rir +E A Vv UT a GRAY HERBARIUM OF HARVARD U sit ANY U NIV!I CRSITY CON ual CONTRIBUTIONS Nos. 185-191 1959-62 ipo ite BorTaANicak ARDEN LIBRAR Rx CONTRIBUTIONS FROM THE GRAY HERBARIUM OF HARVARD UNIVERSITY Edited by Reed C. Roilins and Robert C. Foster NO. CLXXXV THE WILLOWS OF BOREAL WESTERN AMERICA By Huecu M. Raup SPORANGIA OF THE FERN GENERA ALLIED © WITH POLYPODIUM AND VITTARIA Br Kenneth A. WILSON THE SOMATIC CHROMOSOMES OF RUDBECKIA AND RELATED GENERA OF THE COMPOSITAE By € Ropert E. Pexpvr, Jn. Published by _ THE GRAY HERBARIUM OF HARVARD > UNIVERSITY oe a CAMBRIDGE, » MASS., U. SA A a CONTRIBUTIONS FROM THE GRAY HERBARIUM OF HARVARD UNIVERSITY Edited by Reed C. Roilins and Robert C. Foster NO. CLXXXV THE WILLOWS OF BOREAL WESTERN AMERICA BY HuGu M. Raup SPORANGIA OF THE FERN GENERA ALLIED WITH POLYPODIUM AND VITTARIA By KENNETH A. WILSON THE SOMATIC CHROMOSOMES OF RUDBECKIA AND RELATED GENERA OF THE COMPOSITAE By RoBERT E. PERDUE, JR. Published by THE GRAY HERBARIUM OF HARVARD UNIVERSITY CAMBRIDGE, MASS., U.S. A 1959 Issued March 27th 129 THE WILLOWS OF BOREAL WESTERN AMERICA By HuGuH M. Raup CONTENTS Introduction 3 Problems in the definition and arrangement of species .................+. Y Hybrid willows ..... 15 Notes on the geographic distribution of the species ............:::000+ 15 Notes for an arrangement of species in terms of form, size, and habitat preference 19 The collection of specimens 21 The use of this paper 23 Acknowledgments 27 Key to species and groups, based upon well-developed female flowering or fruiting specimens 28 Key to species and groups, based upon sterile leafy plants exclusive of rank-growing sprouts 33 Description of the species 39 Literature cited 93 INTRODUCTION The present paper is designed primarily for the descrip- tion and identification of the willows known to occur in the northern part of North America west of Hudson and James Bays. The southern limits of the area treated form an ir- regular line connecting the southern end of the Alaska Pan- handle, the upper Peace River, Lesser Slave Lake, Atha- baska and McMurray on the Athabaska River, thence run- ning eastward across northern Saskatchewan and Manitoba to about long. 96°W., and southeastward from there to in- clude the southwestern shores of James Bay. It will also cover present knowledge of the willows of the western arc- tic islands. Ina paper published in 1943, I covered to the best of my knowledge the willow flora of boreal America westward to include the western shores of Hudson and James Bays, and 4 HUGH M. RAUP to include that part of the District of Keewatin that lies in the basins of the Kazan River and Baker Lake, and in the basin of the Dubawnt River up as far as Dubawnt Lake. In northern Manitoba and Ontario that treatment covered the country westward approximately to long. 96°W. and south- ward to about lat. 51°. The areas just noted, therefore, bor- dering Hudson and James Bays on the west, constitute a region of overlap between my earlier paper and the present one. I have drawn freely from the works of several recent stu- dents of northwestern American flora, notably those of Dr. A. E. Porsild and Dr. Eric Hultén. Porsild has traveled and collected widely in the Northwest Territories, Yukon and Alaska. His published papers contain a wealth of material pertinent to the present study (1943, 1945, 1951, 1955, 1957). The most comprehensive treatment of any large proportion of the northwestern American willows is in Hultén’s Flora of Alaska and Yukon (1942, 1949). These papers, particularly that of Hultén, are so comprehensive, and cover so well the knowledge of the genus in the regions treated up to the times at which they were written, that I have used them as a sort of “datum plane” for the present study. There are, however, many instances in which I have differed from them in their interpretations of species and relationships, and on these occasions it has been necessary to go back into some of the earlier literature. Further materials are in my own studies of the vascular plants of the southern and central parts of the Mackenzie drainage basin (1934, 1935, 1936, 1942, 1947), and addi- tional specimens and notes gathered in the years 1943, 1944 and 1948 by my field associates and myself along the Alaska Highway between Dawson Creek in east