeth mountains. It is to be regretted that we did not have oppor- tunity for examination of the lake shore and Sawteeth moun- tains at some points between Lutsen and Grand Portage, but doubtless the number of additional species would not have been large after a thorough examination of the shore both to the ex- treme north and toward the south of the area studied. The whole region is one of extreme interest to the lichenist because of the diversity of natural conditions which gives a flora rich in individuals as well as variations within certain species which attracted special attention. The great masses of igneous and metamorphic rocks along the Superior and inland shore lines, the same rocks back from shore lines and the coniferous _and various other trees together with diversity as to temperature, moisture and elevation, all help to produce a flora richer in lichen species than I had expected to find. Though the annual precipitation of moisture for the area is not large, yet the com- paratively impervious nature of the rocks causes the water to collect in depressions of surface, forming a multitude of lakes of various sizes whose moist borders are a veritable paradise for lichens and especially for lithophytic species. The dense forests also hold moisture and favor lichen growth. When one finds single branches of Usnea longissima Ach. five feet long, as we collected on Grand Portage island, he realizes the signifi- cance of the name. Here and in some other localities of the region studied the dying conifers especially are literally covered with this plant, other species of the genus and A/ectorza jubata (L.) Nyl., all growing in a tangled profusion which obscures the host and when wet with rain or dew furnishes a view of sur- 220 MINNESOTA BOTANICAL STUDIES. passing beauty. Hardly less remarkable is the growth of Cla- donia rangiferina (L..) Hoffm. in open woods near Mt. Josephine, single clusters measuring three or four feet across and reaching a foot in height. ‘This plant was also common on rocks and in crevices exposed to wind and sun, but was always much smaller in such locations. It is evidently not a natural pioneer among lichens, but grows after other plants have attacked the rocky substratum, or on a thin layer of soil in crevices, and best of all after trees or shrubs have grown sufficiently to protect it some- what from wind and sun and have not yet become large enough or thick enough to kill it out. This same kind of ecological relation favors Cladonza furcata (Huds.) Fr., a variety of which was found fruiting only in such environment. More is given below about other C/ladonias, and the observation could be extended to Stereocaulon. After fires have passed over a region destroying the trees and small scattered second growth begins to appear to furnish some protection, Cladonza cristatella Tuck. and a large variety of forms of C. gracilis (L.) Nyl. soon begin to grow in great pro- fusion on old stumps, prostrate logs and bits of decaying wood lying upon, or more or less sunken into the soil. Only a few of the many varieties of the latter plant allowed by European lichenists are recognized in the list of species though forms closely resembling other varieties, so called, are represented in my collections. Nothing seems to be gained by carrying the *¢ splitting” process to extremes without a study of life histor- ies. C. gracilis (L.) Nyl. in regions recently burned showed much less variation than in places where the species had been established longer since the burning, and a careful study of a large number of individuals in this region, extending over a ser- ies of years would enable one to trace the growth and variation within single individuals and thus establish varieties with cer- tainty. Great variety was observed in the plants in regions that had burned 15 or at most 20 years ago so that a study ex- tending over 10 years should be sufficient to give the desired data. Like Cladonia rangtferina (L.) Hoffm., C. créstatella Tuck. is extremely sensitive to environment. In regions where the plants are exposed to sun and wind and in stations of high ele- vation, the plants are much smaller than in better shaded and less elevated places. The relation of size to amount of protec- Fink : LICHENS OF THE LAKE SUPERIOR REGION. Pp | tion offered by trees and shrubs may be observed in many places between Gunflint and Lutsen, especially at Gunflint and in the Misquah hills. The effect of elevation, or rather the combined effect of elevation and exposure, was especially noted on Mt. Josephine and on Carlton peak. In both of these places the stunted condition was also noticeable in other Cladonzas and in lichens belonging to other genera. The part that lichens play in rock decay and soil formation was studied in a general way in the Grand Portage area, and some of the most noticeable facts are stated below. Grand Portage island contains 57 acres of land and furnishes sufficient variety as to substrata suitable to lichen growth to make the study interesting. The crustaceous lichens furnish most of the species which first gain a footing on the rocks, and of these were found on the island three or four species of Placodium, a half dozen or more rock Lecanoras as well as a larger number of Lvatoras, Lecideas and Buellias. Of the foliaceous lichens the U/7bzl2- cartas are most characteristically rock pioneers; but these were very rare on the island, which did not furnish the high bluffs that they seek especially. As soon as rock decay has begun, the less strictly crustaceous species begin to appear. Of these Pannaria microphylla (Sw.) Delis was especially noticed some- times growing on quite firm rock, but more frequently on rotten rock or a residual product of rock decay still 2” sz¢z and pro- tected by the lichen though sometimes several inches deep. Next come the typically foliaceous and fruticulose species as the Peltigeras and Cladonias. Finally enough soil is established so that smaller Spermaphytes and finally trees and shrubs be- come established, these larger ones in turn furnishing substrata for epiphytic lichens. At the present time, trees grow at one end of the ridge of highest land extending across the island while the other end is bare of trees and soil to a large extent and yet supports many strictly lithophytic lichens. At the shore line one finds amphibious Azdocarpons and a Collema while typically xerophytic species cover the remainder of the island. The analysis could be extended to include a statement of differ- ent sorts of woody substrata which result in giving diversity of lichens growing on wood and, indeed, to give a detailed account of substrata including that of each one of the 88 species and varieties listed from the island. But this would lead to more detail than can be undertaken here, and for more minute ac- 222, MINNESOTA BOTANICAL STUDIES. count I have been compelled to select very small islands, though not offering so much diversity as to substrata, and have even then confined the analysis to the lithophytic and a few epigean species. For this study of islands three were selected in the Snowbank lake area, and the lithophytic species were carefully noted on two of them and on the other also the decrease in number due to the establishment of an arboreal flora. It is to be regretted that the study could not have been extended to more islands and to include epiphytic and epigean species as well as lithophytic. Island number one is situated in Sucker lake, 30 feet from the shore, in.the N. W..4% of S. W ¥ of S. E. Y ob Seen a Pian N:, R. 9 W. The size of the island is about 7ox 75) feet) Phe surface is rocky with soil in a few places formed zz sztu or washed in from the lake, so that C7adontas were well estab- lished. About twenty shrubs were growing on the island and two rather small pines. The species noted in a short time are as follows: 1. Cladonia rangiferina (L.) Horr. 2. Cladonia rangiferina (L.) Horrm. var. sylvatica L. 3. Cladonia rangiferina (L.) Horr. var. alpestris L. 4. Cladonia pyxidata (L.) Fr. 5. Cladonia gracilis (L.) FR. 6. Cladonia uncialis (L.) Fr. 7. Stereocaulon paschale (L.) Fr. 8. Umbilicaria muhlenbergii (Acu.) Tuck. 9g. Endocarpon fluviatile DC. 10. Parmelia conspersa (EuruH.) AcuH. 11. Parmelia saxatilis (L.) Fr. 12. Parmelia caperata (L.) Acu. 13. Physcia‘sp. 14. Physcia stellaris (L.) Tuck. 15. Physcia speciosa (Wutr., Acu.) Nyt. 16. Physcia obscura (Enuru.) Nyv. 17. Ephebe solida Born. 18. Pannaria microphylla (Sw.) DeE.is. Fink: LICHENS OF THE LAKE SUPERIOR REGION. 223 1g. Urceolaria scruposa (L.) Nyt. 20. Placodium vitellinum (Euru.) Narc. and Hepp. 21. Lecanora rubina (ViLL.) Acu. 22. Lecanora cinerea (L.) SOMMERF. 23. Buellia petrea (Fior., Koers.) Tuck. Island number two is in Snowbank lake, 50 feet from the snore, in the N.-W. 4 N. E. Y of Sec. 29, T. 64 N., R. 8 W. near the outlet of the lake. The size of the island is about 80x 100 feet, and it is thickly covered with trees and shrubs except in a few spots where Cladonia rangiferina (L.) Hoffm. persists. The species listed below for this island are excepting the Cladonza, confined to a circle of rock extending around the island and up from the water three inches to one foot. The species are as follows: 1. Cladonia rangiferina (L.) HorrM. . Endocarpon fluviatile DC. . Parmelia conspersa (Euru.) Acu. . Parmelia caperata (L.) Acu. . Physcia obscura (Euru.) Nyt. . Leptogium lacerum (Sw.) Fr. Placodium aurantiacum (Licutr.) NArEc. and Hepp. . Lecanora subfusca (L.) Acu. Wel (eo) (Si sen Cr OS Se 1S . Lecanora cinerea (L.) SoMMERF. Island number three is in Disappointment lake, about 200 feet from the shore, in the N. E. 4 of the S. E. 4% of S. E. Y% of nec. 33, T. 64 N., R. 8 W. The size is 50x75 feet. The surface is rocky, with a few small shrubs growing in crevices, and is literally covered with rock lichens, and a few Cladonzas and Stereocaulons growing along crevices and beginning to spread in one or two places. The following species were easily detected. . Cladonia rangiferina (L.) Horrm. Ll . Cladonia rangiferina (L.) Horr. var. sylvatica L. . Cladonia pyxidata (L.) Fr. . Cladonia uncialis (L.) Fr. . Cladonia furcata (Hups.) Fr. Up Se TS 224 MINNESOTA BOTANICAL STUDIES. 6. Stereocaulon paschale (L.) Fr. 7. Umbilicaria muhlenbergii (Acu.) Tuck. 8. Umbilicaria pustulata (L.) Horrn. g. Endocarpon fluviatile DC. 10. Parmelia conspersa (Euru.) Acu. 11. Parmelia saxatilis (L.) Fr. 12. Parmelia caperata (L.) Acu. 13. Physcia sp. 14. Physcia stellaris (L.) Tuck. 15. Physcia cesia (Horrm.) Nyt. 16. Leptogium lacerum (Sw.) Fr. 17. Ephebe pubescens Fr. 18. Ephebe solida Born. 19. Pannaria microphylla (Sw.) DELIs. 20. Urceolaria scruposa (L.) Nyu. 21. Placodium vitellinum (Euru ) Narc. and Hepp. 22. Rinodina oreina (Acu.) Mass. 23. Lecanora rubina (Vi1LL.) AcH. 24. Lecanora cinerea (L..) SOMMERF. 25. Buellia petrea (FLor., Korrs.) Tuck. Comparing the lichens easily detected on islands numbers one and three—those which give character to the flora—whatever rare species may have escaped notice, we find that, of a total of 23 species and varieties on the first and 25 on the second, 19 are com- mon to both islands, separated by several miles. The lists as a whole show a large number of foliaceous and fruticulose species ; and we evidently have not the primitive post-pleistocene lichen population of these rocky islands, which indeed must have dis- appeared centuries ago. It is the more remarkable that practic- ally the same species have succeeded in replacing a former flora on the two islands. I regret that time was wanting for the study of more of these islands, and especially of some farther from the shore line. The growth of larger forms of vegetation is probably beginning to effect a decrease in lichen species on island number one for otherwise, being larger, it should have Fink: LICHENS OF THE LAKE SUPERIOR REGION. 220 furnished more species than number three rather than a smaller number. But it was only on number two that we found the un- mistakable evidence of the effects of the arboreal vegetation in exterminating the lichens. Here too the species existing are all but two the same as those found on one or both of the other islands, but the number is reduced to little more than one-third -as many as occur on either of them. The succession of species is as apparent upon trees as upon rocks and is constantly in evidence in this largely undisturbed region where trees of various ages grow side by side. Some of the crustaceous lichens, of such genera as Pyrenula, Ar- thonia and Graphis, were usually found on young trees with smooth bark. As the substratum becomes more rugged with the increasing age of the tree, these are gradually replaced by foliaceous and fruticulose species as /tamalinas, Usneas, Par- melas, etc. Finally as the trees die certain species of Calicium Cladonta, Peltigera, Parmelia, etc., become the dominant types. It is not possible, nor is it necessary here, to give a detailed ac- count of relation between each epiphytic lichen and its host, but a few of the most apparent relationships are in order. Acer spicatum Lam. supports Arthonia dispersa (Schrad.) Ny]. over the whole area. Populus tremulotdes Michx. and P. bal- samifera L. bear Pyrenula leucoplaca (Wallr.) Kbr. commonly. Some conifers, as Prnus restnosus Ait., P. strobus L., Thuja occidentalis L.., serve for substrata for those species of the genus Calictum which grow on living trees. The most luxuriant growths of Usnea were found on Picea mariana (Mill.) and Abies balsamea (L.) Mill. Graphis scripta (L) Ach. var. recta (Humb.) Nyl. was almost wholly confined to Betula lutea Michx. and this same tree also supports Sagedia oxyspora (Nyl.) Tuck and two or three Pyrenuadas. A close analysis of the distribution of species within the area studied reveals much of interest even though it is a rather a re- stricted region. Of the 258 species and varieties listed, 96 were found only in one place, 32 in two, 31 in three, and the remain- ing 99 were collected along lake Superior and also inland, in four or more localities and are known to be generally distributed over the whole of the two counties. Also of those found in in two or three localities, 34 species were collected at some point along lake Superior and also beyond the ridge of high land formed by the Mesabi range and the Misquah hills and 226 MINNESOTA BOTANICAL STUDIES. are doubtless quite generally distributed over the territory sur- veyed. Of these found in two or three places, 13 more were found along the lake and inland, but none beyond the divide mentioned above. ‘These are doubtless generally distributed between this highest land and lake Superior, and of course may occur northwest of this high area as well. Of course the 133 or more species most generally distributed over the area largely determined the character of its flora and are interesting in studying the relation of the flora of the region to that of others. But for the study of distribution within the area, as influenced by natural conditions, the chief interest cen- ters around the 96 species found in one place only and those found in two or three areas and yet not generally distributed over the whole region. I give below alist stating the whole number collected in each principal collecting ground and also the number found at each one and not elsewhere. It will readily be seen that the last datum for each locality simply bears a close relative proportion to the first, or in other words, that no one area shows a large relative proportion of the rare species. Of course the data as to occurrence of these rare lichens can not be relied on fully; but about two-thirds of them are species of size large enough to be easy of detection, and while these may occur in other places, they are surely not common in the area. The table of species is as follows: Grand Portage island, total collected, &8, not found elsewhere, 15 Grand Portage, “6 66 Rey 6 -5 South Fowl lake, “6 6“ LAs eae rT - Rose lake, és 6“ 20, 66 | ee és 3 Gunflint 6 rT Tih. $e anes 6 s u ’ Misquah hills, “6 “< rg, eae 6 i Tofte, 66 +6 85; 10 ‘6 66 8 Beaver Bay, 6 6 G2, 4 Seay ee 66 13 Palisades, ois 6 o25 3 13 6G I Two Harbors, 6 v6 14 Soo 6 2 Ely, 66 a An, ae 66 a 3 Snowbank lakearea, ‘ 6s re em < aS The rather high per cent. of forms collected at Grand Port- age only is due to the fact that the attempt was made to collect here especially species not found on the island. At an average about one-eighth of the species collected in each locality were not found elsewere. As stated above, two-thirds of these are Fink: LICHENS OF THE LAKE SUPERIOR REGION. Pepal conspicuous forms. The remaining one-third are some of the less conspicuous S#zatoras, Lecideas, Puellias, Graphises, Pyrenulas, etc., which are not easily found. When we consider the limited size of the area studied, the restriction of certain of the rarer species to certain parts of it rather than to others is worthy of careful study. The area lies on two sides of a divide extending approximately east and west and formed of the Mesabi range and the Misquah hills. The alpine or sub-alpine species not generally distributed over the area are mostly confined to the portion lying between the divide and lake Superior and to the Misquah hills on the divide, and also those found along the lake are for the most part found toward the northeastern portion of the territory traversed. The rarer temperate region species on the other hand are most numerous to the north and west of the divide, or toward the south of the portion between the lake and the divide. All of the data given above as to distribution within the area studied are based upon carefully prepared lists showing the distribution of each species. They can not all be reproduced, but parts of them must be. First of all, the facts concerning the species found only in one place can only be properly presented for con- sideration by the somewhat laborious table below, giving the various localities and species for each. GRAND PoRTAGE ISLAND. Usnea cavernosa Tuck., N. Physcia hispida (ScHREB.) FR., N. Solorina saccata (L.) Acu., N. Lecanora calcarea (L.) Sommerr. var. contorta Fr., T. Cladonia gracilis (L.) Nyx. var. symphycarpia Tuck., T. Cladonia squamosa Horrm. var. phyllocoma Rasenn., T. Cladonia deformis (L.) Horrm., N. Bzomyces byssoides (L.) Scuarr., N. Biatora turgida (F'r.) Nyu., T. Lecidea spirea Acu., N. Endocarpon miniatum (L.) Scuaer., T. Lecidea crustulata Acu., N. Lecidea enteroleuca Fr. var. achrista SomMeERrF., T. iw) 28 MINNESOTA BOTANICAL STUDIES. Staurothele drummondii Tuck., T. Pyrenula cinerella (FLot.) Tucx., T. GRAND PORTAGE. Parmelia perforata (JAca.) Acu. var. hypotropa Nyu., T. Physcia adglutinata (FLoERK.) Nyv., T. Umbilicaria hyperborea Horrn., N. Nephroma levigatum Acu. var. parile Nyv., N. Placodium murorum (Horrm.) DC. var. miniatum Tuck., T. Lecanora muralis (SCHREB.) SCHAER var. diffracta Fr., T. Biatora leucophea FLorrk. var. griseoatra Korrs., N. Biatora lucida (Acu.) Fr., N. Lecidea lapicida Fr. var. oxydata Fr., N. Thelocarpon prasinellum Nyt., T. Verrucaria nigrescens PERs., i Dy: Pyrenula cinerella (FLotr.) Tuck. var. quadriloculata var. nov. (?). SouTH FowL LAKE. Alectoria sepincola (Euru.) Acu., N. ROSE LAKE. Heterothecium sanguinarium (L.) Fuor. var. affine Tuck., N. Opegrapha varia (Pers.) Fr. var. notha Acu., T. Usnea barbata (L.) Fr. var. dasyopoga Fr., T. GUNFLINT. Physcia cesia (Horrm.) Nyt., T. Pannaria nigra (Hups.) Nyu., T. Placodium cinnabarrinum (Acu.) Anz., T. Lecanora pallida (ScurEB.) SCHAER., T. Lecanora hageni Acu., T. Cladonia symphycarpa Fr. var. epiphylla (Acu.). Nyu., T. Cladonia fimbriata (L.) Fr., T. Cladonia gracilis (L.) Ny. var. cervicornis FLoERK., T. Biatora glauconigrans Tuck., T. Fink: LICHENS OF THE LAKE SUPERIOR REGION. 229 Lecidea acclinis Fior., T. Buellia petrea (FLor., Korrs.) Tuck.var. grandis FLoERk., N. Arthonia patellulata Nyv., T. MisQuaAH HILLS. Parmelia centrifuga (L.) Acu., N. Collema flaccidum Acu., T. Placodium murorum (Horrm.) DC., T. Lecanora subfusca (L.) Acu. var. hypnorum Scuarr., T. Pertusaria glomerata (AcH.) ScHArrR., N. Biatora oxyspora (TuL.) Nyu., T. Biatora schweinitzii Fr., T. Lecidea lapicida Fr., N. Lecidea albocerulescens WuLr.) ScHAER., N. Lecidea platycarpa Acu., N. Graphis scripta (L.) Acu. var. limitata Acu., T. TORT; Parmelia perforata (JAca.) Acu., T. Sticta limbata (Sm.) Acu., N. Leptogium myochroum (Euru.) ScHAErR., T. Lecanora elatina Acu., T. Stereocaulon coralloides Fr., N. Cladonia cespiticia (PErs.) Fu., T. Cladonia digitata (L.) Horrm., N. Calicium chrysocephalum (Turn.) Acu. var. filare Acu., T. BEAVER Bay. Ramalina calicaris (L.) Fr. var. fastigiata Fr., T. Ramalina pollinarella Nyv., T. Placodium cerinum (HEepw.) Narc. and Hepp. var. pyracea Nw, 1. Placodium vitellinum (Enru.) Narc. and Heprp., T. Lecanora calcarea (L.) Sommerr., T. bo 30 MINNESOTA BOTANICAL STUDIES. Cladonia mitrula Tuck., T. Cladonia delicata (Euru.) FL., T. Cladonia decorticata FLorErK., T. Biatora coarctata (Sm., Nyu.) Tucx., T. Biatora myriocarpoides (Nyv.) Tuck., T. Biatora negelii Hepp., T. Buellia myriocarpa (DC.) Mupp., T. Sagedia oxyspora (NyL.) Tuck., T. PALISADES. Cetraria islandica (L.) Acu., N. Two HARBORS. Physcia aquila (Acu.) Ny., T. Buellia dialyta (Nyt.) Tuck., T. ELY. Lecanora tartarea (L.) Acu., T. Buellia myriocarpa (DC.) Mupp. var. polyspora WittEy., T. Calicium trichiale (Acu.) var. stemoneum Nyt. T. SNOWBANK LAKE AREA. Ramalina calicaris (L.) Fr. var. canaliculata Fr., T. Ramalina pusilla (Prev.) Tuck., N. | Parmelia tiliacea(Horrm.)FLorrk. var. sublavigata Nyu.,T. Collema pycnocarpum Nyt., T. Leptogium lacerum (Sw.) Fr., T. Leptogium lacerum (Sw.) FR. var. pulvinatum Move. and Nespra 1. Rinodina sophodes (Acu.) Nyu. var. confragosa Nyu., T. Gyalecta fagicola (HEepp.) Tuck., T. Biatora spheroides (Dicxs.) Tuck., T. Biatora fuscorubella (HorrM.) Tuck., T. Biatora muscorum (Sw.) Tuck., T. Lecidea cyrtidia Tuck., T. Buellia parmeliarum (Sommerr.) Tuck., T. Fink: LICHENS OF THE LAKE SUPERIOR REGION. 231 Coniocybe pallida (PEers.) Fr., T. Verrucaria epigea (Pers.) Acu., T. In the above table all of the arctic and subarctic species are marked (N.) and the species common in temperate regions at low elevations (T.). Beginning then with the areas between the divide formed by the Mesabi range and the Misquah hills and lake Superior and toward the north of this region, of a total of 15 species found only on Grand Portage island, seven or one less than half are characteristic of northern regions. Of 12 confined to Grand Portage, five or about 42 per cent. are northern species. A single one collected only at South Fowl] lake and one of three at Rose lake are also northern. Of the 12 found only at Gunflint and the 15 found only in the Snow- bank lake area, only one strictly northern species is restricted to each place, while for Ely of the three restricted species not one is northern. Consideration of the figures will show that for the five localities along the international boundary there is a de- crease in proportion of rare arctic or alpine species in passing westward along the boundary. Again passing southward from Grand Portage we found three northern of a total eight species confined to Tofte, and the one species taken only at the Pali- sades is northern. The thirteen species found only at Beaver Bay and the two collected only at Two Harbors are all distinctly temperate region plants. Thus it appears that the northern species give way to those more characteristic of temperate re- gions in passing southward even along the shores of lake Su- perior where the cold lake winds have greatest influence upon the flora. An elevation of more than 1,000 feet was reached at Beaver Bay without finding northern species while at Grand Portage about 150 miles northeast they descend to the lake level. At Tofte a short distance northeast of Beaver Bay we reached an elevation of 1,529 feet on Carlton peak in the Sawteeth mountains and found three northern species. Only one of the three species was taken at the summit of Carlton peak, but this is because the top of the peak is burned over. The other two species were collected at a considerable distance above the base of the peak and are doubtless to be found on unburned portions of the Sawteeth mountains near by. In the discussion of the table thus far the Misquah hills area has not been considered. As stated elsewhere this area is the 232 MINNESOTA BOTANICAL STUDIES. highest portion of the State, reaching 2,230 feet, and the region gave five arctic or subarctic species of a total of eleven collected only here. The influence of elevation becomes apparent when we compare the Misquah hills area with the Gunflint region as to number of northern species. The latter locality lies 20 miles north of the former and about 400 feet lower, only a small por- tion of it near the Paulson mines reaching 2,000 feet. The Misquah hill area which is a more extended region of higher elevation furnished the goodly proportion of northern forms noted above while the Gunflint area gave only one such species in a total of twelve found only in the area. Of all the species found only in one place 25, or more than one-fourth, are arctic or subartic and 71, or approximately three-fourths are plants characteristic of temperate regions. Without entering into the yet more complicated analysis which a consideration of these rare temperate region plants would involve, a mere inspection of the table will show in a general way that their distribution is just the reverse of that of the north- ern species, or that they are especially characteristic of that small portion of the region studied which lies to the north of the divide and of the southern portion of the region lying between the divide and lake Superior. Of course it could also be shown that they are more especially characteristic of the lower elevations. Of the 63 species collected only in two or three places, only a half dozen are arctic or subarctic species, and it would have been useless to give the whole 63 in tabular form as no safe data could be obtained from so small a proportion of northern species. However, the 6 northern species are as follows: Ramalina pusilla (Prev.) Tuck. var. geniculata Tuck. Parmelia encausta (Sm.) Nyt. Sticta scorbiculata (Scop.) Ac. Lecanora frustulosa (Dicxs.) Mass. Lecanora sordida (Pers.) Tu. Fr. Buellia geographica (PERs.) Tuck. In order that we may have all of the northern species before us for a final consideration, I shall give a list of those generally distributed as follows: Lecidea lactea FL. Fink: LICHENS OF THE LAKE SUPERIOR REGION. 230 Buellia petrea (FLor., Korrs.) Tuck. Buellia petrea (FLor., Koers.) Tuck. var. montagnei Tuck. Umbilicaria vellea (L.) Nyv. Nephroma tomentosum (Horrm.) Neck. Pannaria epidiota Tu. Fr. Stereocaulon paschale (L.) Fr. Cladonia amaurocrea (FL.) SCHAER. Bzomyces zruginosus (Scor.) DC. Heterothecium sanguinarium (L.) Fior. Taking into account the above table, we find that of a total of 99 species and varieties generally distributed over the area studied only 10, or one-ninth, are arctic or subarctic, and con- sidering both of the last two tables we see that of 162 species and varieties more or less widely distributed 16, or nearly one- ninth, are arctic or subarctic. It has been stated that about one-fourth of the species found only in one place are such north- ern forms. Thus we find that the more general the distribution of a series of plants in the area the smaller the per cent. of northern species, and conversely the larger the per cent. of temperate region species. In other words the prevailing species are those characteristic of temperate regions, and as a whole the rarer ones are the more northern floral elements. Since the in- troduction of new species is commonly a more rapid process than the complete extermination of others in a given region, the existing conditions above stated seem to prove, as one would naturally suppose, that the present lichen flora of the region is in general of temperate region elements and that the more northern elements of the flora are the persisting for most part in a few favor- able spots. This supposition also explains the existence of the northern species in isolated regions further south as I have done for Taylors Falls. Professor Conway MacMillan has con- sidered the spermaphytic flora of this region as a south-bound one,* or at least that of the portion between the divide and lake Superior. My observations here and at Taylors Falls do not indicate that this is generally true of the lichens. However, because of somewhat milder temperature, lower elevation and perhaps more early retreat of the ice sheet in the western half * MacMillan C. Observations on the distribution of plants along the shore at Lake of the Woods. Minn. Bot. Stud. 1: 954. 1897. 234 MINNESOTA BOTANICAL STUDIES. of the State, the temperate region lichens have no doubt driven or followed the northern species farther north there and doubt- less very few of the latter elements now exist in western Min- nesota south of Lake of the Woods, or indeed anywhere in the western half of the state. Since gaining possession of the north- western part of the state these temperate region lichens have doubtless been moving southward over the Mesabi range to meet similar floral elements of a generally northward bound lichen flora. The western half of the state remains to be ex- plored for lichens, but the above statement as to the character of the flora, based on observations recorded in this paper for localities north of the Mesabi range and some knowledge of the general character of the region, I regard as sufficiently secure. Like the Taylors Falls region this one of course at one time contained only arctic species, and the present more numerous species characteristic of temperate regions have gained the ascendency in quite recent time. However, the problems in- volved in the struggle betwen the contending floral elements do not force themselves upon the observer so strongly in this larger area and must be dismissed with a much briefer state- ment. As in the Taylors Falls region the persisting northern species are largely lithophytic. This is shown in the following exhibit of substrata for the 42 species : Arctic or subarctic lichens confined to rocks..:..., .-0-s.eeeeeeee 25 ee 2 me 3 ‘* trees... eee eee 8 es ee ee oe 36 CC earth, 2. -2.-:ere eee 4 Ba ae a a ‘¢ dead) wood.7-es-eeaa--e 2 G27) ies “ a He ‘¢ dead wood and trees 1 oT aa oe Be 2 a ‘¢ earth and rocks, gs I 66 iad 66 66 66 ee trees 66 ce I The greater persistence of the lithophytic species is doubtless due in part at least to the greater stability of the rock surfaces and also probably in part to the fact that the arctic and sub- arctic species became more thoroughly established on the rocks, which were present for them to attack immediately at the close of the Pleistocene before the advent of large trees and temper- ate-region lichens. Thus far I have given a detailed account of habitat for the northern species only. Of the whole 258 species and varieties o Fink: LICHENS OF THE LAKE SUPERIOR REGION. 235 listed in this paper, so far as observation showed, 83 are epi- phytic, 80 lithophytic and 29 may occur on either trees or rocks. Another 29 are epigean, 22 were found only on dead wood, six are lithophytic or epigean, three are found on earth and dead wood, three are parasitic on other lichens and two were found on living and dead wood. ‘The above analysis of substrata is somewhat different from that used in the second paper of this series for comparison of substrata at Minneapolis and at Fayette, Iowa; but when reduced to that form shows that the percentage of lichens growing on rocks is somewhat higher than that for the two areas farther south, while the percentage of those grow- ing on wood is considerably lower. The larger proportion of rock lichens in the Superior region is due to at least three things. First, the more extensive exposure of rock surfaces, though this is offset in part at least by the existence of three distinct kinds of rock in the Minneapolis region, viz.: the igneous or meta- morphic boulders, the limestone outcrops and the Saint Peter sandstone. Second, the rock surfaces become warmer, each day in the warm portion of the year, than the trees, because of rapid absorption of heat; and this doubtless favors lichen de- velopment on rocks in this northern region. Third, general moisture of much of the surface due to the fact that the rocks are comparatively impervious to water, so that much of it col- lects in lakes and swamps, favors good development of rock lichens as compared with the region about Minneapolis. At Minneapolis unshaded rocks bear very few lichens, but in the Superior region rocks are well populated with them at all eleva- tions and in all sorts of environment at or above the water line, except where killed by fire. The total number of genera for the region is 39, while the number for Minneapolis and Taylors Falls, so far as is known, is 34. The whole number for Illinois, as given by Wolf and Hall* is 40 and for Iowa as recorded by the writer in two papersf is 38. This total number of genera for the Superior region is seen by the comparisons with both larger and smaller areas further south to be rather large for a somewhat limited northern * Wolf, John and Hall, Elihu. A List of the Mosses, Liverworts and Lich- ens of Illinois. Bull. Ills. State Lab. Nat. Hist. 2: 18-34. June, 1878. {+ Fink B. Lichens of Iowa. Bull. Lab. of Nat. Hist., State Univ. of Iowa 3: 70-88. Mr. 1895, and Notes concerning Iowa Lichens, Proc. Iowa Acad. Sci. “5: 174-187. 1897. 236 MINNESOTA BOTANICAL STUDIES. area, but a locality where northern and more temperate floral ele- ments meet seems to compensate fully at least for difference in latitude. The genera giving most of the species new to the state are the following, given in tabular form with the total number of species collected in each genus and the number new to the State. feamalina, collected 6, new to the State 4. Cetaria, “ mat) yee Ci eC Stzcta, ‘< an pt fo eae Nephroma, v6 Ase g 6 Ge ae Solorina, 6 iis OG Cc mee Lecanora, coop ae wa ae cc) ee ines Gyatlecta, rz righ 03° cet acre aed Beaomyces, 66 pony 36 oti,!! Nicer Mie Biatora, SRNR ee CoA ile Page Fleterothectum, ‘‘ Dir hee cc: yin c ee Lecidea, Kc) sree ae c6 WoT gh Calicium, “ Q;° 1% co Ba! Coniocybe, 66 Tay ote Gol Ua aa Sagedia, 66 ts a 66 66 rt, Of these genera Solor7na, Baeomyces, Heterothectum, Calic- zum, Conzocybe, and Sagedza are new to the State. An inspec- tion of the above list shows that the genera are for the most part those furnishing large numbers of arctic and subarctic species, or species hitherto supposed to be confined to New England. The genera Stereocaulon and Umbilicarta are equally characteristic of northern and eastern areas, but two- thirds of the species of these genera here recorded were listed for Minnesota in the first paper of this series. By comparing the present list of species and varieties with those recorded for Minneapolis and Taylors Falls we find that that there are 152 lichens growing in the territory considered in this paper and not found in either of the two areas named above, while there are 33 found in them and not in the northeastern Minnesota area under consideration. ‘This leaves only 73 lichens known to be common to central and northeastern Min- nesota. In the comparison between Minneapolis and certain localities in northeastern Iowa it was shown that no species have been found at the former place and not in the latter region, though Minneapolis is 150 miles north of Fayette, the principal Iowa area considered. In passing about 200 miles north from Fink: LICHENS OF THE LAKE SUPERIOR REGION. Da Minneapolis, on the other hand, we find a lichen flora, about three-fifths of whose species and varieties are not found at Min- neapolis and about half of which are new to the state. The region of rapid transition in lichen species lies between the Minneapolis and Superior areas and has only been touched in the study of its rock lichens at Taylors Falls. As stated in the third paper of this series, this region is one of special interest for tracing the distribution of species. The cause of the great difference in lichen flora between Minneapolis and the Superior region is scarcely due in any great measure to difference in latitude since an almost equal difference in latitude to the south of Minneapolis caused no appreciable difference in the flora. Also, I have shown in this paper that in three localities lying in the northern part of the region studied in the paper, viz.: Snowbank lake, Ely and Gunflint, very few typically northern species are found. These regions at the north of the area are more closely related to the Minneapolis region as to lichen flora than others 50 or 75 miles further south. The difference in lichen flora between central and northeastern Minnesota seems then to be due chiefly to three factors. The first is difference in substrata. The limestones of the Minneapolis regions, as well as the sandstone, are almost entirely wanting in northeast- ern Minnesota, being replaced by an abundance of igneous or metamorphic rocks. The conifers, which abound in the north- ern part of the State, and which serve for substrata for quite a number of species not found southward, form the other chief difference as to substrata. Location in the valley of lake Superior, where the region is shut off from warmer regions west and north as well as south, is another factor that has caused much of the difference in flora. It has been shown that the number of arctic and subarctic species for a given elevation decreases in passing southwestward along the lake. This I suppose to be due not so much to difference in latitude as to the fact that in the northeastern part of the region studied along the lake the winds coming from the broader expanse of water, on this largest American fresh water area, are rendered cooler than farther down where the lake is not so wide. That the cold winds are a factor is demonstrated by the occurrence of a large proportion of northern species at the Grand Portage area, and especially on the island which rises little more than 100 feet above the lake, while such inland areas as Gunflint and Snow- 238 MINNESOTA BOTANICAL STUDIES. bank lake, which are somewhat farther north, are almost entirely devoid of such species. A third factor is increase in elevation. The influence of elevation has been discussed in considering the Misquah hills and the Sawteeth mountains. Concerning cold lake winds and elevation, it is significant that of the 25 arctic and subarctic species found only in one place, 16 are found at stations along the lake, and that of the remain- ing 9, 5 are found in the Misquah hills, the region of highest elevation. Thus all but 4 of these 25 species were collected where one or both of these factors have most influence. It is not possible to state just which ones of the many species found either in central or in northeastern Minnesota and not in the other area would be of most interest in studying the territory lying between the two regions. Of course, the foliaceous and fruticulose species are most easily found, and some of these are most likely to be collected. I may add that interest would centre chiefly about species which are common and give char- acter to the flora in one of the two areas and are not found in the other. Not attempting to select from some 150 species col- lected in the Superior area and not farther south in the state, I will name, from 27 or 28 species found to the south and not to the north of the unexplored area, Theloschistes concolor (Dicks.) Tuck. and Physcia granulifera (Ach.) Tuck. as two species that any botanist can soon learn to distinguish in the field, which are common in the south half of Minnesota and not known farther north in the state, and whose distribution be- tween Minneapolis and Two Harbors would be of special inter- est. Nearly all of the remainder of the 27 ef 26 jspeciessare either infrequent or rare about Minneapolis, are confined to sub- strata not existing in northeastern Minnesota, or are so incon- spicuous as to render their study in the field difficult. As a fitting close to these observations on the distribution of lichens in different regions of the state I may give some notes concerning certain species for most part characteristic of more southern portions and found also in northeastern Minnesota. Parmelia borreré Turn. was not found fruited along the lake north of Beaver Bay. Peltigera aphthosa (L.) Hoffm. seeks high ground in the southern stations from which it is recorded and in the northwestern as well and is one of the species whose farther southern and western extent in the state would be espe- cially worthy of study. Parmelia tiliacea (Hoffm.) Tuck. be- Fink: LICHENS OF THE LAKE SUPERIOR REGION. 239 comes somewhat common at the southwest and more so at the northwest portion of the territory studied. It is one of the southern intrusions which extend farther north at some distance from lake Superior. Trees common farther south in Min- nesota were noted north of the Mesabi range as Quercus macro- carpa Michx. Crategus sp. and Fraxinus sp. On Fraxinus was found Conzocybe pallida (Pers.) Fr. which is common in northeastern Iowa, and on the same host Pyrenula leucophaca (Wallr.) Kbr. also becomes common for the first time in this northern area. Other lichens in the list of those found only north of the Mesabi range or the Misquah hills could be se- lected for special treatment, and on the whole this portion of the Superior region shows a closer floral connection with central Minnesota than does the most southern point reached, viz., Two Harbors. The list of 258 species and varieties is a large one for a rather limited area to yield, especially when it is stated that only about 300 lichens have been listed for Minnesota, including the present list. Yet the undisturbed portions of the region are more remarkable for richness in individuals than for large numbers of species. The Grand Portage area gave 132 species and varieties and the Snowbank lake area 121. These two areas are the ones best studied and are perhaps as thoroughly explored as the Minneapolis area, which gave 113 species and varieties. Both of the former two are like the latter small areas and the comparison seems to indicate that the lake Superior region is, area for area, only slightly richer in species than the Minneapolis region. Professor L. H. Bailey* found that the species of Sfermaphytes and higher Archegonzates of the region are only about one-tenth as numerous at lake Vermilion as in similar areas six degrees farther south, and we should of course expect some decrease in liehen species rather than an increase in passing northward in the state. The reverse con- dition existing is largely due to diverse conditions within the region as to elevation and temperature and as to surface moisture, all of which factors have been duly considered. The Snow- bank lake area studied is a larger one than the Grand Portage. About equal time was spent at the two places, and we did much more of the time-consuming quantity-collecting at Grand Port- * Bailey, L. H., in Arthur, J.C. Report on botanical work in Minnesota in 1886. Geol. and Nat. Hist. Surv. of Minn., Bull. 3:8. 10O., 1887. 240 MINNESOTA BOTANICAL STUDIES. age. Yet fewer forms of lichens were collected about Snow- bank lake. This seems to indicate that the lake shore is richer in lichen species than in interior areas of the territory studied. It is well known that a large portion of the species of lichens of the interior of North America are those found also in regions bordering upon the Atlantic ocean along our eastern border. This was brought out by the writer in a previous paper,* but all the species recorded in that paper were temperate region lichens. It has remained for the present paper to record a large number of more northern lichens previously for the most part known only in arctic or subarctic regions, or descending from mountains farther south only along our Atlantic border. Of the 258 species and varieties listed below 46 are new to the North American interior or to the interior of the United States, and of these six are new west and north of New Eng- land. In treating of distribution the expression ‘‘ the interior of North America” means the area lying between the Appalachian system of mountains on the east and the Rocky Mountains on the west. A number of species noted as new to the interior have been reported from New York or Canada, and doubtless a few of them were previously found a short distance west of the Appalachian system of mountains. . LIST OF SPECIES AND VARIETIES. 1. Ramalina calicaris (L.) Fr. var. fastigiata Fr. On trees, rare. Beaver Bay, July 13, 1897, no. 677. 2. Ramalina calicaris (L.) Fr. var. canaliculata Fr. On trees, rare. Snowbank lake area, July 23, 1897, no. 895. Not previously reported from Minnesota. 3. Ramalina calicaris (L.) Fr. var. farinacea SCHAER. On rocks, common or frequent over whole area and rarely found on trees also. Grand Portage island, June 23, 1897, no. 106. Gunflint, June 30, 1897, no. 257. Misquah hills, July 3, 1897, no. 419, and July 7, 1897, no. 539. Tofte, July 10, 1897, no. 624. Palisades, July 15, 1897, no. 763. Snowbank lake area, July 24, 1897, no. 934. All except no. 763 were lighter colored than other forms of the species. No. 106 occasionally and no. 763 quite commonly * Fink, B. Lichens of Iowa. Bull. Lab. of Nat. Hist., State Univ. of Iowa. 3: 70-78. Mr., 1895. Fink: LICHENS OF THE LAKE SUPERIOR REGION. 241 are irregularly branched and with dilated terminal soredia like R. pollinaria (Acu.) Tuck. 4. Ramalina pusilla (PReEv.) Tuck. On trees, frequent. Grand Portage island, June 19, 1897, no. 34. Snowbank lake area, July 20, 1897, no. 844. Not previously reported from Minnesota and new to the in- terior of North America. 5. Ramalina pusilla (Prev.) Tuck. var. geniculata Tuck. On trees, infrequent or rare. Gunflint, July 2, 1897, no. 375. Beaver Bay, July 13, 1897, no. 675. Not previously reported from Minnesota and new to the in- terior of the United States. 6. Ramalina pollinarella Nyv. On rocks, rare. Beaver Bay, July 13, 1897, no. 681. Not previously reported from Minnesota and new to the/in- terior of North America. 7. Cetraria aurescens Tuck. On trees, rare. Tofte, July 10, 1897, no. 636. Snowbank lake area, July 21, 1897, no. 869. Not previously reported from Minnesota and new to the in- terior of North America. 8. Cetraria islandica (L.) Acu. On earth above the Palisades, rare, July 15, 1897, no. 76s. Not previously reported from Minnesota. g. Cetraria ciliaris (Acu.) Tuck. On trees, abundant on Grand Portage island, elsewhere only rare or frequent. Grand Portage island, June 18, 1897, no. 10. Gunflint, July 2, 1897, nos. 387 and 396 at Misquah hills, July 3, 1897, no. 427. Snowbank lake area, July 22, 1897, no. 883, and July 26, 1897, no. 948. Ely, July 28, 1897, no. 1022. 10. Cetraria lacunosa Acu. On trees, common. Top of bluff at south end of South Fowl lake, June 26, 1897, no 206. Gunflint, July 2, 1897, no. gor. Misquah hills, July 3, 1897, no. 442. Tofte (Carlton peak), July 10, 1897, no. 556. Beaver Bay, July 14, 1897, no. 733. Snowbank lake area, July 21, 1897, no. 946. Not noted at Grand Portage where the last above was abun- dant, but seeming to replace it in part elsewhere, being com- mon in the localities noted above. 242 MINNESOTA BOTANICAL STUDIES. 11. Cetraria juniperina (L.) Acu. var. pinastri Acu. On trees and old logs. Common at Grand Portage and Mis- quah hills, elsewhere infrequent or rare, sterile. Grand Portage island, June 18, 1897, no. 15. Gunflint, June 30, 1897, no. 261. Misquah hills, July 3, 1897, no. 411. Beaver Bay, July 13, 1897, no. 679. Snowbank lake area, July 21, 1897, no. 865.. Ely, July 28, 1897, no. 1026, Not previously reported from Minnesota and new to the inte- rior of North America. 12. Cetraria sepincola (Euru.) Acn. On trees, rare. South Fowl lake, June 27, 1897, no. 201. Not previously reported from Minnesota and new to the in- terior of North America. 13. Evernia furfuracea (L.) Mann. On trees, rare, sterile. Gunflint, July 23 1607, me-sage Misquah hills, July 3, 1897, no. 434. ‘Tofte (Carlton peak), July 10, 1897, no. 573. Not previously reported from Minnesota. 14. Evernia prunastri (L.) Acu. On trees, common. Only seen fertile once. Grand Portage island, June 22, 1897, no. 81. Gunflint, June sje, ecoyem2- 267. Misquah hills, July 3, 1897, no. 428.9" Moten(@anionu peak), July 10, 1897, no. 554. Tofte, July 10, 1897, no. 635. Beaver Bay, July 14, 1897, no. 717. Two Harbors, July 17, 1897, no. 789. Snowbank lake area, July 20, 1897, no. 841. Ely, July 28, 1897, no. 1002. 15. Usnea barbata (L.) Fr. var. florida Fr. On trees, common or abundant but sterile. Grand Portage island, June 17, 1897, no. 8. Misquah hills, July 3, 1897, no. 420. 'Tofte (Carlton peak), July 10, 1897, nos. 557 and 607. Beaver Bay, July 14, 1897, no. 722. Snowbank lake area, July 14, 1897, No. 814. The last has the minute and numerous fibrils of var. Azrta Fr., but it is not sorediate; while the forms given below under that variety are sorediate, but the fibrils are seldom minute. 16. Usnea barbata (L.) Fr. var. hirta Fr. On trees, common or abundant but sterile. Grand Portage island, June 16, 1897, no. 9. Gunflint, July 1, 1897, no. 356. Misquah hills, July 5, 1897, no. 477. Tofte (Carlton peak), July ‘10, 1897, nos. 562 and 569. ‘Tofte, July 12, 1897, no. Fink: LICHENS OF THE LAKE SUPERIOR REGION. 243 649. Beaver Bay, July 13, 1897, no. 671. Snowbank lake area, July 19, 1897, nos. 806 and 851. Ely, July 28, 1897, no. IOOI. 17. Usnea barbata (L.) Fr. var. ceratina ScHAER. On trees, common or abundant but sterile. Gunflint, June 30, 1897, no. 263. Misquah hills, July 3, 1897, no. 423. Tofte, (Carlton peak), July 10, 1897, no. 558. Beaver Bay, July 13, 1897, nos. 665 and 672. Snowbank lake area, July 19, 1897, no. 807. Ely, July 28, 1897, no. 993. Not previously reported from Minnesota. 18. Usnea barbata (L.) Fr. var. dasypoga Fr. On trees, common. Rose lake, June 28, 1897, no. 213. 19. Usnea barbata (L. FR. var. plicata Fr. On trees, common. Snowbank lake area, July 19, 1897, no. 805. Ely, July 28, 1897, no. 994. 20. Usnea trichodea Acu. On trees, common. Grand Portage island, June 23, 1897, no.157. English portage, June 26, 1897, no. 190. Rose lake, June 28, 1897, no. 212. Gunflint, July 2, 1897, no. 381. Mis- quah Hills, July 3, 1897, no. 432, and July 5, 1897, no. 544. Tofte (Carlton peak), July 10, 1897, no. 547. Tofte, July 12, 1897, no. 659. Beaver Bay, July 13, 1897, no. 667. Two Harbors, July 17, 1897, no. 793. Not previously reported from Minnesota. 21. Usnea longissima Acn. On trees, common along lake Superior and possibly in Mis- quah hills. Sterile. Grand Portage island, June 18, 1897, no. 19. Misquah hills, July 3, 1897, no. 422. Tofte, July 12, 1897, no. 649. Beaver Bay, July 13, 1897, no. 649a. Two Harbors, July 17, ie no. O18 7 Snowbank lake area, July 20, 1897, no. 850. Specimens were collected at Grand Portage island five feet long. The plant breaks with its own weight and hangs abun- dantly over branches unattached. Not previously reported from Minnesota. 22. Usnea cavernosa Tuck. On trees, common. Grand Portage island, June 17, 1897, nos. 6 and 18. 23. Alectoria jubata (L.) Tuck. 244 MINNESOTA BOTANICAL STUDIES. On trees abundant on Grand Portage island, infrequent to common elsewhere. Sterile. Grand Portage island, June 18, 1897, no. 17. Gunflint, July 1, 1897, no. 354. Tome rsimiy: 12, 1897,/no. 658. Beaver Bay, July 13, 2807, mosnoso- Snowbank lake area, July 20, 1897, no. 837. Ely, July 28, 1897, no. 996. 24. Alectoria jubata (L.) Tuck. var. chalybeiformis Acu. On trees and old wood, frequent or common throughout. Sterile. High bluff at south end of South Fowl lake, June 26, 1897, no. 194. Gunflint, June 30, 1897, no. 258, and July 1, 1897, no. 316. Misquah hills, July 5,5) 1807, nese Beaver Bay, July 15, 1897, no. 786. Snowbank lake area, July 19, 1897, no. 815, and July 20, 1897, no. 842. Ely, July 28, 1897, no. 998. 25. Alectoria jubata (L.) Tuck. var. implexa Fr. On trees, infrequent. Sterile. Misquah hills, July 3, 1897, no. 421. Beaver Bay, July 14, 1897, no. gaa. Not previously reported from Minnesota. 26. Theloschistes polycarpus (Euru.) Tuck. On trees, infrequent or rare west of Gunflint, elsewhere fre- quent or common. Grand Portage island, June 18, 1897, no. 20. Gunflint, July 2, 1897, no. 393. Misquah hills, July 5, 1897, nos. 452 and 487. Beaver Bay, July 13, 1897, nos. 678 and 686. Ely, July 28, 1897, no. 985. 27. Theloschistes lychneus (Nyu.) Tuck. On rocks, rare. Grand Portage island, June 23, 1897, no. 147. Snowbank lake area, July 19, 1897, no. 830. 28. Parmelia perlata (L.) Acu. On rocks and rarely on trees, rare to infrequent except at Gunflint, where the species seemed to be common. Sterile. Grand. Portage, June 24, 1897, no. 169. Portage between South Fowl lake and Pigeon river, June 26, 1897, no. 205. Gunflint, June 7, 1897, no. 368. Misquah hills, July 5, 1897, no. 543. Beaver Bay, July 14, 1897, no. 725. Snowbank lake area, July 24, 1897, no. 925. 29. Parmelia perforata ( Jaca.) Acu. On trees, common. Sterile. Tofte (Carlton peak), July EO, 1907, DO.15 72- A puzzling plant with sorediate margined lobes and otherwise Fink: LICHENS OF THE LAKE SUPERIOR REGION. 245 resembling the last. However, the margins of the lobes are quite strongly ciliate, and the lower surface of the thallus inter- ruptedly so; possibly might be referred to P. perlata (L.) Acu. var. celiata. DC. Thallus rather thinner than my her- barium specimens of P. perforata. 30. Parmelia perforata (JAca.) Acu. var. hypotropa Nyt. On rocks, rare. Grand Portage, June 23, 1897, no. 116. Not previously reported from Minnesota. 31. Parmelia crinita Acn. On rocks and once collected on trees, rare or infrequent. Sterile. Grand Portage, June 23, 1897, no. 114. Gunflint, July z, 1897, nos. 361 and 362. Tofte, July 10, 1897, no. 627. The plant. differs from my Iowa and Ohio specimens in that it is strongly ciliate on the upper surface of the thallus among the branchlets and granules. 