central British Columbia, and Fairbanks in central Alaska. For data on the eastern part of the area, I have used for the most part my own study of The Willows of the Hudson Bay Region and the Labrador Peninsula, and a paper by Dr. H. J. Scoggan of the National Herbarium of Canada on The Flora of Manitoba (1957). Many suggestions for range exten- sions have come from the excellent series of manuscript maps accumulated by Dr. A. E. Porsild of the National nave been available to me, and have been of e for the filling in of ranges and general knowl- e species. Of particular note are those of the late Il or more or less localized collections made in. THE WILLOWS OF BOREAL WESTERN AMERICA 5 J. P. Anderson between Tanacross and Whitehorse in 1944, of Dr. C. H. D. Clarke in the vicinity of Kluane Lake and along the Haines Road in 1943 and 1944, of M. P. and R. T. Porsild at Johnson Crossing, Whitehorse, and on the Haines Road in 1944, of Dr. J. E. Cantlon and his associates along the Lower Colville River in 1953, and of Dr. T. M. C. Taylor and his associates along the Alaska Highway between Dawson Creek and Haines Junction, and along the Haines Road, in 1956. Likewise I have had access to the recent collections of Dr. H. J. Secoggan in northern Manitoba, and to those of Dr. W. K. W. Baldwin in northern Ontario and Manitoba. Most of the herbarium material studied has been found at Harvard (GH, A), the New York Botanical Garden (NY), and especially in the rich collections of the National Museum of Canada at Ottawa (CAN). Other herbaria consulted are at the Bailey Hortorium at Cornell (BH), the Philadelphia Academy of Science (PH), the Universities of British Columbia (V) and Alberta (ALTA), the U. S. National Museum in Washington (US), the Department of Agriculture (Science Service) of Canada at Ottawa (DAO), the Naturhistoriska Riksmuseet at Stockholm (S), the British Museum (BM), Kew (K), and the University of Copenhagen (C). The keys and dispositions of species have been strongly influenced by the work of Schneider, Ball, and Fernald on American willows during the past forty years. The only recent attempt at a complete monograph of the American species is that of Dr. Camillo Schneider, published between 1918 and 1921. The basic organization of my treatment of our willows, and much of the structure of the keys, are de- rived from Schneider’s monograph. Material from Dr. M. L. Fernald’s studies comes mainly from his treatment of the genus in the 8th edition of Gray’s Manual (1950), though frequent reference has been made to several of his earlier papers published in Rhodora. Several papers by Dr. C. R. Ball and Dr. P. A. Rydberg have been used freely, and will be referred to in appropriate places in the text. The inherent variability within species of willows makes definitive keys difficult to construct. Many species must be reg , not on single unit characters, but often upon - elaborate combinations of characters. Thus the keys must be in large measure descriptive. Further, the grouping of yecies into natural sections is at present inadequate, so that synoptic keys are less workable than purely artificial 6 HUGH M. RAUP ones. I have made no attempt at a key to male specimens. This is due to the fact that our knowledge of variability in the male flowers is poorly developed, and to the fact that for several species no male flowers have ever been collected. The key to sterile specimens, especially, is far from satis- factory. Its use must be restricted to well-developed, normal leafy material, for it does not take into account the often abnormally large and unusually-shaped leaves of rank- growing sprouts. The fickleness of vegetative characters in the willows is well illustrated by this key. Any major sub- division that might be chosen would leave a number of species in which the characters used would be combined: leaves serrate vs. entire, green beneath vs. glaucous be- neath, pubescent vs. glabrous, obovate or oblanceolate vs. ovate or lanceolate, etc. Many species must therefore ap- pear at several points in the key, probably at more places than I have given them. Such difficulties pyramid as the willow flora becomes larger. For small floras, containing twenty-five species or less, keys to sterile material are relatively simple. Fully as troublesome as the one