32. Parmelia tiliacea (Horrm.) FLOERK. On trees, rare or infrequent. Gunflint, July 2, 1897, no. 4o7a. Tofte (Carlton peak), July 10, 1897, no. 620. Bea- ver Bay, July 14, 1897, no. 724a. Ely, July 28, 1897, no. 1018. Really more frequent along boundary at west. Was fre- quently noted in Snowbank lake area and failure to get speci- mens was an oversight. 33. Parmelia tiliacea (HorrM.) FLoeRK. var. sublevigata NYL. On trees, rare. Snowbank lake area, July 23, 1897, no. 896. Not previously reported from Minnesota. 34. Parmelia borreri TuRN. var. rudecta Tuck. On rocks and trees, rare or infrequent, except common in the Snowbank lake area. All sterile except no. 744 on rocks. Grand Portage, June 24, 1897, no. 188. Gunflint, July 1, 1897, no. 369. Misquah hills, July 3, 1897, no. 431. Tofte (Carlton peak) July 10, 1897, no. 612. Palisades, July 15, 1897, nos. 744 and 762. Snowbank lake area, July 30, 1897, no. 839. Ely, July 28, 1897, no. 983. 35. Parmelia saxatilis (L.) Fr. On trees and rarely on rocks. Abundant or common and fre- quently fruited. Grand Portage island, June 17, 1897, no. 7, and June 21, 1897, no. 57.. Misquah hills, July 3, 1897, no. 246 MINNESOTA BOTANICAL STUDIES. 406. Beaver Bay, July 14, 1897, no. 741. Snowbank lake area, July 21, 1897, no. 873. 36. Parmelia saxatilis (L.) Fr. var. sulcata Nyv. On trees, probably common. Grand Portage island, June 17, 1897, no. 7a. Gunflint, June 30, 1897, no. 240. Not previously reported from Minnesota. 37. Parmelia physodes (L.) Acu. On trees and rocks, common. Gunflint, July 1, 1897, no. 383. Snowbank lake area, July 2, 1897, no. 883. Doubtless occurring over the whole region studied, but taken for P. saxatilis (L..) Fr. modified by some peculiarity of sub- stratum. 38. Parmelia encausta (Sm.) Nyt. On trees, common. Grand Portage island, June 6, 1897, no. 143. Tofte (Carlton peak), July 10, 1897, no. 565. Not previously reported from Minnesota and new to the in- terior of North America. 39. Parmelia olivacea (L.) Acu. On trees, common. Grand Portage island, June 18, 1897, no. 11. Grand Portage, June 23, 1897, no. 100. Gunflint, June 30, 1897, no. 260, and July 2, 1897, no. 395. Tofte (Carlton peak), July 10, 1897, nos. 585 and 591. Beaver Bay, July 13, 1897, no. 712. Snowbank lake area, July 20, 1897, no. 848. Ely, July 28, 1897, no. 1004. 40. Parmelia olivacea (L.) Acu. var. prolixa Acn. On rocks, common or frequent. High bluff at south end of South Fowl lake, June 26, 1897, no. 197. Gunflint, June 3o, . 1897, no. 290. Misquah hills, July 5, 1897, no. 491. Tofte (Carlton peak), July 10, 1897, no. 574. Beaver Bay, July 13, 1897, no. 703. Palisades, July 15, 1897, no. 742a. Snow- bank lake area, July 20, 1897, no. 832. Not previously reported from Minnesota and new to the in- terior of North America. 41. Parmelia caperata (L.) Acu. On trees and rocks, common. Grand Portage island, June 23, 1897, no. 107. Gunflint, June 30, 1897, no. 254a. Mis- quah hills, July 3, 1897, no. 403. Tofte (Carlton peak), July 10, 1897, no 608. Beaver Bay, July 14, 1897, no. 728. Pali- sades, July 15, 1897, no. 768. Snowbank lake area, July 19, 1897, no. 804 and July 24, 1897, no. 916. Fink: LICHENS OF THE LAKE SUPERIOR REGION. 247 More commonly fruited than farther south and especially well fruited in the Snowbank lake area. 42. Parmelia conspersa (Enru.) Acu. On rocks, abundant or common. Grand Portage island, June 23, 1897, no. 103. Gunflint, June 30, 1897, no. 289. Misquah hills, July 5, 1897, no. 506. ‘Tofte (Carlton peak), July 10, 1897, no. 621. Palisades, July 15, 1897, no.° 755. Snowbank lake area, July 27, 1897, no. 967. 43. Parmelia centrifuga (L.) Acn. On rocks, rare and sterile. Misquah hills, July 5, 1897, no. 496a. Not previously reported from Minnesota and new to the in- terior of the United States. 44. Physcia speciosa (WuLF. Acu.) Nyt. On rocks and more rarely on old wood or trees, infrequent or frequent. Grand Portage island, June 19, 1897, no. 27. Gun- flint, June 30, 1897, no. 270 and July 1, 1897, nos. 342 and 348. Misquah hills, July 3, 1897, no. 438, and July 5, 1897, no. 489. Tofte (Carlton peak), July 10, 1897, nos. §71 and 601. Snow- bank lake area, July 24, 1897, no. g11. Nos. 348, 438 and 489 fruited. 45. Physcia ciliaris (L.) DC. On rocks or high bluffs, rare. Grand Portage, June 24, 1897, no. 180. Bluffs at south end of South Fowl lake, June 26, 1897, no. 200. Palisades, July 15, 1897, no. 774. Not previously reported from Minnesota. 46. Physcia aquila (Acu.) Nyt. On trees, rare. Two Harbors, July 17, 1897, no. 791. The only distinct specimen noted and not found farther north. =—_ Not previously reported from Minnesota. 47. Physcia pulverulenta (ScureEs.) Nyv. On rocks infrequent or rare, and not often fruited. Grand Portage island, June 23, 1897, nos. 145 and 145a. Grand Port- age, June 23, 1897, no. 173. Gunflint, July 2, 1897, no. 374. Snowbank lake area, July 26, 1897, no. 964. A very variable plant. No. 1452 is the typical form with re- spect to the upper surface of the thallus, being lighter colored than the others and pruinose. Like the others it is usually dark 248 MINNESOTA BOTANICAL STUDIES. colored below. No. 173 yielded spores of the usual size meas- uring — mic. and much constricted in the middle. No. 374 gave spores only - oa mic. and scarcely constricted. 48. Physcia pulverulenta (Scures.) Nyv. var. leucoleiptes Tuck. On rocks and trees, probably rare. Sterile. Grand Portage island, June 23, 1897, no. 119. Misquah hills, July 5, 1897, no. 496. Snowbank lake area, July 27, 1897, no. 966. ‘Thal- lus black below. Not previously reported from Minnesota. 49. Physcia stellaris (L.) Tuck. On trees and rocks, frequent or infrequent. Grand Portage “island, June 23, 1897, nos. 105 and 142. Gzunflint, June 30, 1897, no. 283a, and, July 2, 1897, no. 394. Misquah hills, July 5, 1897, no. 460. Tofte (Carlton peak), July 10, 1897, no. 570. Beaver Bay, July 13, 1897, no. 7oo. Two Harbors, July 17, 1897, no. 801. Ely, July 28, 1897, no. 1012. Occasionally the tree forms show dark fibrils and even dark thallus below so that the rock growing variety, below, could only be distinguished certainly by the crenulate border of the apothecia. The plant is much more variable than farther south in Minnesota and Iowa. No. 570 yielded apothecia that were somewhat ciliate below, but the plant is white below and other- wise like the present plant rather than P. obscura (Ehr.) Nyl. 50. Physcia stellaris (L) Tuck. var. apiola Nyu. On rocks, frequent or infrequent. Grand Portage island, June 21, 1897, no. 108. Gunflint, June 30, 1897, no. 284. Misquah hills, July 5, 1897, no. 469. Tofte, July 12, 1897, nos. 640a and 641. Beaver Bay, July 13, 1897, no. 704. Snowbank lake area, July 26, 1897, no. 953. 51. Physcia tribacia (Acu.) Tuck. On rocks, rare and sterile. Grand Portage, June 23, 1897, no. 92. Misquah hills, July 5, 1897, no. 449. Snowbank lake area, July 20, 1897, no. 847. 52. Physcia hispida (ScureB. Fr.) Tuck. On trees, locally infrequent and poorly fruited. Grand Port- age island, June 21, 1897, no. 63. Fink: LICHENS OF THE LAKE SUPERIOR REGION. 249 53. Physcia cesia (HorrMm.) Nyt. On rocks, frequent locally. Sterile. Gunflint, June 30, 1897, nos. 282 and 292. 54. Physcia obscura (Euru.) Nyt. On rocks and trees, rare. Grand Portage island, June 23, Fog7, 00, 145b. © Grand Portage, June 24, 1897; no. 272: Tofte, July 12, 1897, no. 642. 9733 mic., and other- 13 No. 145b yielded spores a little large, a wise looks somewhat like P. fulverulenta (Schreb.) Nyl., but is black below. Apothecia have hispid.borders, and the thallus is much smaller than that of any plants referred to the latter. 55. Physcia adglutinata (FLoERK.) Nyu. On trees, rare. Grand Portage, June 23, 1897, nos. 82a and gfa. 56. Pyxine sorediata Fr. On rocks and trees, rare but widely distributed. Collected in fruit three times. Gunflint, July 1, 1897, no. 364, and July 2, HOO7,,.no.' 385. Misquah hills, July 35 1897; no. 437. ~“Tofte, ily HO, 1897, no. 633.) Palisades, July 15, 1807, no. 746: Two Harbors, July 7, 1897, no. 790. Snowbank lake area, July 24, 1897, no. 912. 57. Umbilicaria muhlenbergii (Acu.) Tuck. On rocks, common or abundant. Grand Portage (Mt. Jose- phine), June 21, 1897, no. 53. Gunflint, June 30, 1897, nos. 245 and 246. Misquah hills, July 5, 1897, no. 537. Tofte (Carl- ton peak), July 10, 1897, no. 602. Palisades, July 15, 1897, no. 771. Snowbank lake area, July 24, 1897, no. 938. 58. Umbilicaria vellea (L.) Nyu. On rocks, common or frequent. Grand Portage (Mt. Jose- poime), june 21, 1897, no.) 57, and) June 24, 1897, nosphye. Gunflint, July 1, 1897, no. 367. Misquah hills, July 3, 1897, no. 430, and July 5, 1897, no. 538. Palisades, July 15, 1897, no. 766. Snowbank lake area, July 24, 1897, no. 931. 59. Umbilicaria dillenii Tuck. Onrocks, common. Grand Portage, June 24,1897, no. 170. Gunflint, July 1, 1897, no. 370. Misquah hills, July 3, 1897, no. 429, and July 5, 1897, no. 445. Tofte (Carlton peak), July 250 MINNESOTA BOTANICAL STUDIES. 10, 1897, no. 599. Palisades, July 15, 1897, no. 775. Snow- bank lake area, July 24, 1897, no. 937. 60. Umbilicaria pustulata (L.) Horrm. var. papulosa Tuck. On rocks, rare to frequent. Spores reaching 100 mic. in length. Gunflint, July 1, 1897, no. 371. Misquah hills, July 3, 1897, no. 542. Palisades, July 15, 1897, no. 756. Snow- bank lake area, July 24, 1897, no. 932. 61. Umbilicaria hyperborea Horr. On rocks, rare, Grand Portage (Mt. Josephine), June 23, 1897, no. 104. Not previously reported from Minnesota and new to the in- terior within the United States. 62. Sticta amplissima (Scop.) Mass. On trees or rocks, rare or infrequent. Grand Portage, June 23, 1897, no. 115. Tofte (Carlton peak), July ao, 1ée7emue: 609. Beaver Bay, July 14, 1897, no. 720. Two Harbors, July 17, 1897, no. 802. | Ely, July 28, 1807,;00 4008. 63. Sticta pulmonaria (L.) Acn. On trees or rocks, common or frequent. Grand Portage island, June 19, 1897, no. 23. Rose lake, June 28, 1897, no. 225. Gunflint, July 1, 1897, no. 365. Misquah hills, July 3, 1897, no. 443. Tofte (Carlton peak), July 10, 1897, no. 396. Beaver Bay, July 14, 1897, no. 724. Two Harbors, July 17, 1897, no. 795. Snowbank lake area, July 19, 1897, no. 808. Ely, July 28, 1897, no. 984. 64. Sticta limbata (Sm.)iAcu. On trees, very rare. Tofte, July 10, 1897, no. 626. Only two plants collected with thallus also much like that of the Euro- pean Stzcta fuliginosa. Spores brown, two-celled, constricted, 19-23 8-9 Not previously reported from Minnesota and only once from North America. (Oregon, by J. W. Eckfeldt.) 65. Sticta crocata (L.) Acu. On trees and rocks, rare. Tofte (Carlton peak), July 10, 1897, no 597. Beaver Bay, July 13, 1897, no. 685. Snow- bank lake area, July 24, 1897, no. 940. Not previously reported from Minnesota and new to the in- terior within the United States. mic. Fink : LICHENS OF THE LAKE SUPERIOR REGION. 251 66. Sticta scorbiculata (Scop.) Acu. Mossy rocks and trees, rare except at Grand Portage where frequent. Grand Portage, June 24, 1897, no. 176. Tofte, July 10, 1897, no. 629. Not previously reported from Minnesota and new to the in- terior of North America. 67. Nephroma tomentosum (Horrm.) Korrs. On rocks and trees, frequent. Grand Portage island, June 19, 1897, no. 26. Misquah hills, July 5, 1897, no. 541. Tofte, July 10, 1897, no. 634. Snowbank lake area, July 19, 1897, no. 803. Not previously reported from Minnesota and new to the in- terior of North America. 68. Nephroma helveticum Acu. On rocks and occasionally on trees and earth, frequent. Grand Portage island, June 23, 1897, no. 133. Gunflint June, 29, 1897, no. 235, and June 30, 1897, no. 250. Misquah hills, July 5, 1897, no. 540. Beaver Bay, July 14, 1897, no. 740. Tofte (Carlton peak), July 10, 1897, no. 605. Snowbank lake area, July 24, 1897, no. 921, and July 26, 1897, no. 961. Ely, July, 28, 1897, no. 1005. 69. Nephroma levigatum Acu. On trees, rare. Misquah hills, July 5, 1897, no. 426. Two Harbors, July 17, 1897, no. 799. Not previously reported from Minnesota. 70. Nephroma levigatum Acu. var. parile Nyt. On rocks, locally common. Grand Portage, June 23, 1897, N06. 113. Not previously reported from Minnesota. 71. Peltigera venosa (L.) Horrm. On earth and mossy rocks, rare, spores reaching 60 mic. in length and occasionally five-celled. Grand Portage, June 23, 1897, no. 150. Portage at south end of South Fowl lake, June 26, 1897, no. 207. 72. Peltigera aphthosa (L.) Horr. On earth or rocks, common or frequent. Grand Portage Island, June 24, 1897, no. 177. Gunflint, June 30, 1897, no. 248. Misquah hills, July 5, 1897, no. 526. Palisades, July 15, 1897,no. 770. Snowbank lake area, July 19, 1897, no. 829. PAT MINNESOTA BOTANICAL STUDIES. 73. Peltigera horizontalis (L.) Horrm. On earth, frequent or common. Grand Portage island, June 19, 1897, no. 30 and June 23, 1897, no. 121. Gunflint, June 30, 1897, no. 247, and July 1, 1897, nos. 338 and 360. Mis- quah hills, July 3, 1897, no. 441. Tofte, July 12, m@ee7-enos. 630 and 631. Beaver Bay, July 14, 1897, no. 723. Palisades, July 15,1897,no. 761. Snowbank lake area, July 24, 1897, nos. 917 andg18. Part of the plants placed here agree somewhat with those reported elsewhere for Iowa and Minnesota as P. pulveru- lenta (Tayl.) Nyl., but though the sterile forms previously seen differ considerably from the fertile ones herein reported per- haps all must eventually be placed here. The sterile forms, occasionally light colored below, are crisped and broken prob- ably from unfavorable conditions which prevented their fruiting. Not previously reported from Minnesota. 74. Peltigera polydactyla (Nrecx.) Horrn. On earth,common. Spores reaching 110 mic. Grand Port- age island, June 23, 1897, nos. 140 and 144. Gunflint, July 1, 1897, no. 336. Misquah hills, July 5, 1897, no. 536. Beaver Bay, July 14, 1897, no 726. Snowbank lake area, July 24, LGO7, 110.9209, and July 27, 16907, no o7me 75. Peltigera canina (L.) Horr. var. spuria Acu. On earth, frequent or common. Grand Portage island, June 19, 1897, no. 29. Grand Portage (Mt. Josephine), June 21, 1897, no. 54.. Gunflint, July 1, 1807, Moos 424-emeecaner Bay, July 13, 1897, no. 663. Palisades; July 135snea 7eane: 760. Snowbank lake area, July 24, 1897, no. 943. 76. Peltigera canina (L.) Horr. var sorediata ScHAER. On earth, frequent. Grand Portage (Mt. Josephine), June 23, 1897 no. 118a. Gunflint, July 1, 1807, mon gaeeeolie- quah hills, July 5, 1897, no. 535. Tofte (Carlton peak), July 10, 1897, no. 589. Beaver Bay, July 13, 1667, mol003: Snowbank lake area, July 24, 1897, no. 902. Ely, July 28, 1897, no. 976. 77. Solorina saccata (L.) Acu. On earth, rare. Grand Portage island, June 24, 1897, no. 179. Not previously reported from Minnesota and new to the interior of North America. 78. Pannaria languinosa (AcH.) Korres. On rocks, common. Grand Portage island, June 17, 1897, Fink : LICHENS OF THE LAKE SUPERIOR REGION. 253 no. r. South Fowl lake, June 26, 1897, no. 195. Gunflint, June 30, 1897, no. 272. Misquah hills, July 3, 1897, no. 435, and July 5, 1897, no. 457. Tofte (Carlton peak), July ro, 1897, no. 561a. Beaver Bay, July 13, 1897, no. 691. Snow- bank lake area, July 19, 1897, no. 824. Nos. 195 and 457 showing the bright sulphur-colored plant common in Europe and only noted in North America by the present writer in the second paper of this series. 79. Pannaria microphylla (Sw.) DELIs. On rocks, frequent on Grand Portage island, June 19, 1897, no. 22. Gunflint, July 1, 1897, no. 330. Beaver Bay, July EAtoo7,,n0. 684. . Hly, July 28,1697, no: 987. 80. Pannaria lepidiota Tu. FR. On rocks and wood, infrequent. Grand Portage, June 24, 1897, no. 175. Gunflint, July 2, 1897, no. 372. Misquah hills, July 5, 1897, nos. 463 and 479. ‘Tofte (Carlton peak), July ro, 1897, no. 582. Snowbank lake area, July 24, 1897, no. gos. Not previously reported from Minnesota. 81. Pannaria flabellosa Tuck. On rocks, rare. Grand Portage island, June 23, 1897, no. 128. Sterile, but having the narrow linear lobed thallus and blue-black hypothallus. The thallus does not show the ex- panded and striated lobes at circumference. Not previously reported from Minnesota and new west of New England. 82. Pannaria nigra (Hups.) Ny. On rocks, rare. Gunflint, July 1, 1897, no. 347. 83. Ephebe pubescens Fr. = On rocks, rare, sterile. Palisades, July 13, 1897, no. 745. Snowbank lake area, July 24, 1897, no. gor. Not previously reported from Minnesota and new to the in- terior of North America. 84. Ephebe solida Born. (?) On rocks, rare and sterile. Misquah hills, July 5, 1897, no. 488. Beaver Bay, July 13, 1897, no. 683. Snowbank lake area, July 20, 1897, no. 849. A short form growing in small dense tufts. 254 MINNESOTA BOTANICAL STUDIES. 85. Collema pycnocarpum Nyt. On trees, rare. Snowbank lake area, July 27, 1897, nos. 968 and 974. 86. Collema flaccidum Acu. (?) On high rocks, rare. Misquah hills, July 5, 1897, no. 495. Sterile, but with the thallus corresponding with tree forms farther south, except that the plant is larger. 87. Collema nigrescens (Hups.) Acu. On trees, especially Populus, common or frequent. Grand Portage, June 19, 1897, no. 36, and June) 23,1607, mo.mon, Rose lake, June 28, 1897, no. 221. Gunflint, July 30, 1897, no. 266. Misquah hills, July 5, 1897, no. 483. Beaver Bay, July 15, 1897, no. 783. Snowbank lake area, July 19, 1897, no. 822, and July 21, 1879, no. 874. Ely, July 28, 1897, no. 1or4. 88. Collema furvum (Acu.) Ny? On wet rocks, frequent. Grand Portage island, June 23, 1897; no. 156. Tofte, July 12, 1897, no. 646. (iNet typical: Not previously reported from Minnesota. 89. Leptogium lacerum (Sw.) Fr. On rocks, rare and sterile. Snowbank lake area, July 21, 1307, NO. G67. go. Leptogium lacerum (Sw.) Fr. var. pulvinatum Move. and NESTL. On rocks, rare. Snowbank lake area, July 29, 1897, -no. 965. Not previously reported from Minnesota. 91. Leptogium tremelloides (L.) Fr. On rocks andrarely ontrees. Widely distributed, but usually rare locally. Portage between South Fowl lake and Pigeon river, June 26, 1897, no. 208. (Gunflint, July ayase7.mos 349 and 358. Misquah hills, July 3, 1897, no. 424. Beaver Bay, July 15, 1897, no. 781. Snowbank lake area, July 20, 1897, no. 843, July 26, 1897, no. 962, and July 27, 1897, no. 974a. Ely, July 28, 1897, nos. 977 and 1008. 92. Leptogium myochroum (Euru., ScHAER.) TucK. On trees and rocks, frequent. Grand Portage island, June 19, 1897, no. 24. Gunflint, July 1, 1807, non 341-eeome (Carlton peak), July 10, 1897, no. 598. Snowbank lake area July 19, 1897, no. 820. Not previously reported from Minnesota. Fink : LICHENS OF THE LAKE SUPERIOR REGION. 25b 93. Leptogium myochroum (Enurn., Scuarr.) Tuck. var. tomentosum SCHAER. On trees, rare. Tofte (Carlton peak), July 10, 1897, no. 559. Not previously reported from Minnesota. 94. Placodium elegans (Linx.) DC. On rocks, common. Grand Portage, June 21, 1897, no. 69. Grand Portage island, June 23, 1897, nos. 93 -and 98. Gun- flint, June 30, 1897, no. 294, and July 1, 1897, no. 328. Mis- quah hills, July 5, 1897, no. 444. Palisades, July 15, 1897, no. 747. Snowbank lake area, July 22, 1897, no. 887, no. 444, looking toward the next in having orange-red apothecia. 95. Placodium murorum (HorrM.) DC. On rocks, rare. , Misquah hills, July 5, 1897, no. 450. Not previously reported from Minnesota. 96. Placodium murorum (Horrm.) DC. var. miniatum Tuck. On rocks, rare. Sterile. Grand Portage, June 23, 1897, no. 88. Not previously reported from Minnesota. 97. Placodium cinnabarinum (Acu.) Anz. On rocks, common at the one locality. Gunflint, July 2, TO. +370. 98. Placodium citrinum (Horrn.) Lretenr. On rocks, infrequent. No. 68 well fruited but with thallus nearly obsolete in some specimens. Grand Portage island, june 21, 1697, 10.68. ‘Grand Portage, June 23, 1897, n0. 117. Misquah hills, July 5, 1897, no. 461. 99. Placodium aurantiacum (Licutr.) Narc. and Hepp. On rocks, rare at first locality, frequent at second. Grand Portage island, June 23, 1897, no. 127. Gunflint, June 30, 1897. nos. 296 and 298, and July 2, 1897, no. 389. Beaver Bay, July 13, 1897, no. 661. No. 296 with a white thallus and otherwise not typical, but I can place it nowhere else. 100. Placodium cerinum (Hrepw.) Narc. and Hepp. On trees, frequent. Grand Portage, June 23, 1897, no. 84. Guntint; June 30, 1897, no. 253, and) July: 2, 1897; mo.1392- Beaver Bay, July 13, 1897, no. 669. . Placodium cerinum (Hrepw.) Nase. and Eee var. pyra- cea Nyt. On old wood, common locally. Beaver Bay, July 13, 1897, no. 682. 256 MINNESOTA BOTANICAL STUDIES. 102. Placodium vitellinum (EurRu.) Narc. and Hepp. On old wood, frequent. Beaver Bay, July 13, 1897, no. 660. Spores simple or two-celled and reaching 30 in each ascus. 103. Placodium vitellinum (Euru.) Narc. and Hepp. var. aurellum AcH. Onrocks, common at Gunflint, rare elsewhere. Grand Port- age island, June 24, 1897, no. 163. Gunflint, June 30, 1897, no. 279, and July 1, 1897, no. 329. ‘Tofte (Carleton peak), July 10, 1897, no. 618. Snowbank lake area, July 24, 1897, no. 903. Spores reaching 20 in asci. 104. Lecanora rubina (VILL.) Acu. On rocks, common or frequent. Grand Portage (Mt. Jose- phine), June 19, 1897, no. 47. Grand Portage island, June 21, 1897, no. 73. Gunflint, June 30, 1897, no. 265. Misquah hills, July 5, 1897, no. 486. Beaver Bay, July 13, 1897, no. 705. Snowbank lake area, July 26, 1897, no. 954. 105. Lecanora rubina (ViLL.) Acu. var. heteromorpha Acnu. - On rocks, frequent locally. Grand Portage (Mt. Josephine), June 19, 1897, no. 47a. Gzunflint, July 1, 1897, no. 326. Pali- sades, July 15, 1897, no. 748. Not previously reported from Minnesota. 106. Lecanora muralis (ScHREB.) SCHAER. var. saxicola SCHAER. On rocks, common or frequent. Grand Portage island, June 21, 1897, no. 75. Rose lake, June 29, 1607, no: 7224-0 eMis— quah hills, July 5, 1897, no. 513. Tofte, July 12, 1897, no. 643. 107. Lecanora muralis (ScHREB.) SCHAER. var. diffracta Fr. On rocks, rare. Grand Portage (Mt. Josephine), June 19, 1897, no. 46. Not previously reported from Minnesota. 108. Lecanora pallida (ScHREB.) SCHAER. On trees, infrequent. Gunflint, July 1, 1897, no. 337. Not previously reported from Minnesota. 10g. Lecanora frustulosa (Dicxs.) Mass. On rocks, frequent. Grand Portage, June 21, 1897, no. 78, and June 23, 1897, no. 153. Gzunflint, July 2, 1897, no. 376. Beaver Bay, July, 13, 1897, no. 706. Not previously reported from Minnesota and new to the in- terior of North America. Fink : LICHENS OF THE LAKE SUPERIOR REGION. 207 110. Lecanora sordida (Pers.) Tu. Fr. Rocks, common at the last two locations which were high bluffs. Grand Portage, June 24, 1897, no. 185. South Fowl lake, June 26, 1897, no. 198. Misquah hills, July 5, 1897, nos. 465 and 505. Not previously reported from Minnesota and new to the in- terior of North America. 111. Lecanora subfusca (L.) Acu. On trees and rocks, common on the former. Grand Portage, June 23, 1897, no. 131. South end of South Fowl lake, June 26, 1897, no. 202. Misquah hills, July 5, 1897, no. 500. Gunflint, June 30, 1897, no. 274, and July 1, 1897, no. 305. Tofte (Carlton peak), July 10, 1897, nos. 588 and 595. Beaver Bay, July 15, 1897, no. 780. Snowbank lake area, July 19, 1897, no. 825, and July 21, 1897, no. 855. 112. Lecanora subfusca (L.) Acu. var. hypnorum ScHarr. Among moss on a cedar tree, rare. Misquah hills, July 5, 1897, no. 494a. Not previously reported from Minnesota. 113. Lecanora subfusca (L.) Acu. var. coilocarpa Ach. On trees and rocks, frequent. Grand Portage island, June 21, 1897, no. 67. Gunflint, June 30, 1897, no. 301, and July Peeogy., 10: 3309: . Misquah hills; July 95, 2807, no. 481. Beaver Bay, July 13, 1897, no. 698. 114. Lecanora varia (EuHRH.) Nyu. On rocks, rare or infrequent and some specimens perhaps approaching var. folytropa Nyl. Grand Portage, June 19, 1897, no. 41. Gunflint, July 1, 1897, no. 307, and July 2, 1897, no. 380. 115. Lecanora varia (Eurun.) Nyv. var. sepincola Fr. On wood, common at first locality. Beaver Bay, July 14, 1897, no. 742. Snowbank lake area, July 22, 1897, no. 889. Not previously reported from Minnesota. 116. Lecanora varia (Euru.) Nyu. var. symmicta Acu. On old wood, rare. Grand Portage island, June 18, 1897, no.12. Gunflint, July 2, 1897, no. 400. Tofte (Carlton peak), July ro, 1897, no. 580. 117. Lecanora hageni Acu. On rocks, rare. Gunflint, July 1, 1897, no. 357. 118. Lecanora hageni Acu. var. sambuci (Pers.) Tuck. 258 MINNESOTA BOTANICAL STUDIES. On trees common locally. Grand Portage island, June 19, 1897, no. 25. Misquah hills; July 5, 1807,;nom5ou-meorte (Carlton peak), July 10, 1897, no. 575. A puzzling plant with exciple commonly entire or excluded and looking quite as much like forms of Z. subfusca (L.) Ach. or L. varia. (Ehrh.) Nyl. Spores reaching sixteen in asci. Not previously reported from Minnesota and new to the in- terior of North America. 11g. Lecanora elatina Acu. On trees, rare... Notte, July 10, 1897, mon Gao. Not previously reported from Minnesota and new to the in- terior of North America. 120. Lecanora pallescens (L.) ScHAER. On trees, infrequent or rare, but widely distributed. Mis- quah hills, July 3, 1897, no. 412, and July 5, 1897, no. 494. Tofte (Carlton peak), July 10, 1897, no. 592. Beaver Bay, July 13, 1897, no. 674. ‘Two Harbors, July 17, 1897, no. 798. Snowbank lake area, July 21, 1897, no. 864, and July 2%, 1897, no. 973. Not previously reported from Minnesota. 121. Lecanora tartarea (L.) Acu. On rocks, rare. Ely, July 28, 1897, no. 988. Not previously reported from Minnesota. 122. Lecanora cinerea (L.) SOMMERF. On rocks, common or abundant. The thallus varying in color from ash-color to a dull black. Grand Portage (Mt. Josephine), June 19, 1897, no. 39. Grand Portage island, June 21, 1897, no. 74. Gunflint, June 30, 1897, nos. 285, 293a, 295: and 300. Misquah hills, July 5, 1897, nos. 458, 468 and 493a. Tofte (Carlton peak), July 10, 1897, nos. 615 and 622. Palisades, July 15, 1897, no. 752. Beaver Bay, July 15, 1897, no. 778. Snowbank lake area, July 20, 1897, no. 833. Ely, July 28, 1897, no. 9go. 123. Lecanora cinerea (L.) SomMeErRF. var. levata FR. On rocks, rare. - Grand Portage, June 23)9i607,umo0mo4- Gunflint, June 30, 1897, no. 286. Misquah hills, July 5, 1897, no. 471. Snowbank lake area, July 20, 1897, no. 836. 124. Lecanora cinerea (L.) SomMeERF. var. gibbosa Nyt. On rocks, rather rare. Grand Portage island, June 23, 1897, no. 79. Gunflint, July 1, 1897, no. 309a. Misquah hills, July 5, LOO7, no. 4472. Fink : LICHENS OF THE LAKE SUPERIOR REGION. 259 125. Lecanora calcarea (L.) SomMerr. (?) On rocks, rare. Thallus almost obsolete and spores only 10-1 ; 707'4 mic. Beaver Bay, July 13, 1897, no. 707. oat | Not previously reported from Minnesota. 126. Lecanora calcarea (L.) SOMMERF. var. contorta Fr. On rocks, rare. Grand Portage island, June 23, 1897, no. PS 5: Not previously reported from Minnesota. 127. Lecanora fuscata (ScHRAD.) Tu. FR. On rocks, frequent. Grand Portage (Mt. Josephine), June I9, 1897, no. 44a. Grand Portage, June 21, 1897, no. 71. Gunflint, July 1, 1897, no. 331. 128. Lecanora fuscata (ScHRAD.) Tu. Fr. var. rufescens Tu. FR. : On rocks, frequent at second locality. Misquah hills, July 5, 1897, no. 446. Beaver Bay, July 14, 1897, no. 718, and July EeeoO 7, NO. 777: Not previously reported from Minnesota. 