I have attempted here is that of Schneider for all the American willows known to him at the time of his studies (1921: 107- 16). The terminal points in the main keys are in some cases individual species, and in others groups of species. Keys to the species within these groups will be found in the ensuing text immediately preceding the description of the first species named in each group. No attempt has been made to give complete citations of Synonyms, or of the many papers in which lists of willows or discussions of species have been published. Likewise I have omitted the citation of a great many of the specimens that I have seen in the preparation of the paper. However, comparable with the more recent papers on the willows of the region. Literature necessary to discussions of the author, year and page, and will be found listed alphabeti- i S ° * by my own field parties in the vicinity of the Alaska High- wey i I A ae I THE WILLOWS OF BOREAL WESTERN AMERICA 7 PROBLEMS IN THE DEFINITION AND ARRANGEMENT OF SPECIES Salix is abundant throughout the whole region, not only in number of individuals, but also in number of species. Its species make up a major part of the shrubby flora of mus- kegs, prairie and swamp margins, open woods, and tundra. The genus is notorious for the variation that occurs within species, and for poor definition of morphological bounda- ries between many of the commonly recognized species. In the present treatment I have recognized 98 species and lesser categories. Of the 98 taxa, 61 have been given speci- fic names A century and a half of study in so variable a group of plants as the American willows, together with a strong tendency to use a binomial system of nomenclature, have led inevitably to the description of many species of un- equal taxonomic soundness. Much of the reputation of the willows as a difficult and frustrating genus to work with __ terra incognita to most botanists and collectors — is due to lack of understanding of relationships among the many microspecies that have been described and given binomials. If it were possible to make a logical and widely accepted synoptic arrangement of the species in well-defined sections or subgenera, some of the confusion could be resolved. But the subgenera of Salix are not clearly defined, nor is there general agreement on how they should be constituted. S. interior, S. Setchelliana, S. polaris subsp. pseudopolaris, S. herbacea, S. glacialis, S. Turnorii, S. Farrae, S. cordata, S. commutata, S. Chamissonis, S. pyrifolia, S. Barrattiana, S. amplifolia, S. candida, S. Bebbiana, S. fallax, S. gracilis, S. arbusculoides, S. sitchensis. Some of them, such as S. Setchelliana, S. Turnorii, S. amplifolia, and possibly S. fallax, are highly localized or more widespread endemics so far as present knowledge goes. Others, such as S. Bebbiana, are among the commonest and most wi stribute Se region. Most of the variants that have been _ 8 HUGH M. RAUP described within these well-defined species have proved to have poorly-defined geographic segregation or none at all. None of the remaining 41 species can be isolated in the above manner. They can, however, be arranged in groups, each containing two or more closely related species whose ranges commonly overlap, and among which intermediates are common. It is unknown whether these intermediate forms are the result of introgressive hybridization, or are brought about by genetic segregation within large species populations rich in biotypes. Usually the species included in the groups show fairly well-defined geographic distribu- tional patterns, while minor variants within the species have none or very poorly-defined ones. A few of the groups, such as those related to S. glauca and S. rigida, would be en- larged by additional species if a larger geographic area were involved, but I have included only the forms known to occur in the region upon which this paper is based. The following arrangement of the 41 species into 16 groups is tentative, intended to be merely suggestive of what I believe to be the probable relationships. If put into practice it would involve a trinomial nomenclature, or quad- rinomial if minor varietal trends were recognized. The spe- cies themselves (as treated in the present paper) would become subspecies within the groups. Although