129. Rinodina oreina (Acu.) Mass. On rocks, rare, preferring high perpendicular rocks or larger masses of talus. No. 750 is an unusually coarse form, but must be referred here. South end of South Fowl lake, June 26, 1897, no. 203. Misquah hills, July 5, 1897, no. 518. Pali- sades, July 15, 1897, nos. 750 and 753. Snowbank lake area iuby 20,.1897, no. 854. 130. Rinodina ascociscana ‘Tuck. On, tees, rare. Gunilint, July 15.1897, mo.- 340: “Notte (Carlton peak), July 10, 1897, no. 594. Not previously reported from Minnesota. 131. Rinodina sophodes (Acu.) Nyt. Abundant on drift pebbles at Beaver Bay, infrequent on wood elsewhere. Grand Portage, June 23, 1897, no. 95. Tofte, uly £2, 1697, no. O5t. Beaver Bay, July /13,.1897, nox yoo: Snowbank lake area, July 20, 1897, no. 853. 132. Rinodina sophodes (Acu.) Nyt. var. confragosa Nyu. On wood, rare. Spores 25 to 34 mic. in length. Snowbank lake area, July 21, 1897, no. 875. Not previously reported from Minnesota. 260 MINNESOTA BOTANICAL STUDIES. 133. Pertusaria velata (Turn.) Nyt. On trees, infrequent. Portage, between Rose and Rove lakes, June 27, 1897, no. 210a. Tofte (Carlton peak), July 10, 1997, no. 577. Ely, July 26519907, no. moaze 134. Pertusaria multipuncta (TurN.) Nyt. On trees, frequent at second locality. Tofte (Carlton peak), July 10, 1897, no. 611. Snowbank lake area, July 28, 1897, no. goo. 135. Pertusaria multipunctata (TurN.) Ny. var. laevigata Turn. and Borr. On trees, probably frequent. Grand Portage, June 23, 1897, no. 158. Rose lake, June 28, 1897, no. 2174, (Gumiliata ily 2, 1897, no. 389a. Misquah hills, July 5, 1897, no. 478. Not previously reported from Minnesota and new to North America. 136. Pertusaria communis DC. On trees, nearly always cedars, common or frequent. Port- age, between Rose and Rove lakes, June 27, 1897, no. 210. Rose lake, June 28, 1897, no. 217. Gunflint; Julyerenoo7. no. 336, Misquah hills, July 7, 1897, nos. 453 and 499. Beaver Bay, July 13, 1897,-no. 664, and July ors enca 7am. 782. Snowbank lake area, July 21, 1897, nos. 870 and 888. 137. Pertusaria sp. On trees. Spores nearly like the above in the few apothecia not transformed into soredia. Thallus lighter colored at cir- cumference with frequent two or three dark lines near circum- ference. Misquah hills, July 3, 1897, no. 410. Beaver Bay, July 15, 1897, no. 688. Snowbank lake area, July 19, 1897, no. 826. 138. Pertusaria leioplaca (AcH.) SCHAER. On trees, widely distributed but seldom common in any local- ity. Grand Portage, June 23, 1897, no. 91. Portage between North Fowl lake and Moose lake, June 26, 1897, no. 192. Rose lake, June 28, 1897, no. 218. Gunflint, Jume goneay. no. 275, and July 2, 1897, no. 399. Misquah) sills simives. 1897; no. 491a. Beaver Bay, July 15, 1807, no. 7e4eueae July 28, 1897, no. 1010. Varying greatly according to sub- stratum. On young trees with smooth bark the thallus is thin and smooth and the ostioles frequently indistinct. On older trees with rough bark the thallus is thicker and broken, and Fink : LICHENS OF THE LAKE SUPERIOR REGION. 261 the apothecia are falsely lecanoroid. The last feature is due no doubt to great age of these plants which began growth when the trees were young. The extremes appear macroscopically like distinct species. Not previously reported from Minnesota. 139. Pertusaria pustulata (Acu.) Nyv. On trees, infrequent. Rose lake, June 28, 1897, no. 228. Gunflint, July 1, 1897, no. 310. Misquah hills, July 3, 1897, no. 418. 140. Pertusaria glomerata (AcuH.) ScHAER. On rocks, very rare. Misquah hills, July 5, 1897, no. 490. Not previously reported from Minnesota and new to the in- terior of North America. 141. Gyalecta fagicola (HEpp.) Tuck. On trees, rare. Snowbank lake area, July 22, 1897, no. 880. Not previously reported from Minnesota and new to west of of New England. 142. Urceolaria scruposa (L.) Nyt. On rocks, rare or infrequent, but widely distributed and no. 126 with thallus approaching var. gypsacea Nyl. Grand Port- age island, June 25, 1897, no. 126. Gzunflint, June 30, 1897, no. 302. Misquah hills, July 5, 1897, no. 473. Beaver Bay, July 13, 1897, no. 7o1.. Snowbank lake area, July 23, 1897, no. 899. Ely, July 28, 1897, no. 978. 143. Stereocaulon coralloides Fr. On rocks, frequent. Tofte (Carlton peak), July 10, 1897. no. 549: Not previously reported from Minnesota and new to the in- terior of North America. 144. Stereocaulon paschale (L.) Fr. On earth among rocks, common. Grand Portage (Mt. Josephine), June 19, 1897, no. 49. Grand Portage island, June 21, 1897, no. 50. Gunflint, June 30, 1897, no. 233. Misquah hills,*July 5, 1897, no. 482. Beaver Bay, July 13, 1897, no. 689. Palisades, July 15, 1897, no. 758. Snowbank lake area, July 24, 1897, no. 927. No. 49 seems to approach the above. Also no. 50 has the stout podetia of S. tomentosum (Fr.) Th. Fr. and is somewhat tomentose. Yet it appears nearer herb. specimens of the above. 262 MINNESOTA BOTANICAL STUDIES. 145. Cladonia symphycarpa FR. var. epiphylla (Acu.) Nyt. In crevices in rocks, rare. Habitat unusual but thallus too large for C. cespiticia (Pers.) Fl. Gunflint, July t, 1897, no. 363. Not previously reported from Minnesota. 146. Cladonia mitrula Tuck. On earth, rare. Beaver Bay, July 15, 1897, no. 694. 147. Cladonia cariosa (AcH.) SPRENG. On earth, probably frequent. Grand Portage island, June 23, 1897, nos. 120 and 146. Gunflint, July) te07,.ne1a2 Misquah hills, July 5, 1897, nos. 509 and 533. Tofte (Carl- ton peak), July 10, 1897, no. 606. Beaver Bay, July 14, 1897, NOS..7/3/7 2nd 7.39: The forms listed here seem to me to be partly intermediate between this and the last having the habit of this, but some are rather small with the squamules usually small. C. mztruda Tuck. is the common form in southern Minnesota, but the better de- velopment of this region runs into the present species. Of the specimens here listed no. 146 is the best representative of the species and no. 120 the poorest. Some of the smaller ap- proaches C. symphycarpa Fr., which is itself a doubtful species. 148. Cladonia decorticata FLoERK. On earth, rare. Beaver Bay, July 14, 1897, no. 738. Not previously reported from Minnesota and new to west of New England. 149. Cladonia pyxidata (L.) Fr. . On earth, common. Grand Portage island, June 19, 1897, no. 32. Gunflint, July 1, 1897, nos. 333 and 352. Misquah hills, July 5, 1897, no. 534. Tofte (Carlton peak), July 10, 1897, nos. 567 and 568. Beaver Bay, July 14, 1897, no. 735. Snowbank lake area, July 21, 1897, no. 868, and. July 27, 1807, NO. 972. 150. Cladonia fimbriata (L.) Fr. On earth, rare. Gunflint, June 30, 1897, no. 259. Not previously reported from Minnesota. 151. Cladonia fimbriata (L.) Fr. var. tubeformié Fr. On dead wood and earth, common. Grand Portage island, June 23, 1897, no; 129. Gunflint, June 30, 1807;0nomeoe Misquah hills, July 3, 1897, no. 439. Tofte (Carlton peak), July 10, 1897, no. 561. Snowbank lake area, July 24, 1897, no. 930. Fink: LICHENS OF THE LAKE SUPERIOR REGION. 268 152. Cladonia fimbriata (L.) Fr. var. radiata Fr. On earth and old wood, frequent. Grand Portage island, June 19, 1897, no. 33 and June 24, 1897, no. 162. Gunflint, June 30, 1897, no. 234. Snowbank lake area, July 20, 1897, no. 840. Not previously reported from Minnesota. 153. Cladonia gracilis (L.) Nyw. On earth, common or abundant and extremely variable. Grand Portage island, June 19, 1897, no. 35. Gunflint, July I, 1897, nos. 325, 344 and 345. Misquah hills, July 5, 1897, nos. 508 and 522. Tofte (Carlton peak), July 10, 1897, nos. 550, 564 and 604. Beaver Bay, July 14, 1897, no. 734. Snow- bank lake area, July 21, 1897, no 876 and July 24, 1897, nos. 922 and 923. 154. Cladonia gracilis (L.) Nyv. var. verticillata Fr. On earth, rare, Grand Portage island, June 19, 1897, nos. 35b and 35d. ‘Tofte (Carlton peak), July 10, 1897, no. 625. 155. Cladonia gracilis (L.) Nyu. var. symphycarpia Tuck. On earth, infrequent, possibly as near C. degenerans Floerk. Grand Portage island, June 19, 1897, no. 35a. Not previously reported from Minnesota. 156. Cladonia gracilis (L.) Ny. var. cervicornis FLoERK. On earth, rare. Gunflint, June 30, 1897, no. 231. Not previously reported from Minnesota. 157. Cladonia gracilis (L.) Nyv. var. hybrida ScHAEr. On earth, common. Grand Portage island, June 19, 1897, no. 35c. Gunflint, July 1, 1897, no. 346. Misquah hills, July 3, 1897, no. 433. Snowbank lake area, July 19, 1897, no. 821, and July 24, 1897, no. 905. 158. Cladonia turgida (Enru.) Horr. On earth, common at Gunflint. Gunflint, June 30, 1897, nos. 241 and 252. Snowbank lake area, July 24, 1897, no. 928. Not previously reported from Minnesota. 159. Cladonia turgida (Enru.) HorrM.var. conspicua (SCHAER.) INVE: On earth, frequent... Kose lake, June 28, 1897,' no. 237- Misquah hills, July 3, -1897, no. 425, and July 5, 1897, no. 525. Tofte, (Carlton peak), July 10, 1897, nos. 603 and 637. Not previously reported from Minnesota. 264 MINNESOTA BOTANICAL STUDIES. 160. Cladonia squamosa Horrn. On earth, common or abundant. Grand Portage island, June 24, 1897, no. 165. Gunflint, June 30, 1897, no. 232, and July 1, 1897, no. 350, and July 2, 1897, no. 384. Misquah hills, July 5, 1897, no. 529. Tofte, July 12, 1897, nos. 628 and 632. Beaver Bay, July 14, 1897, no. 719. Above Pali- sades, July 15, 1897, no. 772. Snowbank lake area, July 24, 1897, nos. 913, 919 and 924. Ely, July 28, 1897, no. 995. 161. Cladonia squamosa Horr. var. phyllocoma RABENH. On earth, frequent. Grand Portage island, June 23, 1897, no. 141. Misquah hills, July 5, 1897, no. 459. Snowbank lake area, July 24, 1897, nos. 939 and 942. Not previously reported from Minnesota and new to North America. 162. Cladonia cornuta (L.) Fr. On earth, rare. Grand Portage island, June 24, 1897, no. 159. Misquah hills, July 5, 1897, no. 511. Not previously reported from Minnesota. 163. Cladonia delicata (Euru.) FL. On old wood, rare. Beaver Bay, July 13, 1897, no. 692. 164. Cladonia cespiticia (PERs.) FL. On old wood, rare. Tofte (Carlton peak), July 10, 1897, no. 586. 165. Cladonia furcata (Hups.) Fr. On earth, frequent. Gunflint, June 30, 1897, nos. 236 and 237. Misquah hills, July 5, 1897, no. 524. Above Palisades, July 15, 1897, no, 767. Snowbank lake area, July 24, 1897, no. 914. 166. Cladonia furcata (Hups.) Fr. var. crispata FL. On earth, common locally. Grand Portage, June 24, 1897, no. 168. Gunflint, June 30, 1897, no. 249. Palisades, July [55 LOO], MOn 757. 167. Cladonia rangiferina (L.) Horr. On earth, abundant or common. Grand Portage island, June 17, 1897, no. 2, and June 23, 1897, 135. Grand Portage (Mt. Josephine), June 19, 1897, no. 48. Gunflint, June 30, 1897, no. 244. Misquah hills, July 5, 1897, no./521. 9 Botte, (Carlton peak), July 10, 1897, no. 548. Beaver Bay, July 14, 1897, no. 732. Palisades, July 15, 1897, no. 759. Snowbank lake area, July 24, 1897, no. 926. ol Fink ;» LICHENS OF THE LAKE SUPERIOR REGION. 26! 168. Cladonia rangiferina (L.) Horr. var. sylvatica L. On earth, frequent. Misquah hills, July 5, 1897, no. 531. Tofte (Carlton peak), July 10, 1897, no. 553. Snowbank lake area, July 24, 1897, nos. 915 and 936. 169. Cladonia rangiferina (L.) Horr. var. alpestris L. On earth, common locally. Grand Portage, June 24, 1897, no,171. Gunflint, July 1, 1897, no. 351. Above Palisades, ily i5, 1897, no. 773. 170. Cladonia amaurocrea (FL.) ScHAER. On earth and rocks, common or frequent. Grand Portage (Mt. Josephine), June 21, 1897, no. 52. Grand Portage island, June 24, 1897, no. 164. Gunflint, June 30, 1897, nO. 243. Misquah hills, July, 5, 1897, nos. 530 and 532. Tofte (Carl- ton peak), July, 10, 1897, no. 552. Palisades, July 15, 1897, no. 769. Snowbank lake area, July 24, 1897, nos. 935 and OTe: Not previously reported from Minnesota. 171. Cladonia uncialis (L.) Fr. On earth, frequent or common. Grand Portage (Mt. Jose- ping), June 27, 1897, no. 55. Grand Portage, June 24, 1897, no.167. Gunflint, June 30, 1897, no. 239. Misquah hills, July 5, 1897, no. 475. Misquah hills, July 5, 1897, no. 528. Tofte (Carlton peak), July 10, 1897, no. 551. Beaver Bay, July 14, 1897, no. 727. Palisades, July 15, 1897, no. 764. Snowbank lake area, July 24, 1897, no. 933. 172. Cladonia cornucopioides (L.) Fr. On earth, rare but widely distributed. Grand Portage, June 24, 1897, no. 166. Gunflint, July 30, 1897, no. 238. Misquah hills, July 5, 1897, nos. 497 and 527. Tofte (Carlton peak), July 10, 1897, no. 563. Above Palisades, July 15, 1897, no. 754: = 173. Cladonia deformis (L.) Horr. On earth, rare. Grand Portage island, June 23, 1897, no. EGt. Not previously reported from Minnesota and new to the in- terior of North America. 174. Cladonia digitata (L.) Horr. On an old stump, rare. Tofte, July 12, 1897, no. 655. Not previously reported from Minnesota and new to the in- terior of North America. 266 MINNESOTA BOTANICAL STUDIES. 175. Cladonia macilenta (Euru.) Horr. On old wood and earth, rare. Misquah hills, July 5, 1897, no. 510. Above Palisades, July 15, 1897, no. 776. 176. Cladonia cristatella Tuck. On earth and old wood, abundant. Grand Portage island, June 18, 1897, no. 31. Gunflint, July 1, 1897, nos. 322 and 332. Misquah hills, July 3, 1897, no. 417. Tofte (Carlton peak), July 10, 1897, no. 610. Beaver Bay, July 14, 1897, no. 736. Snowbank lake area, July 19, 1897, no. 827. 177. Beomyces byssoides (L.) ScHArEr. On rocks, rare. Grand Portage island, June 21, 1897, no. 109. Not previously reported from Minnesota, and new to the in- terior of North America. 178. Beomyces eruginosus (Scop.) DC. On rotton wood, common. Grand Portage island, June 18, 1897, no. 16. Gunflint, July 1, 1897, no. 309. Misquah hills, July 3,.1897, no. 404. Tofte, July 12, 1897; no. 63g; beaver Bay, July 14, 1897, no. 721. Snowbank lake area, July 19, 1897, no. 823. Not previously reported from Minnesota, and new to the in- terior of North America. 179. Biatora rufonigra Tuck. On rocks, frequent. Grand Portage (Mt. Josephine), June 19, 1897, no. 37. Grand Portage island, June 21, 1897, no. 64. South Fowl lake, June 26, 1897, no. 193. Misquah hills, July 5, 1897, no. 456. Snowbank lake area, July 21, 1897, no. 859. 180. Biatora coarctata (Sm., Ny.) Tuck. On rocks, rare. Beaver Bay, July 13, 1897, no. 709. Not previously reported from Minnesota. 181. Biatora viridescens (SCHRAD.) FR. On old wood, common locally. Misquah hills, July 3, 1897, NO AEA Snowbank lake area, July 20, 1897, no. 846. Not previously reported from Minnesota, and new to the in- terior of North America. 182. Biatora vernalis (L.) FR. On old wood, mosses and trees, frequent. Grand Portage Fink: LICHENS OF THE LAKE SUPERIOR REGION. 267 island, June 21, 1897, no. 61, and June 23, 1897, no. 149. Gunflint, June 30, 1897, no. 262. Not previously reported from Minnesota. 183. Biatora sanguineoatra (F'r.) Tuck. On earth common. Grand Portage island, June 19, 1897, no. 21 and 28. Grand Portage, June 24, 1897, no. 181. Gun- flint, June 30, 1897, no. 269, and July 1, 1897, no. 359. Mis- quah hills, July 5, 1897, no. 413. Tofte (Carlton peak), July 10, 1897, no. 578. Snowbank lake area, July 22, 1897, no. 884. Not previously reported from Minnesota. 184. Biatora turgidula (Fr.) Nyt. On old wood, rare. Grand Portage island, June 18, 1897, no. 14. Not previously reported from Minnesota. 185. Biatora leucophea (FLOERK).” On rocks, infrequent. Grand Portage, June 23, 1897, no. 139, and June 24, 1897, no. 187. Gunflint, July 1, 1897, no. 306. Not previously reported from Minnesota. 186. Biatora leucophea FLorrK var. griseoatra Korrs. On rocks, rare. Grand Portage, June 24, 1897, no. 186. Not previously reported from Minnesota and new south of Arctic America. 187. Biatora uliginosa (ScHRAD.) Fr. On earth, abundant near Disappointment lake. Beaver Bay, July 13, 1897, no. 708. Snowbank lake area, July 24, 1897, no. 944. Ely, July 28, 1897, no. 997. Not previously reported from Minnesota. 188. Biatora spheroides (Dicxs.) Tuck. On old wood, rare. Snowbank lake area, July 24, 1897, no. oii ae = Not previously reported from Minnesota. 189. Biatora glauconigrans Tuck. | On trees, rare. Gunflint, July 1, 1897, no. 398. Not previously reported from Minnesota and new west of New England. 190. Biatora arthropurpurea (Mass.) Hepp. On trees, rare. Grand Portage island, June 21, 1897, no. 56. Snowbank lake area, July 22, 1897, no. 811. Ely, July 28, 1897, no. 1007. 268 MINNESOTA BOTANICAL STUDIES. 191. Biatora oxyspora (Tuu.) Nyv. On Parmelia colpodes, rare. Misquah hills, July 3, 1897, no. 416. Not previously reported from Minnesota and _ new to the in- terior of North America. 192. Biatora lucida (Acu.) Fr. On damp rocks, rare. Grand Portage, june 23, 1887, no. 183. Not previously reported from Minnesota and new to the in- terior of North America. 193. Biatora myriocarpoides (Nyv.) Tuck. On old wood, frequent locally. Beaver Bay, July 13, 1897, no. 673. Apothecia larger than usual. 194. Biatora flavidolivens Tuck. On old wood, frequent locally. Rose lake, June 28, 1897, no. 214. Misquah hills, July 5, 1897, no. 504. Not previously reported from Minnesota and new west of New England. 195. Biatora hypnophila (Turn.) Tuck. On trees, rare. Gunflint, June 30, 1897, no. 273. Snow- bank lake area, July 26, 1897, no. 956. 196. Biatora negelii Herr. On old wood, rare. Beaver Bay, July 13, 1897, no. 715. Not previously reported from Minnesota and new west of New England. . 197. Biatora rubella (EHRH.) RABENH. On trees, rare or infrequent. ‘Tofte (Carlton peak), July 10, 1897, no. 546. Beaver Bay, July 13, 1897, no. 640. Snow- bank lake area, July 19, 1897, no. 812, July 21, 1897, no. 872, and July 24, 1897, no. 906. Ely, July 28, 1897, no. 1021. 198. Biatora fuscorubella (Horrm.) Tuck. On trees, infrequent. Snowbank lake area, July 17, 1897, no. 816, and July 21, 1897, no. 861. 199. Biatora schweinitzii Fr. On cedars, rare. Misquah hills, July 5, 1897, no. 493. Not previously reported from Minnesota. 200. Biatora incompta (Borr.) HEpp. On trees, probably common locally. Rose lake, June 28, 1897, no. 222. ,Guntlint, July 1, 1897, no. 335, Not previously reported from Minnesota. Fink: LICHENS OF THE LAKE SUPERIOR REGION. 269 201. Biatora muscorum (Sw.) Tuck. On trees with moss, rare. Snowbank lake area, July 26, 1897, no. 963. 202. Heterothecium sanguinarium (L.) For. On old wood and occasionally on trees or rocks, common ex- cept in last region. Gunflint, July 1, 1897, nos. 318 and 319. Misquah hills, July 3, 1897, nos. 409 and 415, and July 5, 1897, nos. 492a and 498. Tofte (Carlton peak), July 10, 1897, no. 581. Beaver Bay, July 15, 1897, no. 695. Two Harbors, July 17, 1897, no. 796. Snowbank lake area, July 19, 1897, no. 810. Not previously reported from Minnesota and new to the in- terior of North America. 203. Heterothecium sanguinarium (L.) Fvor. var. affine Tuck. On wood, rare. Rose lake, June 28, 1897, no. 222a. Not previously reported from Minnesota and new to the in- terior of North America. 204. Lecidea lactea FL. On rocks along lake Superior, common especially north. Grand Portage island, June 21, 1897, no. 76, and June 23, 1897, no. 136. Grand Portage, June 23, 1897, no. 182. Tofte, July 12,1897, no. 652. Palisades, July 15, 1897, no. 751. Not previously reported from Minnesota and new to the in- terior of North America. 205. Lecidea crustulata Acu. On rocks, rare. Grand Portage island, June 23, 1897, no. 137. Not previously reported from Minnesota and known else- where in North America only from Labrador by Eckfeldt and Arnold. 206. Lecidea lapicida Fr. On rocks, probably common locally. Misquah hills, July 5, 1897, no. 448. Not previously reported from Minnesota and new to the interior of North America. 207. Lecidea lapicida Fr. var. oxydata Fr. On rocks, rare. Grand Portage, June 24, 1897, no. 174. Not previously reported from Minnesota and new to the in- terior of North America. 270 MINNESOTA BOTANICAL STUDIES. 208. Lecidea speirea Nyt. On rocks along lake shore, rare. Grand Portage island, umes 235 1607, 10. 110. Not previously reported from Minnesota. 209. Lecidea albocerulescens (WuLF.) SCHAER. On rocks, rare. Misquah hills, July 5, 1897, no. 484. 210. Lecidea platycarpa Acu. On rocks, rare. Misquah hills, July 5, 1897, no. 474. Not previously reported from Minnesota. 211. Lecidea enteroleuca Fr. On trees and rocks, common. Grand Portage island, June 21, 1897, nos. 59b and 60, and June 23, 18947, no. 111. Grand Portage, June 23, 1897, no. 148. English Portage) june 20, 1897, no. Ion. Misquah hills, July 55) 18077 emomeag2- Beaver Bay, July 13, 1897, no. 700. Snowbank lake area, July 24, 1897; no. 909. 212. Lecidea enteroleuca FR. var. achrists SOMMERF. On trees, infrequent. Grand Portage island, June 21, 1897, no. 59a. Grand Portage, June 23; 1307, nomaye Not previously reported from Minnesota. 213. Lecidea melancheima Tuck. On old wood, rare. Gunflint, July 2, 1897, no. 390. Mis- quah hills, July 5, 1897, no. 454. Snowbank lake area, July 20, 1897, no. 835. Not previously reported from Minnesota. 214. Lecidea cyrtidia ‘Tuck. On pebbles, rare, thallus reduced and hence the black hypo- thallus prominent. Snowbank lake area, July 27, 1897, no. 969. Not previously reported from Minnesota. 215. Lecidea acclinis Fior. On cedars, rare. Gunflint, July 1, 18975 movas5ey Not previously reported from Minnesota. 216. Buellia alboatra (Horrno.) Tu. Fr. On rocks, rare. Grand Portage, June 23, 1897, no. 9g. Not previously reported from Minnesota. 217. Buellia parasema (Acu.) Tu. FR. On trees, common. Ely specimen having spores reaching — = mic. Grand Portage island, June 16, 1897, no. 4, and ' Fink: LICHENS OF THE LAKE SUPERIOR REGION. PT a June 21, 1897, no. 65. South Fowl lake, June 26, 1897, no. 196. Gunflint, July 1, 1897, nos. 311 and 312. Misquah hills, July 5, 1897, no. 480. Beaver Bay, July 15, 1897, no. 676. Snowbank lake area, July 19, 1897, nos. 811a, 813 and 817. Ely, July 28, 1897, no. 98ga. 218. Buellia parasema (Acn.) Tu. FR. var. triphragmia Nyv. On trees, infrequent. Gunflint, June 30, 1897, nos. 255 and 27olotte, July 10, 197, no. 613. Not previously reported from Minnesota. 219. Buellia dialyta (Nyv.) Tuck. On pines, rare. Two Harbors, July 17, 1897, no. 792. Not previously reported from Minnesota, and new to the in- terior of North America. 220. Buellia myriocarpa (DC.) Mupp. On old wood, abundant locally. Beaver Bay, July 14, 1897, no. 716a. 221. Buellia myriocarpa(DC.) Mupp. var. polyspora WiLLEyY. On ittees, rare. “Ely, July 28, 1997, no. 1011. 222. Buellia petrea (FLotr., Korrs.) Tuck. On rocks, common or abundant. Grand Portage island, mie r7, 1597, no. 5. “Gunilint, June’ 30), 1897; nos 281, 283 and 291. Misquah hills, July 5, 1897, no. 470. Beaver Bay, July 15, 1897, no. 779. Ely, July 28, 1897, no. 989. 223. Buellia petrea (Fior., Korrs.) Tuck. var. grandis FLOERK. On rocks, rare. Gunflint, June 30, 1897, no. 293, and July ROO], Or 304. 224. Buellia petrea (FLor., Korers.) Tuck. var. montagnei Tuck. On rocks, common or abundant. Grand Portage (Mt. Jose- phine), June 18,-1897, no. 42. Gunflint, July 2, 1897, no. 379. Misquah hills, July 5, 1897, no. 467. Beaver Bay, July 13, 1897, no. 662. Palisades, July 15, 1897, no. 749. Snowbank lake area, July 20, 1897, no. 834. 225. Buellia geographica (L.) Tuck. On rocks, rare and approaching var. /ecanorina Floerk Gunflint, June 30, 1897, no. 303. Palisades, July 15, 1897, no. 743: Not previously reported from Minnesota, and new to the in- terior of North America. I (P MINNESOTA BOTANICAL STUDIES. 226. Buellia parmeliarum (SoMMERF.) ‘Fuck. On Parmelia borrert, rare. Snowbank lake area, July 22, 1897, no. 885. Not previously reported from Minnesota. 227. Opegrapha varia (Pers.) Fr. On trees, common on cedars except along lake Superior. Gunflint, July 1, 1897, no. 334. Misquah hills, July 5, 1897, no. 502. ‘Tofte (Carlton peak), July 10, 1897, no. 593. Snow- bank lake area, July 21, 1897, no. 857. 228. Opegrapha varia (Pers.) Tuck. Fr. var. notha Acu. On cedars, locally abundant. Rose lake, June 28, 1897, no. 220. Not previously reported from Minnesota. 229. Graphis scripta (L.) Acn. On trees, frequent or common. Grand Portage island, June 21, 1897, no. 66. Gunflint, July 1, 1807, mnosauaiaeameanye Tofte (Carlton peak), July 10, 1897, no. 576. Beaver Bay, July 15, 1897, no. 716. Snowbank lake area, July 21, 1897, no. 858. 230. Graphis scripta (L.) Acu. var. recta (Hums.) Nyt. On birch trees, infrequent. Grand Portage, June 24, 1897, no. 189. Misquah hills, July 3, 1897, no. 405. Tofte, July 12, 1897, no. 650. Snowbank lake area, July 23, 1897, no. 897. 231. Graphis scripta (L.) Acu. var. limitata Acu. On trees, very rare. Resembles G. dendritica externally as to apothecia. Misquah hills, July 5, 1897, no. 451. Not previously reported from Minnesota. 232. Arthonia dispersa (ScuRAD.) Nyt. On Acer spicatum, abundant. Grand Portage, June 23, 1897, no. 86. Tofte (Carlton peak), July 10, 1897, no. §90. Beaver Bay, July 13, 1897, no. 714. Snowbank lake area, July 20, 1897, no. 845. Ely, July 28, 1897, no. 979. Not previously reported from Minnesota. 233. Arthonia radiata (Pers.) Tu. Fr. On trees in low places, common. Grand Portage island, June 23, 1897, ne. 123. Rose lake, June 26, 9ico7;mmoueeue: Gunflint, July 1, 1897, no. 315, and July 2, 1607, "moo: Tofte (Carlton peak), July 10, 1897, no. 584, Beaver Bay, Fink ; LICHENS OF THE LAKE SUPERIOR REGION. Zhe July 15, 1897, no. 785. Snowbank lake area, July 20, 1897, moose. Ely, July 26, 1897, no. 101g. 234. Arthonia punctiformis Acu. On trees, locally common. Gunflint, June 30, 1897, no. 277. Misquah hills, July 3, 1897, no. 407. 235. Arthonia patellulata Nyt. On trees, rare. Gunflint, June 30, 1897, no. 254. Not previously reported from Minnesota. 236. Calicium trichiale Acu. On trees, common locally. Rose lake, June 28, 1897, no. 230. Beaver Bay, July 15, 1897, no. 696. Snowbank lake area, July 19, 1897, no. 818. Not previously. reported from Minnesota, and new to the in- terior of North America. 237. Calicium trichiale Acu. var. stemoneum Nyt. On pine, common. Ely, July 28, 1897, no. 992. Not previously reported from Minnesota, and new to the in- terior of North America. 238. Calicium brunneolum Acu. On decorticated wood, common locally. Two Harbors, July 17, 1897, no. 800. Snowbank lake area, July 12, 1897, no. 860. Ely, July 26, 1897, no. 1000. Not previously reported from Minnesota, and new to the in- terior of North America. 239. Calicium chrysocephalum (Turn.) Acu. On trees, frequent. Misquah hills, July 5, 1897, no. 447. Two Harbors, July 17, 1897, no. 788. Snowbank lake area, July 22, 1897, no. 882. Ely, July 28, 1897, no: 1003. Not previously reported from Minnesota, and new to the in- terior of North America. 240. Calicium chrysocephalum (Turn.) Acn. var. filare Scu. On cedars, rare. Tofte, July 12, 1897, no. 647. Not previously reported from Minnesota. Variety apparently new to North America. 241. Calicium parietinum Acun. On decorticated wood, common. Grand Portage island, June fp oo7, no. 3: Gunilint; July 2, 1897, no. 382. iMisquah hills, July 3, 1897, no. 408. Not previously reported from Minnesota. 274 MINNESOTA BOTANICAL STUDIES. 242. Calicium quercinum Pers. On dead wood, infrequent. Rose lake, June 28, 1897, no. 229. Tofte (Carlton peak), July 10, 1897, no 583. Not previously reported from Minnesota. 243. Calicium hyprellum Acu. var. viride Nyv. On trees, rare. Misquah hills, July 5, 1897, no. 476. Snow- bank lake area, July 21, 1897, no. 877, and July 22, 1897, no. 893. Ely, July 28, 1807, no, 1015: Not previously reported from Minnesota. Variety new to North America. Stipes sometimes very short. 244. Calicium turbinatum Pers. On Pertusaria communts, rare. Beaver Bay, July 13, 1897,. no. 664a. Snowbank lake area, July 21, 1897, no. 866. Not previously reported trom Minnesota. 245. Coniocybe pallida (PERs.) FR. On Fraxinus, rare. Snowbank lake area, July 19, 1897, no. 831. Not previously reported from Minnesota. 246. Endocarpon miniatum (L.) ScHAER. On rocks, along shore of lake Superior, very rare. Grand Portage island, June 21, 1897, no. 80. 247. Endocarpon miniatum (L.) ScHAER. var. complicatum SCHAER. On rocks, frequently 1000 feet above water level, frequent. Grand Portage (Mt. Josephine), June 19, 1897, no. 38. Grand Portage island, June 23, 1897, no 102. Misquah hills, July 5, 1897, no. 445. 248. Endocarpon fluviatile DC. On rocks frequently inundated, common. Rose lake, June 28, 1897, no. 211. Gunflint, July 1, 1897, no. 327. Misquah hills, July 7, 1897, no. 512. Snowbank lake area, July 21, 1897, no. 878. 249. Thelocarpon prasinellum Nyv. On rocks, rare. Grand Portage (Mt. Josephine), June 22, 1597, no. go. The plant agrees here and not with saxicoline species, Euro- pean or American. 250. Staurothele umbrina (Wanv.) Tuck. Wet rocks, common. Misquah hills, July 5, 1897, nos. Fink: LICHENS OF THE LAKE SUPERIOR REGION 275 462, 519 and 520. Snowbank lake area, July 26, 1897, no. 957: Not previously reported from Minnesota. 251. Staurothele drummondii Tuck. On rocks along the shore, frequent locally. Grand Portage iskand, ‘June 21, 1897, no. 72. Not previously reported from Minnesota and new to the in- terior of North America. 252. Verrucaria nigrescens Pers. On rocks, rare. Grand Portage, June 24, 1897, no. 184. 253. Verrucaria epigea (Prrs.) Acn. On earth, rare. Snowbank lake area, July 26, 1897, no. oar: Not previously reported from Minnesota. 254. Sagedia oxyspora (Nyv.) Tuck. Omebirch, care. Beaver Bay, July £3. 1897, no. 607. Not previously reported from Minnesota and new to the in- yerior of North America. 255. Pyrenula punctiformis (Acu.) NaArc. var. fallax Nyt. On trees, common. Gunflint, June 30, 1897, no. 276. Mis- quah hills, July 5, 1897, no. 503. Snowbank lake area, July 21, 1897, no. 871, and July 26, 1897, no. 950. Not previously reported from Minnesota. 256. Pyrenula leucoplaca (WALLR.) Krs. On trees, common to west of region. Between Rose and Rove lakes, June 27, 1897, no. 209. Gunflint, July 2, 1897, no. 391. Misquah hills, July 5, 1897, nos. 499 and 507. Snowbank lake area, July 19, 1897, no. 828, July 20, 1897, no. 852, and July 26, 1897, nos. 955, 958 and 959. _ Ely, July 28, 1897, nos. 100g and 101g. 257. Pyrenula cinerella (FLotT. Tuck. On birch, common. Grand Portage island, June 21, 1897, no. 56. The only American specimens seen by me which show the spores as large as those of the European plant. Spores meas- ured 12-18 by 6-g mic. Spore measurements for the species in America are more commonly 12-17 by 5-7 mic., my Iowa specimens giving 12-16 by 6-7 mic., and T. A. William’s from Nebraska, 15-17 by 5%4-7 mic. Not previously reported from Minnesota. 276 MINNESOTA BOTANICAL STUDIES. 258. Pyrenula cinerella (FLot.) Tuck. var. quadriloculata, var. nov. Spores 12-15 by 5-6% mic., passing from 2 and occasion- ally 3-celled to a much more common 4-celled condition. The apothecia somewhat below normal size for the species. Pyren- ula punctiformis Ach., Naeg. var. fallax Nyl., quite commonly occurs with the species and variety, as it does with the latter in the present instance and with the former both in Minnesota and Iowa. On birch, probably common locally. Grand Portage island, June 24, 1397, no: 35. XIX. CONTRIBUTIONS TO A KNOWLEDGE OF THE LICHENS OF MINNESOTA.—V. LICHENS OF THE MINNESOTA VALLEY AND SOUTHWESTERN MINNESOTA. BruceE FInxK. CONSIDERATIONS OF DISTRIBUTION AND HABITAT. The area considered in this paper was selected with a view to obtaining as complete a knowledge as possible of the lichen flora of the Minnesota river valley and of that of southwestern Minnesota in general. The upper portion of the valley near Minneapolis would, of course, give a flora essentially like that of Minneapolis and vi- cinity already studied. Hence, for the month’s field work, it was thought best to begin operations at a locality a considerable distance from Minneapolis. As an initial place, Mankato, about 60 miles from Minneapolis, was selected. The location of this city is also advantageous in that it lies nearly midway between the Minneapolis and the northeastern Iowa areas compared carefully in the second paper of this series, thus forming a con- necting link between the two areas previously studied. After a careful study of the lichens of the Mankato area both to gain a knowledge of the lichen flora of the region and for the sake of relationships with the areas indicated above, New Ulm was next selected as an area of special interest because of the expo- sures of Cretaceous sandstone and the most southeastward ex- posures of quartzite rocks in the valley. At New Ulm only these two rock formations were studied, as time spent on other substrata present would only be repaid for most part by a repe- tition of the species found upon the same substrata at Mankato, only 30 miles distant. Three days were next spent at Red- wood Falls, Morton and North Redwood with a view to secur- ing rare species and noting the southeastern extension of certain species in the valley. From here I proceeded to Granite Falls. 277 278 MINNESOTA BOTANICAL STUDIES. This being the most northwestern area reached in the survey, its lichen flora was studied carefully. The final task was to study the lichen flora of the pipestone and the Sioux quartzite at Pipestone. A brief statement as to substrata is next in order. About Mankato trees abound, and three kinds of rock—limestone, sandstone and bowlders—are plentiful. I found only the two interesting substrata mentioned above at New Ulm. ‘Trees and bowlders were abundant, but were not studied for the reason al- ready stated. At Redwood Falls, Morton and North Redwood, granite trees and earth were examined for species especially rare or interesting. The great masses of granite, supposed to have been exposed since the close of the glacial age, formed the most interesting substratum at Granite Falls. This is also the most northwestern area in the valley where trees occur in any considerable numbers. The calcareous drift pebbles and cal- careous earth proved also very interesting here. The two sub- strata examined at Pipestone have been mentioned. I need to add only one statement more to make the analysis of substrata complete enough for the present purpose. ‘This is that earth — was examined everywhere and furnished much of interest, as will appear later. The following rare lichens were found only at Redwood Falls, Morton or North Redwood: VPeltigera cantina (L.) Hoffm. var. spongiosa Tuck. and Stereocaulon paschale (L.) Fr. Also the area including the above places forms the most southeastern known extension of the following lichens in the valley: Parmelia olivacea (li.) Ach. var. prolixa Ach.; Pan- naria microphylla (Sw.) Delis; Omphalaria phyllisca (Wahl.) Tuck.; Lecanora frustulosa (Dicks.) Mass., and Buellia pul- lata Tuck. With this much in hasty review I shall pass to lo- calities more thoroughly studied. However, I may add here better than elsewhere in my paper that /t7nodina oreina (Ach.) Mass. and Lecanora xanthophana Nyl. are here and elsewhere in the valley far more abundant than I have ever found them in other regions. In attempting a general comparative study of distribution in the valley the places that present questions of greatest interest are the vicinities of Mankato and Granite Falls, where all sorts of sub- strata were examined. ‘The two areas were about equally well studied, though the former, because of the greater number and Fink: LICHENS OF THE MINNESOTA VALLEY. 279 less accessibility of rocky substrata, required more time. The former area furnished 151 species and varieties and the latter 124. A brief analysis of the causes of the advantage in favor of the former region can be best made by a consideration of the sub- joined table, giving the various substrata for both localities with the number of lichens most commonly found on each. Numbers for Mankato. For Granite Falls. Trees 60 41 Rocks 55 54 Earth 22 17 Dead wood 14 12 A complete analysis introducing per cents as was made ina former paper is not necessary since general likeness except for trees is apparent inthe table. The difference in richness then is due mainly to absence of large areas of trees at Granite Falls. The slight differences in the other three items in the table is doubtless due to difference in moisture, the precipitation -being 30.53 inches annually at Mankato for three years for which I could get data and 21.83 inches annually at Granite Falls for five years for which data were obtained. Difference in moisture doubtless also accounts in small measure for the ad- vantage of the Mankato area as to arboreal lichens. As to rocky substrata favorable to lichen growth little can be definitely given by way of comparison. As to kinds of rocks Mankato has an advantage in having the sandstone which is wanting at Granite Falls, and also in the great masses of lime- stone which are replaced at Granite Falls only by the calcareous drift pebbles anda few bowlders. Yet these two advantages are probably quite overcome by the great masses of exposed gran- ite at Granite Falls, not replaced at Mankato in any way, since granitic bowlders are equally abundant in both places. Comparing the Mankato vicinity with Minneapolis and with Fayette, lowa, two areas compared in a former portion of these studies, we find that it has a much richer lichen flora than the former region which gave only 113 lichen forms and probably nearly as rich as the latter which gave 157 lichens which one could expect to find in a study of limited duration. Minnesota has now furnished more lichens than any other state in the Mississippi Valley, having 351 species and varieties. Illinois with 249 lichens being next in order. Yet the fact that 280 MINNESOTA BOTANICAL STUDIES. northeastern Iowa, a portion of a State not so thoroughly sur- veyed and only having 226 known lichens, has 26 lichens not yet found in Minnesota, shows that the study of Minnesota lichens is by no means yet approximately completed, since a large part of these 26 rare or obscure lichens found already within 50 miles of the state certainly exist within its bord- ers in the southeastern portion, and other unstudied portions of the state may yet be expected to bring additions to the lichen flora in like proportion. A list of these 26 lichens could be added with habitats to aid in their discovery in southeastern Minnesota but an inspection of another paper* will give the names of them. A study of the table above, giving habitats and number of species for each, by per cents, would give a somewhat larger per cent. of lithophytic lichen species for the two areas consid- ered than a former study exhibited for the Minneapolis and Iowa localities and about the same per cent. as the lake Superior region. I subjoin, arranged according to habitat, a list of the 41 lichens added to the state in this paper. From the list it will. be seen that more than half of these species are most common on rocks, and that the great Archean and Algonkian masses exposed throughout the upper valley alone produced one-third of them. For convenience of reference to the above statements I shall now add the table, placing rock species first, and then follow the list with further discussion. New to Minnesota on Archean or Algonkian rocks. Ramalina polymorpha (Acu.) Tuck. Parmelia saxatilis (L.) Fr. var. panniformis (AcH.) SCHAER. Pyrenopsis phzococca Tuck. Pyrenopsis melambola Tuck. Omphalaria phyllisca (Wanv.) Tuck. Leptogium pulchellum (Acu.) Nyv. Lecanora sp. Lecanora subfusca (L.) Acu. var. allophana Acu. Lecanora cinerea (PreRs.) Nyt. var. cinereoalba var. nov. Rinodina sophodes (Acu.) Nyt. var. tephraspis Tuck. * Fink, B. Review of Lichenological Studies in the Upper Mississippi Valley, with suggestions for future investigations. In list to be published in Memoirs of the Torrey Botanical Club. Fink: LICHENS OF THE MINNESOTA VALLEY. 281 Rinodina lecanorina Mass. Urceolaria actinostoma PErRs. Buellia pullata Tuck. New to Minnesota on limestone. Omphalaria kansana Tuck. Omphalaria pulvinata Nyv. Collema plicatile SCHAER. Collema pustulatum Acu. Lecanora bookii (Fr.) Tu. Fr. Rinodina bischoffii (Hrpp.) Korrs. Buellia alboatra (Horrm.) Tu. FR. var. saxicola Fr. Staurothele diffractella (NyL.) Tuck. New to Minnesota on wood. Placodium ferrugineum (Hups.) Hepp. Placodium ferrugineum (Hups.) Hepp. var. pollinii Tuck. Cladonia cristatella Tuck. var. paludicola Tuck. Biatora flexuosa Fr. Biatora suffusa Fr. Buellia turgescens (Nyv.) Tuck. Opegrapha varia (PerRs.) Fr. var. pulicaris Fr. Arthonia sp. Endocarpon arboreum SCHWEIN. Pyrenula gemmata (Acu.) Nake. Pyrenula hyalospora Nyt. Pyrenula quinqueseptata (Nyu.) Tuck. Pyrenula glabrata (Acu.) Mass. Pyrenula megalospora sp. nov. New to Minnesota on earth. Heppia despreauxii (Mont.) Tuck. Heppia polyspora ‘Tuck. Collema tenax (Sw.) Acu. Biatora decipiens (Enru.) FR. Biatora decipiens (Euru.) Fr. var. dealbata Aucr. The list of species new to the state shows a large number of Pyrenulas, the genus being unusually well represented in the valley, especially at Mankato. It will also be seen that the 282 MINNESOTA BOTANICAL STUDIES. gelatinous lichens, the Co//eme7, are especially conspicuous in the genera Pyrenopsis, Omphalaria, Collema and Leptogium. This happens because part of the valley is more favorable for their development as to substrata and moisture than other studied portions of the state. The part of the studied por- tion of the valley most favorable for their development is the Mankato vicinity where most of the gelatinous lichens were found. ‘The whole number of Co//emer found in the valley is 17. Richness is apparent when we add that only four were found about Minneapolis, 11 in the lake Superior region and that only 16 are known in Iowa. It may be added that a large proportion of the species added to the state flora are of special interest for various reasons. Thus the Omphalarias are not commonly collected; Lecanora book (¥r.) Th. Fr. is a difficult lichen to detect; the Pyrenulas are difficult to distinguish macroscopically and are therefore commonly overlooked; members of the genus Pyrenofpszs are seldom reported; while Urceolaria actinostoma Pers., Buellia pullata Tuck. and Heppia polyspora Tuck. are very rare lich- ens. /tinodina lecanorina Mass. is reported for the first time from North America, and Lecanora cervina (Pers.) Nyl. var. cinereoalba var. nov. is interesting because new. It may be noted in passing that the region shows some of the Arctic or sub-Arctic species found at Taylor’s Falls and already discussed in a former paper. These are Bvratora rufonigra Tuck., two forms of Buellia petrea (Flot., Koerb.) Tuck. and an £pheée, though not the species reported from Taylors Falls. As in the Taylors Falls region the Buellz is the most common of these species being a crustaceous form well adapted to resist unfavorable conditions. The Svatora is next in fre- quency of occurrence and the Aphebe, a fruticulose form, was only seen once. So far as I was able to ascertain by careful search the foliaceous forms, Umézlicaria and Vephroma, found at Taylors Falls have not succeeded in persisting in the Min- nesota valley. This failure of northern forms to persist so suc- cessfully may be accounted for perhaps in a very small degree by more southern position of the area now under consideration, but no doubt is due much more to climatic and edaphic factors which have allowed plant migrations to proceed northward more rapidly in the Minnesota valley than farther east in the state since the last retreat of the glaciers. This matter has been Fink: LICHENS OF THE MINNESOTA VALLEY. 283 touched upon by Professor C. MacMillan.* It is interesting to note that the strictly crustaceous Luella is the only one of the more northern forms found in the state which persists as far south as Pipestone. Indeed, its abundance here and records of occurrence elsewhere well southward in low altitudes since Tuckerman wrote lead to the suspicion that it. may not be so strictly sub-Arctic in distribution as I have supposed. It may be added that the Bella is the only one of these northern species persisting in the valley, which was found on bowlders at any considerable distance from the large masses of Archean and Algonkian rocks, which are supposed to have been exposed continuously since the close of the glacial epoch, and that it was only found once in very small quantity on a bowlder re- mote from these larger masses. It has been my plan to introduce in each paper of the series some feature regarding distribution which could be especially well illustrated by the area under consideration. In the study of the Minnesota valley and southwestern Minnesota I was able to keep in mind a variety of ecologic factors and to pre- serve the data necessary for their solution. This I had pre- viously done in part for several areas in Minnesota and Iowa so that in the present paper interesting and instructive comparisons can be made. Leaving other questions, then, thus briefly stated, I shall now pass to a consideration of the lichen for- mations of the region, causes of their peculiar make-up, and comparisons with similar formations within and outside the area under consideration. Aside from the purely scientific interest of the analysis to follow, it has a practical bearing, in that knowledge of the re- lation between ecologic factors and distribution enables the col- lector to predict in the field about what species of lichens he may expect to find in a spot having a given set of environmen- tal features. In the study species rarely found in the forma- tions have not been considered when there appeared to be doubt as to whether they were collected on their usual substrata, and rarer varieties have been omitted when showing the same habi- tat as other forms of the species. It will be readily granted that the commoner forms which give character to the flora are the ones which should receive attention in such a study. In * MacMillan, C., Observations on the Distribution of Plants Along the Shore at Lake of the Woods. Minn. Bot. Stud. 1: 967. 1897. 284 MINNESOTA BOTANICAL STUDIES. the analysis, especially as to amount of illumination and the roughness of ligneous substrata, it will be seen that lines can not be drawn very closely without entailing an amount of minutia which would be confusing and therefore unprofitable. With the above briet statement as to the main purpose of the present paper, I shall begin the consideration of lichen forma- tions with the most distinct ones with which I am acquainted, viz., those of the Sioux quartzite at Pipestone. These forma- tions are distinct because for most part removed from trees from which lichens commonly migrate to rocks nearby, producing tension lines and mixture of formations and because the few young trees found, though large enough to bear the foliaceous lichens which commonly migrate to the rocks, have apparently been isolated from larger areas of trees from the beginning of growth and scarcely bear a lichen of any kind. The rocky substratum is for the most part horizontal and exposed to the sun’s rays. In a few places occur perpendicular rock expos- ures which are more or less shaded by trees, overhanging rocks or north exposure. A few ombrophytic lichens occupy these spots; but they are all strictly lithophytic species, none of them having, for the reason stated above, migrated from trees as we shall find to be the condition ina later analysis of other similar formations. Below I give first the lichen formation of the hori- zontal exposed rocks and second, that of the more or less shaded and damp rocks. Lecanoras predominate in the formations on exposed rocks, which may accordingly be named as follows: Lecanora formation of the hortzontal exposed quartzite (Pipestone). Parmelia olivacea (L.) Acu. var. prolixa Acu., C. Parmelia conspersa (Euru.) Acu., C. Physcia tribacia (Acu.) Tuck., C. Physcia cesia (Horrm.) Nyt. Placodium elegans (Linx.) DC., C. Placodium vitellinum (Enru.) Narc. and Hepp. Lecanora rubina (ViLu.) Acu., C. Lecanora rubina (Vi_u.) Acu. var. heteromorpha Acu., C. Lecanora cinerea (L.) SommeERrF., C. >| Lecanora xanthophana Nyv., C. Fink: LICHENS OF THE MINNESOTA VALLEY. 285 Rinodina oreina (Acu.) Mass., C. Buellia spuria (ScHAER.) ARN., C. Buellia pullata Tuck., C. Buellia petrea (FLor., Korres.) Tuck. var. montagnei ePucK., ©. Endocarpon miniatum (L.) ScHAER. var. complicatum ScHAER., C. The formation on shaded rocks may be designated the Stauro- thele formation, after the prevailing genus. Staurothele formation of shaded or damp quartzite (Pipestone). Endocarpon miniatum (L.) ScHArER. Staurothele umbrina (Waut.) Tuck., C. Staurothele drummondii Tuck., C. The lichen formations of the pipestone lying beside the quartz- ite were studied to ascertain to what extent the difference in chemical composition and hardness of the rocks would influence the distribution of lichens, other ecologic factors being identical. In the above table I have indicated species common to quartz- ite and pipestone by (C.), and the table shows that only three lichens were detected on the quartzite and not on the pipestone. The following three, all growing in exposed places, were found on the latter and not on the former. Placodium cinnabarrinum (Acu.) Auz. Placodium cerinum (HEpw.) Narc. and Hepp. var. sideritis ueK. Lecanora muralis (ScCHREB.) SCHAER. var. saxicola SCHAER. It is worthy of note that the differences are specific and that the formations are identical generically. The appearance of a certain plant in a particular set of ecological conditions is too complicated a matter for exact explanation in many instances, and I can offer no explanation as to why the few plants occur on one kind of rock and noton the other. Possibly the specific acid secreted by a particular species acts more readily on one kind of rock than on the other, but more probably the cause is other than this. Nor do I suppose that I have found, here or in other formations to be considered below, all the lichens grow- ing under a particular set of conditions. Yet the common ones which give character to the various formations were doubtless. 286 MINNESOTA BOTANICAL STUDIES. all detected here as elsewhere, and the fact that 15 of 18 were found on each kind of rock demonstrates that difference in composition of rock in this instance has produced little, if any difference in lichen flora. A similar study of lichen formations on large rock areas of greater difference in composition as granite and limestone lying adjacent would be of special interest. To complete the lichen formations of the area, the earth- lichen formation must be considered. ‘This formation and simi- lar ones elsewhere may be called the Axdocarpon hepaticum for- mations of exposed earth from a plant which is found in such formations in all parts of the state except the lake Superior region. Endocarpon hepaticum formation of exposed earth (Pipestone). Urceolaria scruposa (L.) Nyv. Cladonia pyxidata (L.) FR. Cladonia fimbriata (L.) Fr. Cladonia fimbriata (L.) Fr. var. tubeformis Fr. Biatora muscorum (Sw.) Tuck. Endocarpon hepaticum Acu. Endocarpon pusillum Hepw. var. garovaglii Keun. The region is a comparatively dry one because of small pre- cipitation of moisture, since the rocks lie high where there is little or no standing water to give moisture and because there are few trees to give shade. ‘The lichen formations are accord- ingly rather poor in species, as will appear in comparisons to follow an analysis of similar formations. The rocky surfaces at Granite Falls present a much more complex set of conditions than those just considered, and yet, for my purpose, they may be classified, like the latter, into ex- posed surfaces, usually horizontal, and shaded surfaces, usually more or less nearly perpendicular. I shall now record these formations in the same order as in the last series; but after each shall compare it with the corresponding formation at Pipestone, giving, as far as possible, the probable cause of differences. Lecanora formation of exposed (usually horizontal) granite (Granite Falls). Parmelia olivacea (L.) Acu. var. prolixa Acu. Parmelia conspersa (Euru.) Acu. Fink: LICHENS OF THE MINNESOTA VALLEY. 287 Physcia stellaris (L.) Tuck. var. apiola Nyu., A. Physcia cesia (Horrm.) Nyt. Placodium elegans (Linx.) DC. Placodium murorum (Horrm.) DC., A. Placodium cinnabarrinum (Acn.) Auz. Placodium cerinum (Hrpw.) Narc. and Hepp. var. sideritis (euEK. Placodium vitellinum (Enru.) Narc. and Hepp. Lecanora rubina (ViLu.) Ac. Lecanora rubina (ViLuL.) Acu. var. heteromorpha Acu. Lecanora muralis (SCHREB.) SCHAER., A. Lecanora muralis (ScHREB.) SCHAER. var. saxicola ScHAER. Lecanora frustulosa (Dicxs.) Mass., A. Lecanora subfusca (L.) Acu. var. allophana Acu., A. Lecanora subfusca (L.) Acu. var. coilocarpa Acu., A. Lecanora hageni Acu., A. Lecanora cinerea (L.) SoOMMERF. Lecanora calcarea (L.) Sommerr. var. contorta Fr., A. Lecanora xanthophana Nyt. Lecanora cervina (PEerRs.) Nyt. var. cinereoalba var. nov., A. Lecanora fuscata (ScHRAD.) Tu. FrR., A. Rinodina oreina (Acu.) Mass. Rinodina sophodes (Acu.) Nyt., A. Rinodina lecanorina Mass., A. Urceolaria actinostoma Pers., A. Biatora rufonigra Tuck., A. Buellia spuria (ScHAER.) ARN. Buellia pullata Tuck. Buellia petrea (FLor., Korrs.) Tuck. Endocarpon miniatum (L.) ScHaAER., var. complicatum SCHAER. Comparing this lichen formation with the similar ones of the Sioux quartzite and the pipestone, we find it to contain all lichens found on the two except Physcza tribacia (Ach.) Tuck. and to contain fourteen not found on them, which I have marked as additions (A). The absence of the one species from the Granite Falls formation is doubtless an accident in plant distri- 288 MINNESOTA BOTANICAL STUDIES. bution whose explanation would be very difficult or impossible to trace; but it is quite remarkable that with this exception all the plants found in the two formations sixty miles away should oc- cur in this lichen formation also, especially since there could have been no rocky connection between the two areas since glacial times. It is not strange that the exposed granite lichen formation at Granite Falls should be a much richer one than the two exposed formations at Pipestone combined; for itis a much larger area, is connected with a limestone lichen formation and an epiphytic, and a number of swamps and ponds furnish mois- ture along the borders. Indeed the presence of ten of the fourteen additions may be more or less satisfactorily explained. These I shall proceed to consider sevzatim. Physcia stellaris (L.) Tucx., var. apiola Tucx.—a litho- phytic variety of a species common on adjacent trees. Lecanora frustulosa (Dicxs.) Mass.—a northern lichen not extending so far south as Pipestone. Lecanora subfusca (L.) Acu., var. alliophana AcH.—a variety of a species common on trees near by. Lecanora subfusca (L.) Acu., var. coilocarpa AcH.—as the last above. Lecanora cervina (PeRs.) Nyu., var. cinereoalba var. nov.— has not been seen outside the Minnesota valley. Lecanora calcarea (L.) SOMMERF., var. contorta Fr.—a lichen migrating from the limestone near by. Rinodina sophodes (Acu.) Ny_.—found on trees of the region and perhaps migrating from them. Rinodina lecanorina Mass.—a very rare plant which, therefore, very probably does not exist at Pipestone or was overlooked. Urceolaria actinostoma PErs.—as the last above. Biatora rufonigra ‘’'uck.—a northern form not extending so far south as Pipestone. Though somewhat confusing another similar lichen formation must be introduced here for comparison as follows: Lecanora formation of exposed quartzite (New Ulm). Parmelia conspersa (Euru.) Acu., CTS. Physcia cesia (Horrm.) Nyu., CTS. Placodium cerinum (Hrepw.) NArc. and Hepp. var. sideritis Pock.,-CT; Fink: LICHENS OF THE MINNESOTA VALLEY. 289 - Placodium vitellinum (Eurn.) NAc. and Hepr., CTS. Lecanora rubina (ViLu.) Acu., CTS. Lecanora rubina (ViLu.) Acu. var. heteromorpha Acu., CS. Lecanora subfusca (L.) Acu., S. Lecanora varia (Euru.) Nyv., AS. Lecanora cinerea (L.) Sommerr., CTS. Lecanora xanthophana Ny -., C. Rinodina oreina (Acu.) Mass., C. Rinodina sophodes (Acu.) Nyt. Biatora rufonigra Tuck., T. Biatora myriocarpoides (Nyu.) Tuck., A. Buellia spuria (ScHAER.) Arn., CT. Buellia petrea (Firor., Korrs.) Tucx., CTS. Endocarpon miniatum (L.) ScHArER. var. complicatum SCHAER., C. Comparing the above lichen formation with the similar ones at Pipestone and Granite Falls we find it to contain only two species which are additions to the three at the two places just named. These I have marked (A). It is about as extensive an area as the two at Pipestone combined, has about the same number of lichens as both and has 12 species (marked C) which are common to all the exposed rock lichen formations in the area considered in this paper. In general these 12 species may be regarded as the most constant of the exposed Archean and Algonkian rock lichen formations of southwestern Minnesota. As we multiply areas of comparison and especially as we introduce those at a greater distance the number of common floral elements very naturally decreases. Thus considering the similar forma- tion at Taylors Falls, we find only 8 species (marked T) com- mon to it and all the similar ones previously considered, and passing to the corresponding formation at Gunflint in the lake Superior region, the number found in all these similar forma- tions in widely separated areas of the state is found to be only 6 (marked S). These 6 species may be looked for with con- siderable certainty wherever such lichen formations are well de- veloped in the state. Other elements will vary according to relation to other adjacent formations, position northward or southward and in some instances eastward or westward in the state and to various ecologic factors which cannot be enum- erated fully. 290 MINNESOTA BOTANICAL STUDIES. We may now turn to the lichen formation of shaded or damp rocks at Granite Falls. This includes some flat rock surfaces somewhat shaded or simply wet part of the time, as well as the perpendicular shaded surfaces. I shall divide the formation into three parts—species naturally belonging to the rocks, those which have probably migrated from the trees near at hand and those which have probably migrated from the earth. Here and in another formation we have a mixture of elements, hence the following name is proposed : Mixed formation of shaded (or damp) granite (Granite Falls). A. PROBABLY NATURALLY BELONGING TO THE ROCKS. Ramalina polymorpha (Acu.) Tuck. Ramalina calicaris (L.) Fr. var. farinacea SCHAER. Pannaria microphylla (Sw.) DELIs. Pannaria languinosa (Acu.) KoERB. Omphalaria phyllisca (WanL.) Tuck. Collema furvum (Acu.) Nyt. Leptogium lacerum (Sw.) Fr. Endocarpon muriatum (L.) SCHAER. Staurothele umbrina (WaunL.) Tuck. Staurothele diffractella (NyL.) Tuck. Staurothele drummondii Tuck. B. NEAR TREES AND PROBABLY MIGRATED FROM THEM. Parmelia cetrata Acu. Parmelia crinita Acu. Parmelia borreri TuRN. Parmelia borreri Turn. var. hypomela Tuck. Parmelia saxatilis (L.) Fr. Parmelia saxatilis (L.) Fr. var. sulcata Nyu. Parmelia saxatilis (L.) Fr. var. panniformis (AcH.) SCHAER.. Parmelia caperata (L.) Acu. Physcia speciosa (WuLF., Acu.) NyL. — Physcia pulverulenta (Scures.) Nyt. Physcia obscura (Euru.) Nvyv. Pyxine sorediata Fr. Leptogium myochroum (Euri., ScHAER.) TUCK. Fink: LICHENS OF THE MINNESOTA VALLEY. 291 Placodium aurantiacum (Licurr.) Narc and Hepp. Biatora fuscorubella (Horrm.) Tuck. C. SPprECIES WHICH HAVE PROBABLY MIGRATED FROM EARTH. Peltigera rufescens (NEck.) Horr. Peltigera canina (L.) Horrn. Of the three parts of the formation under consideration only the first can be compared with the similar formation at Pipe- stone, and we find besides the 3 species of the Pipestone for- mation, 8 additional forms as a result of greater areas studied, more moist conditions near the Minnesota river, and where abundant ponds and marshes situated near the rocks give moist- ure, and where trees are numerous in some parts of the area and increase the shade. I must add that presence of the feamalinas here, and their absence from shaded rocks at Pipe- stone leads to the suspicion that they may have sprung from Reamalina calicaris (L.) Fr. of the region, migrating from trees to rocks and acquiring the varietal, and in one instance the specific characters as an adaptation to changed environment. The question is as to whether these lichens are sufficiently plastic to acquire such new characters since trees have grown in the valley in post-glacial time. I can only say that I believe that they may be, and that it is quite as likely that the two Aama- finas should be placed in the second division of the formation as in the first. As to plants of the second portion of the formation, which I have designated as having probably migrated from trees, in some instances they are locally more abundant and luxuriant on the rocks than on trees. Hence a hasty consideration would lead to the conclusion that they have not migrated. But the luxuriant condition obtains on the rocks in Parmelia borrer? Turn., a lichen seldom seen on rocks elsewhere, and many of these lichens grow on mossy rocks where lichens are commonly large. Also it is to be taken into account that these lichens are those usually found on large trees with rough bark. The larger trees were for most part destroyed years ago by man or fires, and these lichens, formerly common on trees, are preserved on rocks better than on the less permanent trees. Hence some of them are more common now on the rocks than on the trees, which are for most part second growth and not large. The 292 MINNESOTA BOTANICAL STUDIES. third division, consisting of two Peltigeras, scarcely needs any special consideration. I shall next consider the similar shaded rock formation at New Ulm, which may be divided into those lichens naturally belonging to the rocks and those probably migrating from trees. Mixed lichen formation of shaded rocks (New Ulm). A. NATURALLY BELONGING TO THE ROCKS. Pannaria languinosa (AcH.) KoERB. Collema flaccidum Acu. Collema furvum (Acu.) Nyt. B. NEAR TREES AND PROBABLY MIGRATED FROM THEM. Theloschistes lychneus (Nyu.) Tuck., CTS. Parmelia crinita Acu., CTS. Parmelia borreri TurN., CTS. Parmelia saxatilis (L.) Fr., CTS. Parmelia saxatilis (L.) Fr. var. panniformis (AcH.) SCHAER., Parmelia caperata (L.) Acu., CTS. Physcia speciosa (WuLF. Acu.) Nyu., CTS. Physcia pulverulenta(Scures.) Nyu., CTS. Physcia stellaris (L.) Tucx., TS. Physcia obscura (Euru.) Nyu., CTS. As to the shaded rock lichen formations of the region sur- veyed considering only plants naturally belonging to the rocks, there is not a single lichen that is common to all of them. Pan- narta languinosa (Ach.) Koerb. is the most constant element of such formations, which as a whole might be named for this plant were it not quite as common in shaded limestone forma- tions otherwise quite different from any of those on the rocks under consideration at present. Of the lichens of the shaded rock formation at New Ulm, which have probably migrated from trees, the nine marked common (C), may be taken as the ones most commonly occurring, as they were found also at Granite Falls in the similar formation. Those marked (T) all but one of the nine, occur in the similar formation at Taylors Falls. Other elements vary more with change in various eco- logic factors. The similar partial formation was noted at Grand Portage, especially on the island, and adding those Fink : LICHENS OF THE MINNESOTA VALLEY. 293 lichens (S) of it found in the corresponding ones considered above, subtracts none from the number of common species. Therefore, these eight lichens may be regarded as the elements of that portion of the shaded rock lichen formations which have probably migrated from trees, most widely occurring in such formations over the state. Only one day was spent in study- ing the New Ulm formations. A second day would have added somewhat to the list, yet doubtless all the dominant lichen floral elements were secured. Without entering into a detailed analysis, it will appear from an inspection of the lichens composing the formations for shaded and for exposed rocks that the species occurring in the former are for most part foliaceous or fruticulose types, while those given for the latter are in general crustaceous, or if foliaceous, at least closely prostrate on the rocks. This is what would be expected, since shade favors better development of thallus, so that those species showing good thalli crowd out the other species in shaded places, or when unshaded become shaded with the growth of trees. Next in order come the earth lichen formations of the rocky areas of Granite Falls and New Ulm. I shall first record the exposed formations for the two localities and compare with the similar formation already recorded for Pipestone. Then will follow the lichen formations of shaded earth at the first two sta- tions, which is scarcely developed at Pipestone. A consid- eration of calcareous-earth lichen formations follows, the pres- ent being formations of non-calcareous earth. Endocarpon hepaticum lichen formation of exposed earth (Granite Falls). Heppia despreauxii (Monr.) Tuck. Urceolaria scruposa (L.) Nyv. Cladonia pyxidata (L.) Fr. Biatora muscorum (Sw.) Tuck. Biatora icterica Monr. Endocarpon hepaticum Acu. Endocarpon pusillum Hepw. var. garovaglii Keun. Endocarpon hepaticum lichen formation of exposed earth (New Ulm). Cladonia pyxidata (L.) Fr., CTS. 294 MINNESOTA BOTANICAL STUDIES. Cladonia turgida (Enru.) Horrn. Biatora uliginosa (ScHRAD.) Fr. Endocarpon hepaticum Acu., CT. Endocarpon pusillum Hepw. var. garovaglii Kpun., C. Comparing these lists with the one given for the correspond- ing formation at Pipestone, we find three common lichens which are marked (C) in the list above. Two of these marked (T) are also found in the similar formation at Taylors Falls, and one marked (S) is common in like formations in the lake Superior region. This plant is the most constant element in the exposed earth lichen formations of the State, and I should be disposed to name these Cladonia pyxidata lichen formations, were it not that the plant, though commonly present in exposed stations, thrives better in shaded ones. I must here emphasize that these, as well as the calcareous-earth lichen formations, grow on earth in rocky places where larger vegetation is scanty and scattered. Next in order come lichen formations of shaded earth, partly composed of plants which grow also, though not so well, in un- shaded places. From their dominant elements, these may be designated as follows: Cladonta-Peltigera lichen formation of shaded earth (Mankato). Peltigera rufescens (NEck.) HorrM. Peltigera canina (L.) Horr. Peltigera canina (L.) Horr. var. sorediata SCHAER. Collema pulposum (BEerNu.) Nyt. Collema tenax (Sw.) Acu. Cladonia pyxidata (L.) Fr. Cladonia fimbriata (L.) Fr. Cladonia gracilis (L.) Fr. Cladonia gracilis (L.) Fr. var. verticillata Fr. Cladonia-Peltigera lichen formation of shaded earth (Granite Falls). Peltigera rufescens (NEck.) Horrm., CT. Peltigera canina (L.) Horrm., CTS. Peltigera canina (L.) Horr. var. sorediata ScuarEr., CTS. Collema pulposum (BeRNu.) Nyw., CT. Fink: LICHENS OF THE MINNESOTA VALLEY. 295 Cladonia pyxidata (L.) Fr., CTS. Cladonia fimbriata (L.) Fr., C. Cladonia fimbriata (L.) Fr. var. tubeformis Fr., TS. Cladonia gracilis (L.) Nyu., CTS. Cladonia gracilis (L.) Nyv. var. verticillata Fr., CTS. These two formations are remarkably similar, having 8 com- mon forms (C) of a total of nine lichens in each formation. Including the similar formation at Taylors Falls (T) we still have 7 lichens common to the similar formations for a large part of Minnesota, and extending the observation to the similar formation on Grand Portage island in the lake Superior (S) region, we yet have 6 lichens common to such formations selected from widely separated areas inthe State. This is the first kind of formation thus far considered which is found in the Minne- apolis area studied. Therefore data from this region have not been introduced thus far. Their use in the present considera- tion would not decrease the number of common elements, and I shall not add them. The three rarer Cladonzas of the region under consideration in the present paper, Cladonia symphycar pia Fr., Cladonta mitrula Tuck. and Cladonia cariosa (Ach.) Spreng. have been purposely omitted, as there is yet doubt as to whether their adaptation is ombrophytic. As to the nature of the lichens composing these earth lichen formations, it is apparent that those of the shaded earth forma- tions are as a whole more foliaceous or fruticulose and better developed as to thallus than those of the exposed earth forma- tion. The explanation is of course the same as that already given for exposed and shaded rock lichen formations. I shall now consider the one remaining earth lichen formation at Granite Falls and compare it with a similar one in another region. Itis that of the earth_among the calcareous drift peb- bles and small boulders on hill sides. From the calcareous nature of the earth and the presence of a Bzatora seldom seen elsewhere than in such formations, the following name has sug- gested itself. Biatora decipiens lichen formation of exposed calcareous earth (Granite Falls). Heppia despreauxii (Monr.) Tuck. Heppia polyspora Tuck. 296 MINNESOTA BOTANICAL STUDIES. Urceolaria scruposa (L.) Nyv. Biatora muscorum (Sw.) Tuck. Biatora decipiens (Enru.) FR. Biatora decipiens (Euru.) Fr. var. dealbata Aucr. Endocarpon hepaticum Acu. Some of the plants of this formation have been found at Mankato and also at Minneapolis, but the formation is not well developed at either place. However, it is beautifully developed at Fayette, Iowa, and because of its remarkable similarity there to the Granite Falls formation about two hundred miles distant, I give it below for the sake of comparative study. Biatora dectpiens lichen formation of exposed calcareous earth (Fayette, Iowa). Heppia despreauxii (Mont.) Tuck., C. Urceolaria scruposa (L.) Nyt., C. Biatora muscorum (Sw.) Tuck., C. Biatora decipiens (Enru.) FR., C. Biatora decipiens (Euru.) Fr. var. dealbata Aucrt., C. Biatora fossarum (Dur.) Monr. Endocarpon hepaticum Acu., C. It will be seen that the two formations are identical except that each one contains one species not found in the other. Again, this slight difference becomes less significant when it is stated that each of these two plants not found in both formations is rather rare in the formation in which it occurs. The six lichens common to both formations I have indicated in the Fay- ette list (C). In both localities the formations are formed on hill sides and seen to be somewhat better developed on south- ward than on northward slopes. I have not seen similar forma- tions well developed elsewhere, but it is probable that they reach their best development on unshaded hill sides where other vegetation is scanty and where the lichens are washed with lime-impregnated water flowing down the slope during rains. Biatora decipiens (Ehrh.) Fr. and Endocarpon hepaticum Ach. are the most common plants of these formations, but the latter is quite as common in another formation of non-calcareous earth, which I have named for it, not confined to hill sides. Fink: LICHENS OF THE MINNESOTA VALLEY. 297 Closely related to the above formations are two occupying the same areas and named for a lichen almost wholly confined to them. They follow below : Lecanora calcarea contorta lichen formation of exposed lime- stone pebbles (Granite Falls). Placodium vitellinum (Euru.) NArc. and Hepp. var. aurel- lum Acu. Lecanora calcarea (L.) SoMMERF. var. contorta Fr. Lecanora privigna (Acu.) Nyu. Lecanora privigna (Acu.) Nyw. var. pruinosa Auct. Endocarpon pusillum Hepw. _ Verrucaria muralis Acu. Staurothele diffractella (NyL.) Tuck. Like the last, this formation is not well developed in other studied portions of Minnesota, and I shall give the similar one for Fayette, lowa, for comparison. Lecanora calcarea contorta lichen formation of exposed lime- stone pebbles (Fayette, lowa). — Placodium cinnabarinum (Acu.) Auz. Placodium vitellinum (Enru.) Narc. & Hepp. var. aurellum NCH. (GC. Lecanora muralis (ScHREV.) SCHAER. var. versicolor FR. Lecanora calcarea (L.) SOMMERF. Lecanora calcarea (L.) SomMeErrF. var. contorta Fr., C. Lecanora privigna (Acu.) Nyt., C. Rinodina bischoffii (Hepp.)Korers. Biatora russellii Tuck. Endocarpon pusillum Hepw., C. Verrucaria nigrescens PErRs. Verrucaria muralis Acu., C. Lichens common to the two formations are marked (C) in the Fayette list, and comparison shows marked similarity in the two formations about 200 miles distant, except that the latter is con- siderably better developed than the former. This is as would be expected when we consider that the Iowa region is one where limestones abound, while the Minnesota is one in which the limestone pebbles are those transported in glacial drift and are 298 MINNESOTA BOTANICAL STUDIES. less numerous. All the species of these formations, except the Biatoras, have been found elsewhere in Minnesota, but not ag- gregated into definite formations. Comparing the last two series of formations, viz., those of calcareous earth and those of drift pebbles of the same areas, it will be noted that the former, because of their position on dry hill-sides, consist as a whole of lichens having small foli- aceous or granular thalli, while those on the yet dryer and harder calcareous pebbles are almost entirely made up of strictly crustaceous plants. The formations of exposed and shaded limestone bluffs come next in natural order, and the analysis is difficult, since some of the lichens of these formations grow about equally well in sun- shine and shade. These | shall indicate by an interrogation point (?). From the prevalence of gelatinous lichens they may be named as follows: Gelatinous lichen formation of shaded (or damp) limestone bluffs (Mankato). Pannaria nigra (Hups.) Nyt. Pannaria languinosa (AcH.) KoERs. Omphalaria kansana T'uck.? Omphalaria pulvinata NyL.? Collema plicatile ScHAER. Collema pustulatum Acu. Leptogium lacerum (Sw.) Fr. Placodium citrinum (Horrm.) Leienr. Biatora inundata Fr. Buellia alboatra (Horrm.) Tu. Fr. var. saxicola Fr. Endocarpon miniatum (L.) ScHAER. Staurothele umbrina (WaAHL.) Tuck. Similar formations do not exist in other surveyed portions of Minnesota, except at Minneapolis, where the development is poor. It is as follows: Gelatinous lichen formation of shaded (or damp) calcareous rocks (Minneapolis). Pannaria nigra (Hups.) Nyv., C. Pannaria languinosa (AcH.) Korrs., C. Fink: LICHENS OF THE MINNESOTA VALLEY. 299 Omphalaria sp. Leptogium lacerum (Sw.) Fr., C. Endocarpon miniatum (L.) ScHarr., C. Placodium citrinum (Horrm.) Leienr., C. The plants of the Minneapolis list are all but one common (C) to both formations and may be regarded as characteristic of such formations. Since the last formation is poorly developed, I may add the similar one for Fayette, lowa, which is better developed than either of the above. Gelatinous lichen formation of shaded (or damp) calcareous rocks (Fayette, Iowa). Pannaria nigra (Hups.) Nyv. Pannaria languinosa (Acu.) Korrs. Omphalaria pulvinata Nyu.? Omphalaria umbella Tuck. ? Omphalaria sp. Collema plicatile ScHAER.? Collema furvum (Acu.) Nyv.? Collema pustulatum Acu.? Leptogium lacerum (Sw.) FR. Leptogium chlorometum (Sw.) Nyt. Placodium citrinum (Horrm.) Leicur. Biatora trachona For. Buellia alboatra (Horrm.) Tu. Fr. var. saxicola Fr. Endocarpon miniatum (L.) ScHAER.? Staurothele umbrina (Wauv.) Tuck.? The introduction of the Fayette formation is of special interest for the following reason. The first Minnesota formation is a mile back from the Minnesota river on a bluff along which the river once flowed, but which now is left dry except for the trees which overhang it and shade the lichens of the formation. The Fayette formationis on a bluff at the water’s edge, and the plants are growing within one to ten feetof the water. Doubt- less this in part causes the greater richness. The Mankato for- mation is an interrupted one, none of the plants persisting in wholly unshaded spots. The Fayette formation on the other hand, extends for miles, without complete interruption, wher- 300 MINNESOTA BOTANICAL STUDIES. ever the bluffs exist. With the greater amount of moisture at the water’s edge, some of the plants of the Fayette formation grow well in sunshine and even on south exposures. These I have indicated by an interrogation point (?). These, for most part gelatinous lichens, require a good amount of moisture ; and if growing far from water seek shade for it. In the Fayette locality many trees have been cut recently along the bluffs so that the plants are more exposed than formerly. The Minne- apolis list can be considered a formation only in the sense of a group of plants growing under like conditions, for owing to somewhat dryer climate the formation is poorly developed as to individuals and may be designated as a scattered formation, only one or two of the species usually growing in one limited area, along the bluffs and long stretches of bluff between these areas frequently not bearing a single plant of the formation. Next in order comes the lichen formation of exposed lime- stone bluffs, which I shall designate as follows from the presence of a large proportion of angiocarpous lichens. Angiocarpous lichen formation of limestone bluffs (Mankato). Theloschistes lychneus (Nyu.) Tuck. Placodium elegans (Linx.) DC. Placodium vitellinum (Eurn.) NArc. and Hepp var. aurel- lum Acu. Placodium aurantiacum (Licut.) Narec. and Hepp. ~ Lecanora hageni Acu. Lecanora erysibe Nyt.? Endocarpon pusillum Hepw.? Endocarpon miniatum (L.) ScHAER.? Staurothele diffractella (NyL.) Tuck. Verrucaria fuscella Fr. Verrucaria nigrescens Pers. Verrucaria muralis Acu. I might add similar formations from Minneapolis and Fayette, Iowa; but the analysis is very uncertain so that the comparisons could have little value. I shall now consider the sandstone bluff formations of certain localities, simply designating them as formations of damp sand- stone since they are found along streams where the rocks are Fink: LICHENS OF THE MINNESOTA VALLEY. 301 well supplied with moisture. The first of the formations is almost completely shaded, but the second is only partially shaded, moisture, the thing really sought by the plants, being sufficient in more or less exposed spots so that the less ombro- phytic plants of the group thrive twenty or thirty feet from the water’s surface, and even the more shade-loving ones are found in exposed spots nearer the water. I shall now record the formations as follows, designating the less ombrophytic plants of the second formation thus (?). For these formations I suggest the following name from a plant almost wholly con- fined to them in Minnesota. Usnea barbata rubrginea lichen formation of damp sandstone bluffs (Minneopa Falls). Ramalina calicaris (L.) Fr. var. farinacea ScHAER. Usnea barbata (L.) Fr. var. hirta Fr. Usnea barbata (L.) Fr. var. rubiginea Micux. Peltigera canina (L.) Horr. var. sorediata Acu. Leptogium chloromelum (Sw.) Nyt. Pannaria languinosa (Acu.) Korrs. Cladonia furcata (Hups.) Fr. Cladonia furcata (Hups.) Fr. var. racemosa FR. Urceolaria scruposa (L.) Nyt. Usnea barbata rubiginea lichen formation of damp sandstone bluffs (Minneapolis). Ramalina calicaris (L.) Fr. var. farinacea ScHArr., C. Usnea barbata (L.) Fr. var. hirta Fr., C. Usnea barbata (L.) Fr. var. rubiginea Micux.? C. Parmelia conspersa (Euru.) Acu.? T. Peltigera canina (L.) Horr. var. sorediata Scuarr., CE. Pannaria languinosa (AcH.) Korrs., C. Lecanora subfusca (L.) Acu. var. coilocarpa Acu.? T. Urceolaria scruposa (L.) Nyu.? C. Cladonia cespiticia (Pers.) Fu., T. Cladonia cornucopioides (L.) Fr.? E. Endocarpon pusillum (HeEpw.) var. garovaglii Kpn., E. 302 MINNESOTA BOTANICAL STUDIES. Comparing the two formations we find six common lichens of a total of nine recorded for the first and eleven for the second. Similar formations occur at Pictured Rocks, Iowa, and at Rap- idan, but I shall not multiply lists. As in the instance of cer- tain formations on shaded granite or quartzite recorded above, both of these formations are more or less mixed, being made up of lichens strictly lithophytic in adaptation and of others which have doubtless wandered from trees or from earth. As I have not been able to study such sandstone bluffs at a distance from trees, I have not attempted a definite analysis of these more limited formations as I did for the formations of the shaded granite and quartzite, but have simply indicated in the second list those which have probably wandered from trees by (T) and those from earth by (E). I have omitted from these sandstone formations some of the rarer plants which I should have in- cluded had I attempted an analysis of these mixed formations. I shall now proceed to the two formations of trees, viz., that of rough barked trees and that of trees having smooth bark. The distinctions are difficult in some instances as certain species grow in both habitats. Consequently, as in some instances, in formations previously considered, some plants are recorded for more than one formation. Moreover, it must be added that some of those recorded for rough barked trees frequently seek the smoother portions of the bark. The subfamily Parmed/ez is especially well developed in the rough bark formations, which may accordingly be named as follows: Parmele lichen formation of trees with rough bark (Mankato). Ramalina calicaris (L.) Fr. var. fraxinea FrR., G. Ramalina calicaris (L.) Fr. var. fastigiata Fr., G. Theloschistes chrysopthalmus (L.) Norm., G. Theloschistes polycarpus (Euru:) Tuck., G. Theloschistes lychneus (Nyu.) Tuck., G. Theloschistes concolor (Dick.) Tuck., G. Parmelia perforata (JAck.) Acu. Parmelia crinita Acu., G. Parmelia borreri Turn., G. Parmelia tiliacea (HorrMm.) FLoERK., G. Parmelia saxatilis (L.) Fr. Parmelia caperata (L.) Acu., G. Fink + LICHENS OF THE MINNESOTA VALLEY. 303 Physcia granulifera (Acu.) Tuck., G. Physcia pulverulenta (ScuresB.) Nyv., G. Physcia stellaris (L.) Tuck., G. Physcia tribacia (Acu.) Tuck. Physcia obscura (Enru.) Nyv., G. Physcia adglutinata (FLOERK.) Nyv. Collema pycnocarpum Nyt., G. Collema flaccidum Acu. Leptogium myochroum (Euru., SCHAER.) Tuck. Placodium aurantiacum (Licur.) Narc. and Hepp., G. Placodium cerinum (HrEpw.) Nate. and HeEpp., G. Lecanora subfusca (L.) Acu., G. Pertusaria pustulata (Acu.) Ny. Pertusaria leioplaca (AcH.) SCHAER. Pertusaria velata (TurN.) Nyv. Biatora rubella (Enru.) RABENH. Biatora fuscorubella (Horrm.) Tuck., G. Biatora subfusca Fr., G. Lecidea enteroleuca Fr., G. Buellia alboatra (HorrM.) Tu. FrR., G. Buellia parasema (Acu.) Tu. FR. Opegrapha varia (PERs.) Fr., G. Graphis scripta (L.) Acu., G. Graphis scripta (L.) Acu. var. limitata Acu., G. Arthonia lecideella Nyv. Arthonia radiata (Pers.) Tu. Fr., G. Coniocybe pallida (Prrs.) Fr. Pyrenula gemmata (Acu.) NaAge., G. Pyrenula hyalospora Nvt., G. Pyrenula nitida Acn. Pyrenula quinqueseptata (NyL.) Tuck. Pyrenula leucoplaca (WALLR.) Kpr., G. Pyrenula megalospora sp. nov., G. In order to avoid reproducing a large portion of the above long list of names, I have for the similar formation at Granite Falls marked those of the list found there (G) and add below the 304 MINNESOTA BOTANICAL STUDIES. only one found in the Granite Falls formation and not at Mankato, viz., Balora naegelit Tuck. Thus the mark (G) will indicate also those common to both formations and as a whole most characteristic of such lichen formations for the Minnesota valley. The Mankato area with its abundance of trees would, of course, be expected to possess richer tree lichen formations than Granite Falls, and with the exception of a single species, the rough bark formation of the latter area is but a partial repe- tition of that of the former. The formation on trees with smooth bark at Mankato con- tains all but two of the species of the similar formation at Granite Falls, and the treatment may be abbreviated as the last two above. The genus Pyrenu/a predominates in the forma- tion, and some of the species are among the lichens most char- acteristic of smooth bark. ‘Therefore, the formations may re- ceive the name which follows: Pyrenula lichen formation of trees with smooth bark (Mankato). Theloschistes polycarpus (Euru.) Tuck. Theloschistes concolor (Dicxs.) Tuck., G. Parmelia olivacea (L.) Acu., G. Physcia adglutinata (FLoERK.) Nyt., G. Placodium cerinum (HEpw.) Narc. and HEpp., G. Lecanora subfusca (L.) Acu., G. . Rinodina sophodes (Acu.) Nyt., G. Biatora fuscorubella (Horrm.) Tucx., G. Lecidea enteroleuca Fr., G. Graphis scripta (L.) Acu., G. Arthonia lecideella Nyt. Arthonia dispersa Nyt., G. Pyrenula punctiformis (Acu.) Naze., F. Pyrenula punctiformis (Acu.) Nage. var. fallax Nyu., F. Pyrenula nitida Acu., F. Pyrenula thelena Acu., F. Pyrenula cinerella (FLor.) Tuck., F. Pyrenula cinerella (ILor.) Tuck. var. quadriloculata var. nov. Pyrenula leucoplaca (WaLLR.) Kpr., GF. Fink : LICHENS OF THE MINNESOTA VALLEY. 305 The two formed on smooth bark at Granite Falls and not at Mankato are Leczdea enteroleuca Fr. var. achrista Schaer. and Arthonia punctiformis Ach. As in the rough bark formations, the one at Mankato is richer for the same reason and, strangely enough, my study of the Granite Falls area only discovered a single Pyrenu/a on smooth bark. My name is scarcely appro- priate for this formation, though it is for the one at Mankato as it would be for others from other localities in Minnesota and Iowa which might be added. Without adding another list or another complete formation, I have indicated by (F) in the above list the Pyrenu/as of that list which occur on smooth bark at Fayette, Iowa. Persons acquainted with lichen species will readily observe in the lists for rough bark and smooth bark lichen formations, that the formation on rough bark is composed principally of the more foliaceous and fruticulose lichens while those of the smooth bark formations are in the main crustaceous lichens. This is possibly due in part to the fact that these foliaceous and fruti- culose lichens more easily gain a foothold on the rough bark which breaks up the thallus of the lichens adapted to smooth bark, thus tending to kill them. However it is probable that light, shade and moisture are also factors, the large trees fur- nishing more shade than the smaller ones. Next in order naturally enough we may consider the lichen formations of old boards and old wood, and the formations are so nearly alike for Mankato and Granite Falls that they may be treated in a single list by giving the Mankato list and marking (G) those common to the Granite Falls formation also. Our Calicet are lichens seldom seen in any other formations, hence the following name may be applied. Calicet lichen formation of old boards and wood (Mankato). Theloschistes chrysopthalmus (L.) Norm., G. Placodium cerinum (Hrpw). Narc. and Hepp. var. pyrocea NDT eo), (or Lecanora hageni Acu., G. Lecanora varia (Euru.) Nyv., G. Rinodina sophodes (Acu.) Nyu., G. Rinodina sophodes (Acu.) Nyu., var. exigua Fr., G. 306 MINNESOTA BOTANICAL STUDIES. Buellia parasema (Acu.) Tu. FR., G. Buellia turgescens (Nyv.) Tuck. Calicium parietinum Acu. Thelocarpon prasinellum Nyt. The additions for Granite Falls are Cetrarza ciliarts (Ach.) Tuck., Lecedea enteroleuca Fr. and Calicium quercinum Ach. As in other instances the common forms are those most charac- teristic of such formations. I have not detected the Caliczum for which I have named the Mankato formations at Granite Falls, where it is replaced by another species, and I shall add the species, Acolzum tigillare (Ach.) Dn., which is one of the Calcez common in the similar formation at Fayette, lowa, and the only one found in the like formation at Minneapolis. It must be admitted that the name used for these formations, while it may be applied, is not so appropriate for the related forma- tions in the lake Superior region where some of the Calcez grow on living bark and yet others on rotting wood. But one formation remains to be considered, viz., that of rot- ting stumps and prostrate logs. In these formations the most common plants are those of the genus C/adonza and the forma- tions may accordingly receive the following name: Cladonia formation of rotten wood (Mankato). Peltigera canina (L.) Horrn., G. Peltigera canina (L.) Horrm., var. sorediata SCHAER. Cladonia fimbriata (L.) Fr., G. Cladonia fimbriata (L.) Fr. var. tubeformis Fr., G. Cladonia gracilis (L.) Nyu., G. Cladonia gracilis (L.) Nyw., var. verticillata Fr., G. Cladonia symphycarpia Tuck. Cladonia macilenta (Euru.) Horr. Cladonia cristatella Tuck. The only species found at Granite Falls in the similar forma- tion and not at Mankato is Bratora flexuosa Fr. and the forma- tion may, with this addition, be indicated by marking (G) those plants of the Mankato formation common to both. Comparison with formations from other localities would show some varia- tion, but the C/adonias would predominate and give character to the formations. Wood commonly rots in moist shady places,, Fink: LICHENS OF THE MINNESOTA VALLEY. 307 furnishing an abundance of moisture, and we find accordingly that the formations on rotten wood are made up in large part of fruticulose lichens. The Cal/rcez formations of old wood are exposed to drier conditions and are composed almost entirely of lichens having poorly developed thalli. I must emphasize here that lichens of nearly all the forma- tions enumerated above enjoy moist places, and that lack of moisture produces a decrease in richness both in size and num- ber of individuals and in numbers of species in the formations. I repeat this, which I have established for some parts of Minne- sota previously, because some persons may suppose that lichens, because of their xerophytic adaptations, thrive as well in the driest spots as in those affording more moisture. The excep- tions to this general statement will appear from a careful study of the analyses made of the various formations. The gelatinous lichen formation of shaded limestone (Minne- apolis) has been called a scattered one, and I have explained what is meant by the expression. Others of the same kind are the Cladonia-Peltigera lichen formations of shaded earth, the angiocarpous lichen formations of exposed limestone bluffs, the Calicec lichen formations of old wood and in some instances the Cladonza lichen formations of rotten wood, though in other in- stances half or more of the species of Cladonza of the forma- tion may be found ona single log. Thus formations of the kind last named and like the one first named in this paragraph differs from the other three named in the paragraph in that they may or may not be scattered while the three always are, so far as I know, except the Calrcec formation which may be found nearly complete on afew rods of old fence in some favorable instances. The two formations of trees are widely extended ; but they are not scattered as I have used the term since one commonly finds a good proportion of the species of either for- mation in passing a short distance in the woods. Also in my classification we have the peculiar condition of two lichen formations occupying the same area. This is illus- trated by the Bratora decipiens lichen formation of exposed calcareous earth and the Lecanora calcarea contorta lichen formation of exposed limestone pebbles, or by the Lecanora lichen formations of exposed granite or quartzite and the Azdo- carpon hepaticum lichen formation of exposed earth. Yet it is apparent that the formations are distinct in both instances, the 308 MINNESOTA BOTANICAL STUDIES. division being based on substratum as well as amount of light and moisture. As a whole, the formations may be said to be azonal and without definite form or extent, both depending upon location of proper substrata, protection from or exposure to light, etc. In my paper I have used the expression ‘‘ lichen formation ” to include lichens only. Of course, these plants are in some instances found growing upon the same substrata and in the same general set of conditions as plants of other groups, and which might have been listed in the formations. However, I may be excused, in a paper on lichen distribution, for omitting other plants than lichens, especially since I could not possibly have treated the other plants with the same detail that I have accorded the lichens. I know of no other paper which has dealt exclusively with lichen distribution as I have done herein, and surely this anal- ysis must be helpful in the study of the lichen flora of other regions. The multiplicity of observations necessary for such a detailed study are not easy to make, and I am sure that much of interest has escaped me. However, I hope that this paper may stimulate others to study the lichens from an ecologic point of view. LIST OF SPECIES AND VARIETIES. 1. Ramalina calicaris (L.) Fr. var. fraxinea Fr. On trees and old wood, infrequent or rare. Mankato, June 23, 1899, no. 55, June 26, 1899, no. 102, and June 28, 1899, no. 164. Granite Falls, July 11, 1899, no. 385 and July 13, 1899, nos. 510 and 533. 2. Ramalina calicaris (L.) Fr. var. fastigiata Fr. On trees and rocks, rare. Mankato, June 23, 1899, no. 54. New Ulm, July 5, 1899, no. 275. Granite Falls, July 14, 1899, no. 518, and July 17, 1899, no. 588. 3. Ramalina calicaris (L.) Fr. var. farinacea SCHAER. On sandstone and granite. Mankato (Minneopa Falls), June 27, 1899, no. 154. Redwood Falls, July 6, 1899, no. 305, and July 8, 1899, no. 349. Granite Falls, July 12, 1899, no. 460. 4. Ramalina polymorpha (Acu.) Tuck. ? On shaded granitic rocks in first locality and on a large bowlder in the second, rare. Granite Falls, July 12, 1899, no. Fink: LICHENS OF THE MINNESOTA VALLEY. 309 456, and July 13, 1899, no. 492. Pipestone, July 19, 1899, no. 641. The plants are placed here provisionally. They resemble in part PRamalina calicaris (L.) Fr. var. fart nacea Schaer. Not previously reported from Minnesota and new to the Mis- sissippi valley. 5. Cetraria ciliaris (Acu.) Tuck. A single sterile specimen collected on an old cedar stump. Granite Falls, July 17, 1899, no. 570. 6. Usnea barbata (L.) Fr. var. florida Fr. On an old stump, only seen once and then sterile. Granite Falls, July 17, 1899, no. 565. 7. Usnea barbata (L.) FR. var. hirta Fr. On sandstone, rare. Mankato (Minneopa), June 27, 1899, no. 151. 8. Usnea barbata (L.) Fr. var. rubiginea Micux. On sandstone and granite rocks, rare. Mankato (Minneopa Falls), June 27, 1899, no. 152. Redwood Falls, July 8, 1899, no. 350. g. Theloschistes chrysopthalmus (L.) Norm. On trees and old boards, rare or infrequent except at Granite Falls, where the plant is frequent. Mankato, June 22, 1899, no. 9. Mankato (Rapidan), June 28, 1899, no. 163. New Ulm, July 4, 1899, nos. 226 and 227. Redwood Falls, July 6, 1899, no. 302, and July 8, 1899, no. 340. Granite Falls, July II, 1899, no. 387, and July 15, 1899, no. 549. 10. Theloschistes polycarpus (Euru.) Tuck. On trees and rocks, rare. Mankato, June 26, 1899, no. 107. Granite Falls, July 12, 1899, no. 447, and July 15, 1899, no. 531. 11. Theloschistes lychneus (Nyu.) Tuck. On trees and rocks, frequent. Mankato, June 22, 1899, no. 5. New Ulm, July 5, 1899, no. 263. Redwood Falls, July 8, 1899, no. 330. Granite Falls, July 11, 1899, no. 438, and July 17, 1899, no. 579. 12. Theloschistes concolor (Dicxs.) Tuck. On trees and old wood, common at the first locality and rare at the second. Mankato, June 22, 1899, no. 7. Granite Falls, uly ri rSo0, n0.-37'7- 310 MINNESOTA BOTANICAL STUDIES. 13. Theloschistes concolor (Dicxs.) Tuck. var. effusa Tuck. On trees rare. Mankato, July 1, 1899, no. 216a. 14. Parmelia perforata ( Jaca.) AcuH. On trees, rare. Mankato, June 26, 1899, no. 134. 15. Parmelia cetrata Acu. On trees and rocks, rare except at the last locality where the plant is frequent. New Ulm, July 4, 1899, no. 228. Redwood Falls, July 8, 1899, nos. 329 and 360. Granite Falls, July 12, 1899, no. 455, and July 17, 1899, nos. 550, 573 and 574. 16. Parmelia crinita Acu. On trees and granitic rocks, rare. Mankato, June 23, 1899, no. 47. Mankato (Minneopa Falls), June 27, 1899, no. 133- New Ulm, July 5, 1899, no. 258. Granite) Halls; ijolyaa. 1899, nos. 400 and 439. 17. Parmelia borreri TurN. On trees and granitic rocks, common. Mankato, June 22, 1899, no. 11. New Ulm, July 5, 1899, no. 288. Redwood Falls, July 8, 1899, no. 335. Granite Falls, July 11, 1899, nos. 383 and 389. 18. Parmelia borreri TuRN. var. rudecta Tuck. On trees and old wood, infrequent. Mankato, June 23, 1899, no. 12, and.) uly, 23, 1890, no. 58. 19. Parmelia borreri TurN. var. hypomela Tuck. On shaded granite rocks, rare and sterile. New Ulm, July 5, 1899, no. 287. 20. Parmelia tiliacea (Horrm.) FLoERK. On trees, infrequent. Mankato, June 23, 1899. 21. Parmelia saxatilis (L.) Fr. On trees and rocks, rare. Mankato, June 26, 1899, no. 106. New Ulm, July 5, 1899, no. 264. Redwood Falls, July 8, 1899, no. 341. Granite Falls, July 11, 1899, no. 414. 22. Parmelia saxatilis (L.) Fr. var. sulcata Ny. On old wood and shaded rocks, rare. Mankato (Rapidan), June 28, 1899, no. 165. Granite Falls, July 17, 1899, no. 586. 23. Parmelia saxatilis (L.) Fr. var. panniformis (Acu. (SCHAER.? On shaded rocks, rare. New Ulm, July 5, 1899, no. 268. Granite Falls, July 17, 1899, no 561. The plant is placed here provisionally. Fink: LICHENS OF THE MINNESOTA VALLEY. ol Not previously reported from Minnesota and new to the Mississippi valley. 24. Parmelia olivacea (L.) Ach. On trees and old wood, rare. Mankato, June 23, 1899, no. 61. Granite Falls, July 15, 1899, no. 537. 25. Parmelia olivacea (L.) Acu. var. prolixa Acu. On granitic rocks, quartzite, pipestone and once collected on earth, frequent except at the first locality, where it is rare. Morton, July 7, 1899, no. 315. Granite Falls, July 11, 1899, no. 405. Pipestone, July 18, 1899, nos. 594, 609 and 621, and July 19, 1899, no. 643. 26. Parmelia caperata (L.) Acu. On trees and granitic rocks, frequent. Mankato, June 23, 1899, no. 53. Mankato (Minneopa) June 27, 1899, no. 153. New Ulm, July 5, 1899, no. 285. Granite Falls, July 11, 1899, no. 441. 27. Parmelia conspersa (Euru.) Acu. On granitic rocks, quartzite and pipestone, common or abun- dant. New Ulm, July 5, 1899, no. 269. Granite Falls, July 12, 1899, no. 449. Pipestone, July 18, 1899, no. 589, and July 19, 1899, no. 637. 28. Physcia speciosa (WuLF., Acu.) NYL. On rocks and mossy bases of trees, infrequent. Mankato, Jume.22, 1899, no. 13. New Ulm; july 5, 1899; no. 28a. Granite Falls, July 11, 1899, no. 374. 29. Physcia granulifera (Acu.) Tuck. On trees, rare. Mankato, June 24, 1899, no. 84. Granite Falls, July 11, 1899, no. 372. 30. Physcia pulverulenta (ScuREB.) NYL. On rocks and trees, frequent. Mankato, June 23, 1899, no. 52. New Ulm, July 5, 1899, no. 286. Granite Falls, July rz, 1899. no. 383. 31. Physcia stellaris (L.) Tuck. On trees and rocks, common or abundant. Mankato, June 23, 1899, no. 1. New Ulm, July 5, 1899, no 297. Granite Falls, July 11, 1899 no. 431, and July 13, 1899, no. 481. 32. Physcia stellaris (L.) Tuck. var. apiola Nyv. On granitic rocks, infrequent. Mankato, June 23, 1899, no. 44. Granite Falls, July 12, 1899, no. 463. See MINNESOTA BOTANICAL STUDIES. 33. Physcia tribacia (Acu.) Tuck. On wood, granite and quartzite, rare. Mankato, June 23, 1899, no. 77, and July 1, 1899, no. 216. Granite Falls, July 14, 1899, no. 517. Pipestone, July 18, 1899, no. 601 and July 19, 1899, no. 634. 34. Physcia cesia (Horrm.) Nyt. On bowlders and all kinds of rocks in the region, except lime- stone, frequent. Mankato, June 23, 1899, no. 76. New Ulm, July 5, 1899, nos. 290 and 296. Granite Falls, July 12, 1899, no. 443. Pipestone, July 18, 1899, nos. 618 and 624. 35. Physcia obscura (Enru.) Nyt. On trees and rocks, common. Mankato, June 1899, no. 76a. New Ulm, July 5, 1899, no. 289. Granite Falls, July 11, 1899, no. 378, and July 17, 1899, no. 583. 36. Physcia adglutinata (FLoERK.) NyL. On trees, frequent. » Mankato, June 23, 1899, no. 45. Gran- ite Falls, July 11, 1899, no. 382 and July 13, 1899, no. 482. 37. Pyxine sorediata Fr. On granitic rocks, rare. Granite Falls, July 17, 1899, no. 578. 38. Peltigera rufescens (NEcK.) Horr. On earth and mossy rocks, frequent. Mankato, June 25, 1899, no. 48. Mankato (Minneopa Falls), June 27, 1899, no. 150. New Ulm, July 5, 1899, no. 251. Granite Falls, July II, 1899, no. 395 and July 17, 1899, no. 559. 39. Peltigera canina (L.) Horrm. On earth and rocks, common. Mankato (Minneopa Falls), June 27, 1899, no. 149. New Ulm, July 5, 1899, no. 262. Granite Falls, July 11, 1899, no. 390, and July 17, 1899, no. 580. 40. Peltigera canina (L.) Horrn. var. spongiosa Tuck. On earth, rare. Redwood Falls, July 8, 1899, no. 357. Not previously reported from Minnesota. 41. Peltigera canina (L.) Horr. var. spuria Acu. Onearth, rare. Mankato (Rapidan), June 28, 1899, no. 161. 42. Peltigera canina (L.) Horr. var. sorediata SCHAER. On earth and old wood, rare, Mankato, June 26, 1899, no. 121. New Ulm, July 5, 1899, no. 260. Granite Falls, July 13, 1899, no. 512. Fink : LICHENS OF THE MINNESOTA VALLEY. oles 43. Peltigera horizontalis (L.) Horrn. On shaded earth, frequent locally. Redwood Falls, July 6, 1899, no. 301. 44. Heppia despreauxii (Mont.) Tuck. On earth, rare at first locality and frequent at second. Man- kato (Rapidan) June 28, 1899. no. 177. Granite Falls, July 11, 1899, no. 394, July 13, 1899, no. 507 and July 14, 1899, no. 522). Not previously reported from Minnesota. 45. Heppia polyspora Tuck. ? On ‘earth, rare. Granite Falls, July 13, 1899, no. 4938: Spores spherical or subspherical, 2 8 mic. This exceeds . = Tuckerman’s measurements. Apothecia occasionally surpass- ing one mm. in diameter. I may later find it necessary to separate this as new species. Not previously reported from Minnesota. 46. Pannaria languinosa (AcH.) Korers. On various rocks, earth and trees in shaded places, common or abundant. Mankato, June 23, 1899, no. 50. New Ulm, July 5, 1899, no. 280. Granite Falls, July 11, 1899, no. 396, and July 17, 1899, no. 569. 47. Pannaria microphylla (Sw.) DeEtts. On shaded granite, infrequent. Redwood Falls, July 8, 1899, no. 345. Granite Falls, July 11, 1899, no. 384. 48. Pannaria nigra (Hups.) Nyt. On limestone, common locally. Mankato, June 24, 1899, no. 95. 49. Ephebe pubescens Fr. On quartzite, rare. New Ulm, July 5, 1899, no. 257. 50. Pyrenopsis pheococca Tuck. On bowlders, rare. Mankato, June 23, 1899, no. 74. Not previously reported from Minnesota and new to the Mississippi valley. 51. Pyrenopsis melambola Tuck. ? On bowlders, frequent. Mankato, June 29, 1899, no. 189. Spores somewhat small (7-10 x 4-5 % mic.). Not previously reported from Minnesota and new to the Mississippi valley. 314 MINNESOTA BOTANICAL STUDIES. 52. Omphalaria kansana Tuck. On limestone, locally frequent. Mankato, June 23, 1899, no. 27. Not previously reported from Minnesota. 53. Omphalaria pulvinata Nyt. On limestone, rare. Mankato, June 27, 1899, no. 148. Not previously reported from Minnesota. 54. Omphalaria phyllisca (WaAunL.) Tuck. On granitic rocks, rare. Redwood Falls, July 6, 1899, no. 360, and July 8, 1899, nos. 338 and 343. Granite Falls, July £7, 1599;, NOS. 572 and 50x: Not previously reported from Minnesota and new to the Mississippi valley. 55. Collema pycnocarpum Nyt. On trees and once on rocks, generally distributed in the Minnesota valley, but rare. Mankato, June 23, 1899, no. 60, and June 24, 1899, nos. 89 and 97. Mankato (Minneopa Falls), June 247, 1899, no. 136. Redwood Falls, July 6, 1899, no. 309, and July 8, 1899, no. 355- Granite Falls, July 11, 189g no. 380. 56. Collema flaccidum Acu. On trees and rocks, rare. Mankato, June 23, 1899, no. 81. Mankato (Minneopa), June 27, 1899, no. 137. New Ulm, July 5, 1899, no. 278. . 57. Collema pulposum (BERNH.) NyYL. On earth and rocks, common in first locality. Mankato, June 22, 1897; no. 3, and June 23, 1899; no.7o, 1 @mamie Falls, July 11, 1899, no. 417. 4 58. Collema tenax (Sw.) Acu. On earth, rare. Mankato (Rapidan), June 28, 1899, no. 169. Not previously reported from Minnesota. 59. Collema plicatile ScHAER. On calcareous rocks, locally frequent. Mankato, June 23, 1899, no. 28. Not previously reported from Minnesota. 60. Collema pustulatum Acu. On calcareous rocks, rare. Mankato, June 26, 1899, no. med. Not previously reported from Minnesota. Fink: LICHENS OF THE MINNESOTA VALLEY. 315 61. Collema furvum (Acu.) Nyt. On shaded rocks, infrequent. New Ulm, July 5, 1899, no. 283. Redwood Falls, July 6, 1899, no. 307. Granite Falls, July 11, 1899, no. 391. Not previously reported from Minnesota. 62. Leptogium lacerum (Sw.) Fr. On various rocks, usually shaded, frequent. Mankato, June 23, 1899, no. 49. Redwood Falls, July 8, 1899, no. 328. Granite Falls, July 13, 1899, nos. 480 and 497 and July 17, 1899, no. 568. 63. Leptogium pulchellum (Acu.) Nyt. Collected once on a large bowlder in a shaded ravine, rare. Mankato, July 1, 1899, no. 212. Not previously reported from Minnesota. 64. Leptogium chloromelum (Sw.) Nyt. On mossy, shaded sandstone, rare. Mankato (Minneopa Falls), June 27, 1899, no. 144. 65. Leptogium myochroum (Euru., ScHarER.) Tuck. On trees and shaded granitic rocks, rare. Mankato, June 26, 1899, no. 126. Mankato (Rapidan), June 28, 1899, no. coo. (‘Grauite Falls, July 11, 1899, no. 392. 66. Placodium elegans (Linx.) DC. On various rocks ; common at Granite Falls, infrequent else- where. Mankato, June 24, 1899, no. 83. Redwood Falls, July 8, 1899, no. 353. Granite Falls, July 11, 1899, no. 440. Pipestone, July 18, 1899, nos. 592 and 603. 67. Placodium murorum (Horrm.) DC. On granitic rocks, rare. Granite Falls, July 12, 1899, no. 452. 68. Placodium cinnabarrinum (Acu.) Auz. On various rocks, frequent or common. Mankato, June 23, 1899, no. 73, and June 30, 1899, no. 200. Morton, July 7, 1899, no. 320. North Redwood, July 10, 1899, no. 369. Gran- ite Falls July 11, 1899, no. 411. Pipestone, July 18, 1899, no. 607. 69. Placodium aurantiacum (Licutr.) Narc. and Hepp. On trees and rocks, common at first locality. Mankato, June 22, 1899, no. 19, and June 23, 1899. no. 38. Granite Falls, July 11, 1899; no. 375. 316 MINNESOTA BOTANICAL STUDIES. 70. Placodium citrinum (Horrno.) Lercur. On limestone, rare. Mankato, June 26, 1899, no. 117. 71. Placodium cerinum (HEpw.) Nagc. and HEpp. On trees and old wood, common. Mankato, June 22, 1899,. no. 18, and June 23, 1899, no. 75. Mankato (Rapidan), June 28, 1899, nos. 178 and 179. Granite Falls, July 11, 1899, nos. 432 and 435. Granite Falls, July 17, 1899, no. 553. 72. Placodium cerinum (Hrepw.) Narc. and HEpp. var. sider- itis Tuck. On granitic rocks and catlinite, common. Mankato, June 23, 1899, no. 41, June 26, 1899, no. 114, ands) ines2a,magn, no. 198. New Ulm, July 5, 1899, no. 247. Granite Falls, July 12, 1899, no. 444. Pipestone, July 18, 1899, no. 613. 73. Placodium cerinum(HrEpw.) Narc. and Hepp. var. pyracea NYL. On old boards, infrequent. Mankato, June 22, 1899, no. 2. Granite Falls, July 11, 1899, no. 430. 74. Placodium ferrugineum (Hups.) Hepp. On old wood, rare. Mankato, June 22, 1899, no. 17. Not previously reported from Minnesota. 75. Placodium ferrugineum (Hups.) Hepp. var. pollinii Tuck. On dead cedars, rare. New Ulm, July 4, 1899, no. 230. Not previously reported from Minnesota. 76. Placodium vitellinum (Euru.) NArc. and Hepp. On granite and quartzite, common. Mankato, June 30, 1899,' no. 203. New Ulm, July. 5, 1899; mo. 292.) "Granite Falls, July 12, 1899, no. 462. Pipestone, July 18, a6qo9,eae- 629. 77. Placodium vitellinum (Euru.) Narc. and Hepp. var. aurel- lum Acu. On granite, quartzite and sandstone, frequent. Mankato, June 22, 1899, no. 10. New Ulm, July 4, 1899, nos. 2335 237 and 239. Granite Falls, July 14, 1899, no. 523: 78. Lecanora sp. On granitic rocks, frequent locally. Granite Falls, July 11, —16 1899, no. 408. Spores = a mic. Seems near Lecanora . —— 5) gelida (L.) Ach., but the thallus is probably too rough and heavy. Fink : LICHENS OF THE MINNESOTA VALLEY. 317 Not previously reported from Minnesota and new to the Miss- issippi Valley. 79. Lecanora rubina (Vitu.) Acu. On granite, quartzite and pipestone, common. Mankato (Rapidan), June 28, 1899, no. 160. New Ulm, July 5, 1899, no. 248. Granite Falls, July 11, 1899, no. 4or and July 12, 1899, no. 451. Pipestone, July 18, 1899, no. 617, and July 19, 1899, no. 644. 80. Lecanora rubina (ViLL.) Acu. var. heteromorpha Acu. With the last, frequent. New Ulm, July 5, 1899, no. 266. North Redwood, July 10, 1899, no. 361. Granite Falls, July II, 1899, no. 409. Pipestone, July 18, 1899, and July 19, 1899, no. 642. 81. Lecanora muralis (ScHREB.) SCHAER. On calcareous rocks, granite and quartzite, common at Granite Falls, rare elsewhere. Mankato, June 24, 1899, no. 85. Granite Falls, July 11, 1899, no. 406. Pipestone, July 18, 1899, no. 630. 82. Lecanora muralis (ScHREB.) SCHAER. var. versicolor Fr. On calcareous rocks, rare. Mankato, June 30, 1899, no. 196. 83. Lecanora muralis (SCHREB.) SCHAER. var. saxicola SCHAER. On granitic rocks and catlinite, frequent. Mankato, June 30, 1899, no. 202. North Redwood, July 10, 1899, no. 370. Pipestone, July 19, 1899, no. 638. 84. Lecanora frustulosa (Dicxs.) Mass. On rocks, rare. Redwood Falls, July 8, 1899, no. 351. North Redwood, July 10, 1899, no. 371. Granite Falls, July II, 1899, no. 410. 85. Lecanora subfusca (L.) Aco. _ On trees and rocks, common at Mankato only. Mankato, June 23, 1899, no. 43, andJune 26, 1899, no. 125. New Ulm, ives, 2699, no. 271. Granite Falls, July 11, 1899;. no. 370- 86. Lecanora subfusca (L.) Acu. var. allophana Acu. On granitic rocks, infrequent. Granite Falls, July 11, 1899, nos. 403 and 407. Not previously reported from Minnesota: 87. Lecanora subfusca (L.) Acu. var. argentata Acu. On trees, rare. Mankato, June 23, 1899, no. 78. 318 MINNESOTA BOTANICAL STUDIES. 88. Lecanora subfusca (L.) Acu. var. coilocarpa Acu. On trees, granite and sandstone, rare. Mankato (Rapidan), June 28, 1899, no. 163a. Mankato, July 1, 1899, no. 210. Granite Falls, July 12, 1899, no. 446. 89. Lecanora subfusca (L.) Acu. var. distans Acu. On sandstone, rare. Mankato (Rapidan), June 28, 1899, no. 182. 90. Lecanora hageni Acn. On calcareous and granitic rocks and on old boards, common. Mankato, June 21, 1899, no gi, and July, 1, 1899, nos. 215, 217 and 218. Granite Falls, July 11, 1899, nos. 426 and 436, and July 13, 1899, no. 506. gi. Lecanora varia (EuRH.) Nyv. On old wood and trees, infrequent. Mankato, June 24, 1899, no. 86. Mankato (Rapidan), June 28, 1899, no. 159. New Ulm, July 5, 1899, no. 280. Granite Falls, July 11, 1899, no. 386. g2. Lecanora erysibe Ny. On granitic rocks, rare. Mankato, June 23, 1899, no. 63, and June 26, 1899, no. 131. Granite Falls, July 13, 1899, no. 504. 93. Lecanora cinerea (L.) SOMMERF. On granite, quartzite and catlinite, common. Mankato, June 6, 1899, no. 62. New Ulm, July 5, 1899, no. 261. Gran- ite Falls, ‘July 13, 1899, no. 493. Pipestone, July 18, 1899, nos. 625 and 633. 94. Lecanora cinerea (L.) SommeERF. var. levata FR. On quartzite, rare. New Ulm, July 5, 1899, no. 277. 95. Lecanora cinerea (L.) SomMERF. var. gibbosa Nyt. On bowlders, rare. Mankato, July 1, 1899, no. 221. 96. Lecanora calcarea (L.) SOMMERF. On limestone, rare. Mankato, June 29, 1899, no. 188. 97. Lecanora calcarea (L.) Sommerr. var. contorta FR. On limestone, drift pebbles and granite, infrequent. Man- kato, June 30, 1899, no. 199. Redwood Falls, July 8, 1899, no. 333. Granite Falls, July 11, 1899, no. 402, and July 14, 1899, no. 524. Fink: LICHENS OF THE MINNESOTA VALLEY. 319 98. Lecanora xanthophana Nyv. On granite, quartzite and pipestone, common. Mankato, June 30, 1899, no. 197. New Ulm, July §, 1899, no. 246. Morton, July 7, 1899, no. 313. Redwood Falls, July 8, 1899, no. 331. North Redwood, July 10, 1899, no. 368. Granite Falls, July 11, 1899, no. 418. Pipestone, July 18, 1899, nos. 598 and 611. 99. Lecanora cervina (Prers.) Nyv. On bowlders and sandstone, infrequent. Mankato (Rapidan), June 28, 1899, no. 179. Mankato, June 29, 1899, no. 192. 100. Lecanora cervina (PERs.) Ny. var. cinereoalba var. nov. On granite, frequent. Mankato, June 29, 1899, no. 190. Granite Falls, July 11, 1899, nos. 385 and 403, and July 12, 1899, no. 404. Thallus gray or grayish white. ror. Lecanora fuscata (ScuraAp.) TH. FR. On bowlders, common at Mankato. Mankato, June 29, 1899, no. I91. Granite Falls, July 12, 1899, no. 450. 102. Lecanora bookii (Fr.) Tu. Fr. On limestone, rare. Mankato, June 29, 1899, no. 193. Not previously reported from Minnesota and new to the Mis- sissippi valley. 103. Lecanora privigna (Acu.) Nyv. On sandstone and calcareous drift pebbles, rare. Mankato (Rapidan), June 28, 1899, no. 171. Granite Falls, July 13, 1899, no. 508. 104. Lecanora privigna (Acn.) Nyt. var. pruinosa AucrT. With last on same substrata, rare. Mankato (Rapidan), June 28, 1899, no. 170. Granite Falls, July 14, 1899, no. 514. 105. Rinodina oreina (Acu.) Mass. On granitic rocks, quartzite and catlinite, abundant. Man- kato, June 30, 1899, no. 201. New Ulm, July 5, 1899, no. 245. North Redwood, July 10, ‘1899, no. 367. Pipestone, July 18, 1899, nos. 602, 603 and 605. 106. Rinodina sophodes (Acu.) Nyt. On trees, old wood and rocks, abundant. Mankato, June 23, 1899, no. 33, June 24, 1899, no. 93, and June 26, 1899, no. 115. New Ulm, July 5, 1899, nos. 267, 295 and 298. Granite Falls, July 11, 1899, nos. 427 and 428, and July 13, 1899, nos. 469, 486, 487 and 4o1. 320 MINNESOTA BOTANICAL STUDIES. 107. Rinodina sophodes (Acu.) Ny. var. tephraspis Tuck. On quartzite, rare. Pipestone, July 18, 1899, no. 632. Not previously reported from Minnesota. 108. Rinodina sophodes (Acu.) Nyt. var. exigua FR. On old wood, locally common. Mankato, June 22, 1899, no. 22. Granite Falls, July 11, 1899, no. 434. 10g. Rinodina bischoffii (Hreppr.) Korres. On limestone and granite, rare. Mankato, June 29, 1899, no. 194. Morton, July 7, 1899, no. 316. Not previously reported from Minnesota. 110. Rinodina lecanorina Mass. On boulders, rare. Mankato, June 26, 1899, no. 127. Not previously reported from Minnesota and new to North America. 111. Pertusaria velata (TurRN.) NyL. On trees, rare. Mankato (Minneopa Falls), June 26, 1899, no. 135: 112. Pertusaria pustulata (Acu.) Nvu. On trees, rare. Mankato, June 23, 1899, no. 30, and July I, 1890, 110: 214. 113. Pertusaria leioplaca (AcH.) SCHAER. On trees, rare. Mankato, June 23, 1899, no. 68. 114. Urceolaria scruposa (L.) Nyt. On earth and rocks, infrequent. Mankato, June 26, 1899, no. 128. Mankato (Rapidan), June 28, 1899, no. 187. Red- wood Falls, July 8, 1899, no. 332. Granite Falls, July 11, 1899, no. 393. Pipestone, July 19, 1899, no. 640. 115. Urceolaria actinostoma Pers. On granite, rare. Granite Falls, July 11, 1899, no. 416. Not previously reported from Minnesota. 116. Stereocaulon paschale (L.) Fr. On mossy rocks, only seen once in small quantity. Red- wood Falls, July 8, 1899, no. 359. 117. Cladonia symphycarpia Fr. var. epiphylla (Acu.) Nyt. On earth, rare. Mankato, June 26, 1899, no. 108. 118. Cladonia mitrula Tuck. On earth, rare. Mankato, June 26, 1899, no. 98. Granite Falls, July 11, 1899, no. 436. The last a small form approach- ing Cladonia cespiticia (Pers.) FI. Fink: LICHENS OF THE MINNESOTA VALLEY. 321 119. Cladonia cariosa (ACH.) SPRENG. On earth, rare. Mankato, June 26, 1899, no. 103. Redwood Falls, July 8, 1899, no. 336. Granite Falls, July 17, 1899, no. 587. - 120. Cladonia pyxidata (L.) Fr. On earth, common or frequent. Mankato, June 26, 1899, no. 104. Mankato (Rapidan), June 28, 1899, no. 168. New Ulm, July 5, 1899, nos. 272 and 276, July 11, 1899, no. 397, July 12, 1899, no, 453, and July 17, 1899, no. 562. Pipestone, July 18, 1899, no. 627. 121. Cladonia fimbriata (L.) Fr. On earth, rare. Mankato, June 26, 1899, no 123. Granite Falls, July 12, 1899, no. 582. Pipestone, July 18, 1899, no. 604. 122. Cladonia fimbriata (L.) Fr. var. tubeformis Fr. On old wood and earth, rare. Mankato, June 26, 1899, no. 124. New Ulm, July 5, 1899, no. 279. Granite Falls, July II, 1899, no. 425, July 13, 1899, no. 495, and July 17, 1899, nos. 551 and 563. 123. Cladonia fimbriata (L.) Fr. var. radiata Fr. On earth, rare. Redwood Falls, July 8, 1899, no. 337. 124. Cladonia gracilis (L.) Nyv. On old wood and earth, frequent at Mankato, elsewhere rare. Mankato, June 22, 1899, no. 4, and June 26, 1899, no. Ioo. Granite Falls, July 13, 1899, nos. 468 and 488, and July 17, 1899, no. 556. 125. Cladonia gracilis (L.) Nyu. var. symphycarpia Tuck. On old wood, rare. Mankato, June 26, 1899, no. gg. 126. Cladonia gracilis (L.) Nyv. var. verticillata Fr. On earth, are Mankato June 26, 1899, no. 101. Granite Falls, July 17, 1899, no. 557. 127. Cladonia gracilis (L.) Ny. var. hybrida ScHAER. On earth, rare. Mankato (Rapidan), June 28, 1899, no. 158. Redwood Falls, July 8, 1899, no. 347. 128. Cladonia turgida (Enru.) Horr. On earth, rare. New Ulm, July 5, 1899, no. 253. 129. Cladonia cespiticia (PErRs.) FL. On earth, rare. Redwood Falls, July 8, 1899, no. 342. Granite Falls, July 17, 1899, no. 555. Sy MINNESOTA BOTANICAL STUDIES. 130. Cladonia furcata (Hups.) FR. On earth, rare. Mankato (Minneopa Falls), June 27, 1899, nos. 155 and 157. 131. Cladonia furcata (Hups.) Fr. var. racemosa FL. On earth in shaded places, rare. Mankato (Minneopa Falls), - June 27, 1899, no. 156. Redwood Falls, July 8, 1899, no. 334. 132. Cladonia furcata (Hups.) Fr. var. pungens FR. On earth, rare. Redwood Falls, July 6, 1899, no. 303. 133. Cladonia rangiferina (L.) Horr. On earth, frequent locally among granitic rocks. New Ulm, July 5, 1899, no..252. 134. Cladonia rangiferina (L.) Horr. var. sylvatica L. On earth, rare. Redwood Falls, July 8, 1899, no. 358. 135. Cladonia macilenta (Euru.) Horr. On old wood, rare. Mankato, June 29, 1899, no. 195. 136. Cladonia cristatella Tuck. On old stumps, rare. Mankato (Rapidan), June 28, 1899, no. 162. Redwood Falls, July 8, 1899, no. 356. Granite Falls, July 13, 1899, no. 467 and July 17, 1899, no. 575. 137. Cladonia cristatella Tuck. var. paludicola Tuck. Once collected on an old log. Mankato, June 26, 1899, no. 122. Squamules not powdery. Not previously reported from Minnesota, and new to the upper Mississippi valley. 138. Biatora decipiens (Enru.) FR. Common on earth containing calcareous drift pebbles. Granite Falls, July 13, 1899, no. 500. Not previously reported from Minnesota. 139. Biatora decipiens (EHRH.) FR. var. dealbata Auct. Common on earth with the last. Granite Falls, July 13, 1899, no. 499. Not previously reported from Minnesota. 140. Biatora icterica Monr. On earth, rare. Granite Falls, July 11, 1899, no. 398, and July 18, 1899, no. 519. 141. Biatora rufonigra Tuck. On granitic rocks and quartzite, common. New Ulm, July 5, 1899, no. 265. Morton, July 7, 1899, no. 325. Granite Falls, July 12, 1899, no. 454. Fink : LICHENS OF THE MINNESOTA VALLEY. 323 142. Biatora coarctata (Sm., Nyu.) Tuck. On limestone and sandstone, rare. Mankato, June 26, 1899, no. 113. Mankato (Rapidan), June 28, 1899, no. 173. 142a@. Biatora coarctata (Sm., Nyt.) Tuck. var. brugeriana, SCHAER. On sandstone, locally abundant. Mankato (Minneopa Falls), June 27, 1899, nos. 139, 142, 145 and 146. Mankato (Rapi- dan), June 29, 1899, nos. 172, 174 and 176. 143. Biatora uliginosa (SCHRAD.) FR. On earth, infrequent. Mankato, June 26, 1899, no. 128. New Ulm, July 5, 1899, no. 250. 144. Biatora myriocarpoides (Nyu.) Tuck. On quartzite, locally common. New Ulm, July 5, 1899, no. 300. 145. Biatora varians (Acu.) Tuck. On trees, probably frequent locally. Granite Falls, July 15, 1899, no. 502. 146. Biatora flexuosa FR. On dead cedar, rare. Granite Falls, July 13, 1899, no. 477. Not previously reported from Minnesota. 147. Biatora hypnophila (TurN.) Tuck. On earth and limestone, rare. Mankato, June 23, 1899, no. 36, and June 26, 1899, no. 120. 148. Biatora negelii Hepp. On trees, infrequent. Granite Falls, July 13, 1899, no. 484, and July 15, 1899, no. 530. 149. Biatora rubella (EHRH.) RABENH. On trees, common locally. Mankato, June 23, 1899, no. 35, and June 26, 1899, no. 130. Mankato (Minneopa Falls), June 27, 1899, no. 138. — 150. Biatora fuscorubella (Horrm.) Tuck. On trees and rocks, common at Mankato, else where rare or infrequent. Mankato, June 23, 1899, no. 29. Mankato (Min- neopa Falls), June 27, 1899, no. 141. New Ulm, July 5, 1899, no. 274. Granite Falls, July 15, 1899, nos. 529 and 536. 151. Biatora suffusa Fr. On trees, rare. Mankato, June 23, 1899, no. 37. Granite Falls, July 15, 1899, no. 540. Not previously reported from Minnesota. 324 MINNESOTA BOTANICAL STUDIES. 152. Biatora muscorum (Sw.) Tuck. On earth, frequent and once on sandstone. Mankato, July I, 3899, no. 220. New Ulm, July 4, 18099,-no7237>5 inca wood Falls, July 8, 1899, no. 344. Granite Falls, July 11, 1899, no. 433. Pipestone, July 18, 1899, nos. 590 and 591. 153. Biatora inundata Fr. On limestone and sandstone, common. Mankato, June 22, 1899, no. 24. New Ulm, July 4, 1899, no. 234. 154. Lecidea enteroleuca FR. On trees, common at Granite Falls. Mankato, June 23, 1899, no. 80, and July 1, 1899, no. 209. New Ulm, July 4, 1899, no. 231. Redwood Falls, July 8, 1899, no. 354. Granite Falls, July 11, 1899, no. 429, July 13, 1899, nos. 476, 479, 485 and 496, and July 14, 1899, no. 516. 155. Lecidea enteroleuca Fr. var. achrista SOMMERF. On trees, frequent. Granite Falls, July 13, 1899, nos. 471 and 475, and July 15, 1899, no. 546. 156. Buellia spuria (ScHAER.) ARN. On granitic rocks, quartzite and pipestone, frequent or com- mon. New Ulm, July 5, 1899, no. 294. Morton, July 7, 1899, no. 324. Granite Falls, July 12, 1899, nos. 445 and 458. Pipestone, July 18, 1899, no. 612. 157. Buellia alboatra (HorrM.) Tu. FR. On trees, especially U/mus, rare at first locality anil more common at second. Mankato, June 22, 1899, no. 15. Gran- ite Falls, July 11, 1899, no. 382, and July 17, 1899, no. 577. 158. Buellia alboatra (Horrm.) Tu. FR. var. saxicola Fr. On limestone, shaded, rare. Mankato, June 22, 1899, no. 16. Not previously reported from Minnesota. 159. Buellia parasema (Acu.) Tu. FR. On trees, infrequent. Mankato, June 23, 1899, no. 34, and June 24, 1899, no. 85. New Ulm, July 3, 1899, no. 224. Granite Falls, July 11, 1899, no. 388. 160. Buellia myriocarpa (DC.) Mupp. On old wood, common or frequent. Mankato, June 22, 1899, no. 20. Mankato (Rapidan), June 28, 1899, no. 184. Granite Falls, July 17, 1899, no. 552. Fink : LICHENS OF THE MINNESOTA VALLEY. 325 161. Buellia pullata Tuck. On rocks, frequent. Morton, July 7, 1899, no. 327. North Redwood, July 10, 1899, nos. 365 and 366. Granite Falls, July 11, 1899, no. 405. Pipestone, July 18, 1899, no. 600. Not previously reported from Minnesota. 162. Buellia turgescens (Nyv.) Tuck. On old boards, rare. Mankato, June 22, 1899, no. 21. _ Not previously reported from Minnesota. 163. Buellia petrea (FLor., Koers.) Tuck. On granite, quartzite and pipestone, abundant. New Ulm, July 5, 1899, nos. 242 and 293. Redwood Falls, July 6, 1899, no. 306. Granite Falls, July 11, 1899, no. 422. 164. Buellia petrea (FLor., Korrs.) Tuck. var. montagnei Tuck. On same rocks as last and even more abundant; however, only a single collection on a bowlder at first locality. Man- kato (Rapidan), June 28, 1899, no. 183. Morton, July 7, 1899, nos. 318 and 321. North Redwood, July 10, 1899, no. 362. Granite Falls, July 11, 1899, no. 422a. Pipestone, July 18, 1899, nos. 593 and 620. 165. Opegrapha varia (PeRs.) Fr. On trees, abundant. Mankato, June 23, 1899, no. 40, June 26, 1899, no. 110, and July 1, 1899, no. 204. Granite Falls, July 11, 1899, nos. 419 and 421, July 13, 1899, no. 483, and July 15, 1899, nos. 539 and 548. 166. Opegrapha varia (PEeRs.) Fr. var. pulicaris (Horr.) FR. On trees, rare. Granite Falls, July 15, 1899, no. 528. Not previously reported from Minnesota. 167. Graphis scripta (L.) Acu. On trees, common at Mankato. Mankato, June 22, 1899, no. 14. Granite Falls, July 13, 1899, no. 503. 168. Graphis scripta (L.) Acu. var. recta (HumB.) Nyt. On birches, rare. Mankato (Minneopa Falls), June 27, 1899, no. I41a. 169. Graphis scripta (L.) Acu. var. limitata Acu. On trees, rare. Mankato, June 22, 1899, no. 23. Granite Falls, July 13, 1899, no 469, and July 15, 1899, no. 538. 170. Arthonia lecideella Nyt. On trees, infrequent. Mankato, June 23, 1899, no. 69. 326 MINNESOTA BOTANICAL STUDIES. 171. Arthonia dispersa (ScHRAD.) Nyt. On trees, common. Mankato, June 22, 1899, no. 25, and July 23, 1899, no. 72. Granite Falls, July 13, 1899, no. 490. 172. Arthonia radiata (PERs.) Tu. FR. On trees, infrequent. Mankato, June 23, 1899, no. 70. Granite Falls, July 13, 1899, no. 489, and July 15, 1899, no. Bao. 173. Arthonia punctiformis Acu. On maples, rare. Granite Falls, July 15, 1899, no. 341a. 173a@. Arthonia sp. On trees, rare. Granite Falls, July 15, 1899, no. 541. With general appearance of Arthonia dispersa (Schrad.) 22-26 7-5-8.5 Nyl., but the colorless spores are four celled and mic. Not previously reported from Minnesota. 174. Calicium parietinum Acu. / On old wood, probably rare. Mankato, June 22, 1899, no. 8, and June 24, 1899, no. 87, Redwood Falls, July 8, 1899, no. 220; 175. Calicium quercinum PrErs. Collected once only, on cedar. Granite Falls, July 13, 1899, no. 478. 176. Coniocybe pallida (PERs.) FR. On a large oak, only once collected. Mankato, July 7 1899, no. 206. 177. Endocarpon miniatum (L.) ScHAER. Abundant on limestone bluffs, frequent on granite and rare on quartzite. Mankato, June 23, 1899 no. 46 and June 25, 1899, no. 59. Redwood Falls, July 6, 1899, no. 312. Granite Falls, July 11, 1899, no. 373, and July 13, 1899, no. 474. Pipestone: July 18, 1899, no. 606. 178. Endocarpon miniatum (L.) ScHAER. var. complicatum SCHAER. On substrata noted above and also on pipestone, frequent. Mankato, June 25, 1899, no. 57. New Ulm, July 5, 1899, no. 249. North Redwood, July 10, 1899, no. 363. Granite Falls, July 12, 1899, no. 448, and July 13, 1899, no. 495. Pipestone, July 18, 1899, no. 615, and July 19, 1899, no. 639. Fink: LICHENS OF THE MINNESOTA VALLEY. oot 179. Endocarpon fluviatile DC. On rocks frequently wet, infrequent. Morton, July 7, 1899, no. 322. Granite Falls, July 12, 1899, 448a. 180. Endocarpon arboreum ScHWEIN. On trees and shaded rocks, once seen on each. Redwood Falls, July 6, 1899, no. 308, and July 8, 1899, no. 339. Not previously reported from Minnesota. 181. Endocarpon hepaticum Acu. On earth and sandstone, common. Mankato (Rapidan), June 28, 1899, no. 175. New Ulm, July 4, 1899, no. 235, and July 5, 1899, no. 244. North Redwood, July 10, 1899, no. 364. Granite Falls, July 11, 1899, no. 384. Pipestone, July 18, 1899, no. 623. 182. Endocarpon pusillum Hepw. On limestone bluffs, sandstone, calcareous drift pebbles, and once on earth, common. Mankato, June 23, 1899, no. 39. New Ulm, July 4, 1899, no. 236. Granite Falls, July 12, 1899, no. 442, July 13, 1899, no. 509, and July 14, 1899, no. 526. 183. Endocarpon pusillum Hrepw. var. garovaglii Kpu. On earth and sandstone, frequent. Mankato, July 1, 1899, no. 219. Mankato (Rapidan), June 28, 1899, no. 186. New Ulm, July 5, 1899, no. 282. Morton, July 7, 1899, no. 317. Pipestone, July 18, 1899, no. 616. 184. Thelocarpon prasinellum Nyt. On old wood and sandstone, frequent. Mankato, June 22, 1899, no. 6, and June 26, 1899, no. 132. I cannot bring that on sandstone under any of the rock species, and it seems to belong here. 185. Staurothele umbrina (WauL.) Tuck. On granite, limestone and quartzite, frequent. Mankato, June 23, 1899, no. 82. Granite Falls, July 11, 1899, no. 412, July 15, 1899, no. 547, and July 17, 1899, no. 566. 186. Staurothele diffractella (NyL.) Tuck. On sandstone, granite, quartzite and calcareous drift pebbles, rare. New Ulm, July 4, 1899, nos. 238 and 240. Granite Falls, July 13, 1899, no. 501, and July 17, 1899, no. 560. Not previously reported from Minnesota. 328 Fink : LICHENS OF THE MINNESOTA VALLEY. 187. Staurothele drummondii Tuck. On granite, quartzite and pipestone, common in damp places at Granite Falls and Pipestone. Redwood Falls, July 8, 1899, no. 327. Granite Falls, July 12, 1899, no. 457, July, 13, 1899, no. 494, and July 17, 1899, no. 554. Pipestone, July 18, 1899, nos. 595, 619, 622 and 628. 188. Verrucaria fuscella Fr. On limestone, infrequent. Mankato, June 23, 1899, no. 42. 188a. Verrucaria nigrescens PERs. On limestone common and once seen on a granite bowlder. Mankato, June 23, 1899, no. 65, June 24, 1899, no. 96, and June 26, 1899, no. 116. 189. Verrucaria muralis Acu. On limestone in bluffs and drift pebbles, abundant at Man- kato. Mankato, June 22, 1899, no. 26. Granite Falls, July 13, 1899, no. 511, and July 14, 1899, no. 525. 190. Pyrenula punctiformis (Acu.) Nake. On trees, infrequent. Mankato, June 24, 1899, nos. go and 94. 191. Pyrenula punctiformis (Acu.) Nace. var. fallax Ny. On birch, infrequent. Mankato, June 24, 1889, no. 66, and June 26, 1899, no. 109. 192. Pyrenula gemmata (Acu.) Nazc. On trees frequent. Granite Falls, July 14, 1899, no. 513. Not previously reported from Minnesota. 193. Pyrenula hyalospora Nyv. On trees, probably rare. Mankato, June 23, 1899, no. 32, and June 25, 1899. Granite Falls, July 13, 1899, no. 470. Not previously reported from Minnesota. 194. Pyrenula nitida Acu. On trees, rare. Mankato, July 1, 1899, no. 211.. New Ulm, July 3, 1899, no. 222. 195. Pyrenula thelena (Acu.) Tuck. On birch, common. Mankato (Minneopa Falls), June 27, 1899, no. I40. 196. Pyrenula cinerella (FLor.) Tuck. On trees, infrequent. Mankato (Minneopa Falls), June 27, 1899, no. 143. Spores reaching 12-16 by 6-8 mic. in one collection. Thus larger than usual American forms. Fink : LICHENS OF THE MINNESOTA VALLEY. 329 197. Pyrenula cinerella (FLor.) Tuck. var. quadriloculata var. nov. On birch, probably rare. Mankato, June 26, 1899, no. 129. Mankato (Rapidan), June 28, 1899, no. 163a. Second time collected in Minnesota and both times from hosts of same genus. 198. Pyrenula quinqueseptata (Nyv.) Tuck. On trees, rare. Mankato, July 1, 1899, no. 208. Spores frequently showing 8 cells, which is not common for the species. Not previously reported from Minnesota. 199. Pyrenula leucoplaca (WALLR.) Kpr. On trees, common. Mankato, June 23, 1899, no. 31. Man- kato (Rapidan), June 28, 1899, no. 180. Mankato, July 1, 1899, no. 207. Granite Falls, July 13, 1899, no. 464, and July I5, 1899, nos. 527, 535 and 543. 200. Pyrenula glabrata (Acu.) Mass. On trees, rare. Mankato, June 24, 1899, no. 88. Not previously reported from Minnesota. 201. Pyrenula megalospora sp. nov. Thallus rather smooth, indeterminate, prominent, gray or grayish white. Apothecia scattered or occasionally aggregated in clusters of two or three, black or brownish black, convex with the ostiole-bearing apex somewhat pointed, semi-immersed or becoming more superficial, .4 to .75 mm. in diameter. Am- phithecitum white. Paraphyses capillary and very distinct. Asci cylindrical, .25 to .3 mm. in length. Spores colorless, 2- celled, oblong with ends obtuse or somewhat pointed, somewhat constricted at the septum, large for 2-celled spores of the genus (35-60 by 14-21 mic.), 8 in asci, crowded and obliquely uni- seriate. On trees, frequent. Mankato, June 26, 1899, no. 112 and July 1, 1899, no. 209. Granite Falls, July 11, 1899, no. 381 and July 11, 1899, no. 576. ee rT a5 * Le uepes WP: ¢ ifs a i PEAR aaa = Tie Pte aria ' e ee cea fe \ fa ory : : ee dh ry a : ; iy a oles py ee P ee | c* ty f yt ‘ | i : ae ) p We een \ ¢ A , ucla TITAS a S. 8 KA uth Or RA parity F ait’, , ieee iis hate ~ 71 RES IST ERR ex. As SYNONYMIC., CONSPECTUS, OF THE NATIVE AND GARDEN AQUILEGIAS OF NORTH AMERICA. hk Ce Davis. The name Aguzlegza (Linn. Sp. Pl. 533, 1753) is probably not from aguz/a, eagle, as commonly given, but from aquzlegus, water-drawer. The name may have been applied from the supposed power of the roots to extract water from rocks, among which they so often grow. They are commonly called Colum- bines. Hardy perennial herbs, mostly with paniculate branches ter- minated by showy flowers ; leaves 1-3 times ternately compound, commonly glaucous; leaflets roundish and obtusely lobed: flowers large, showy, appearing usually in spring or early summer; sepals 5, regular, petaloid; petals concave, produced backward between the sepals forming a hollow spur; stamens numerous; fruit of about 5 many-seeded follicles. About 30 species are distinct; all of the north temperate re- gions of the world. Nearly half of these (12) are natives of North America. Most of the native species and varieties are used in American and European gardens, and ten foreign spe- cies have already been introduced here. Aguz/egza furnishes many useful, ornamental forms eminently fitted for choice mixed borders and beds. 157. Var. nana-alba Horvt., A. flabellata var. flore-albo Horr. Flowers pure white and the plant dwarfish. A. leptoceras Fiscu. & Mryer. Ind. Hort. Petrop. 4: 33. ES A. brachyceras Turcz. ex F. & M. Maund. Bot. Gard. no. 755. About 1842. Stem several flowered, about 1 ft. high: partial-petioles of root-leaves over I inch, leaflets sessile; stem-leaves petioled, biternate: flowers violet, with the tips of the sepals greenish, and tips of the short petal-limbs yellow; spur slender, slightly curved, ¥% inch long, not knobbed; stamens protruding a little beyond the limbs of the petals: follicles slender, glabrous, nearly r inch long. Summer. East Siberia. Bot. Reg. 33: 64. Flor. des Serr. 3: 296. —Little used in America. A. vulgaris Linn. Sp. Pl. 533. 1753. wendersa Minw. Gard. Dict..Svedz no. 3.\ 1768. . silvestris NEck. Delic. Gallo-Belg. 1: 234. 1768. . cornuta GiiB. Fl. Lituan, 2: 286. 1781. . versicolor SALISB. Prod. 374. 1796. . corniculata Viti. Cat. Hort. Strasb. 250. 1807. 4 sellaia Wort) ex Steud. Nom: 1 ed. 61.1822; . elata LEDEB. Ind. Hort. Dorp. Suppl. 41. 1824. airala Koc. in) Flora, 132 1. 118. * 1830. Baa AAA AA 338 MINNESOTA BOTANICAL STUDIES. A. nigricans Baume. Enum. Stirp. Trans. 2: 104. About 1830. A. Haenkeana Kocn. Syn. Fl. Germ. 23. 1837. A. concoler Fiscu. ex Steud. Nom. 2 ed. 1: 115. 1840. A. ecalcarata Hort. ex Steud. l. c. 115. A. elegans Pore ex Steud. 1. c. 115. A. stbtrica Don ex Steud. 1. c. 115. A. subalpina Bor. FI. Centr. Fr. 3 ed. 2: 24. 1840. A. Bernardi Gren. & Gopr. Fl. Fr. 1: 45. 1848. A. Transsilvanica Scour. Vehr. Siebenb. Ver. Nat. 3: 94. 1852. A. sylvestris Scuur. Vehr. Siebenb. Ver. Nat. 4: 4. 1853. A. glaucophylla StTEwp. in Flora, 39: 407. 1856. A. aggericola Jorv. Diagn. 1: 87. 1860. A. collina Jorv. |. c. 84. A. dumeticola Jorn. |. c. 86. A. praecox JoRD. 1. c. 85. A. paraplesita Scour. Enum. Pl. Trans. 28. 1866. A. corsica SOLIER. ex Nym. Consp. 18. 1878. A glaucescens BAKER, l. C. 203. A. Karelint BAKER l. c. 76. A. subscaposa Boruas in Magyar Tudom. Akad. 12, IV, ris peti Coley A. platysepala Reicus. Ic. Fl. Germ. ¢. 4730. About 1886. Stem 1%-2 feet high, many flowered; finely pubescent throughout: root-leaves with 3 partial-petioles 1%,—-2 inches long, secondary branches certain, ultimate leaf lobes shallow and roundish, texture firm; lower stem-leaves petioled and bi- ternate: flowers violet to dark purple; sepals ovate, furnished with a claw, acute, I inch long, one-half as wide; petal-limb 34 inch long, equalling the head of stamens; spur about same length, stout, much incurved, knobbed: follicles densely pubes- cent, r inch long, style half as long. Summer. Europe, Siberia, and naturalized in America. Garden 12, p. 288. Var. nivea Baume. ex Baker Gard. Chron. II, 10: 76. 1878. Var. alba Hort. Often 2-3 feet high: a great profusion of large, pure white flowers. Several weeks in early spring. Var. flore-pleno Horr. ... ...0:.0g;es5 eee Lycoctonum. BB. Leaves divided to the base. C.Melmet hieher than wide\....2.......-3eeeeee variegatum. CC.. Helmet, or hood, broad and low. D. Follicles rarely varying from four........... Na pellus. EMD:; Follicles three to tive... s2.eee ee delphintfolium. AA. Roots fascicled and elongated, or fibrous. B. Stems erect. C,. Blowers yellowishis.20::.0.4.s0-2)