in a few cases the relationships seem sufficiently clear at present to justify setting up such a nomenclature, in most of the groups more detailed study is needed before this can be done properly. Therefore I pee to leave the arrangement tentative for the time bein: Salix lucida group: S. rae S. lasiandra, S. serissima. Salix reticulata group: S. reticulata, S. vestita. Salix rotundifolia group: S. rotundifolia, S. Dodgeana, S. phle- bophy Ua. Baie aries Ereoe: S. arctica, S. arctophil. Salix group: S. ovalifolia, S. stolonifera,S. S. os Salix glauca group: S. glauca, S. niphoe: oclada, S ; f Salix MacCalliana group: S. MacCalliana, S. Tyrrellii. mit: Makendie orom up: SB akin wih 3 ‘group: S. sili Salix poe eee group: “ preety Ss. pedicellaria, S. atha- - Salix discolor group; S. discolor, 'S. Scouleriana. — : te nhvticifol s pug oe ES Salix pellita group: —— = —— EC ap AR ane gr aaa Ee ee ee eer eee O Te ne en nn eee, Ve eee THE WILLOWS OF BOREAL WESTERN AMERICA 9 The 16 species-groups may be arranged in five main categories. First are those whose segregates are eastern and western in the northern part of the continent, with well-defined areas of overlap among them. An arctic-alpine subcategory of these is represented by three groups. The S. reticulata group (Map 3),’ in which S. reticulata itself has a wide transcontinental range, contains also S. vestita which is primarily eastern arctic and subarctic. The latter has also an isolated area in the Rocky Mountains outside the range of the former. The two overlap on the outer Labrador coast and in the southern Hudson Bay region. The S. arctica group (Map 7) likewise has a wide-ranging element (S. arctica, sens. lat.) and an eastern element (S. arctophila) which has its largest development in the eastern Arctic but has a western extension to northeastern Alaska. These two have a large area of overlap in the eastern Arctic. The S. Richardsonii group (Map 18) has a pattern of ranges somewhat similar to that of the S. reticulata group, but S. Richardsonii does not have so extensive a range in the eastern Arctic, and the area of overlap between it and S. calcicola is limited to the northwestern shores of Hudson Bay and southern Baffin Island. However, like S. vestita, S. calcicola has an isolated area in the Rocky Mountains where apparently it has no contact with S. Richardsonii. The second subcategory is composed of four groups which are more or less confined to the fore regions. First among these is the S. lucida group (Map 1) in which but they are connected through the closely related S. seris- sima. The S. Barclayi group (Map 14) asa whole is trans- continental, with S. myrtillifolia the more wide-ranging of the two species. There is a large area of overlap in the 1 qt should be emphasized that the maps presented here are generalized ‘based maps showing collection localities by dots, and are made by connecting the dots that mark the outside limits of the ranges. With the exception of maps 8 and 9, the base map used is by J. Paul Goode, copyrighted and published by the University of Chicago Press Cos 10 HUGH M. RAUP northern Rocky Mountains, Yukon, and eastern Alaska. Salix glaucophylloides and S. padophylla (Map 15) appear to be, respectively, eastern and western variants in a single population of closely related willows. They have an area of overlap in northwestern Ontario and eastern Manitoba. Salix rigida is a strictly eastern representative of its group (Map 12), extending westward approximately to central Ontario. There it overlaps with S. lutea, showing so many intermediate forms that the two species can scarcely be separated. The westernmost element, S. mackenzieana, is primarily an interior species in our region, overlapping S. lutea in the eastern foothills of the northern Rockies and the southern Mackenzie basin. Salix discolor and S. Scou- leriana are well-defined eastern and western segregates in the S. discolor group (Map 25), with a region of overlap in Alberta, central Saskatchewan, and western Manitoba. The third subeategory is a more complex one, involving ranges that are in both forested and arctic-alpine areas. The most elaborate of these is in the S. glauca group (Map 10). Salix glauca (sens. lat.) has the widest range in this group, occupying both forest and tundra. It has its great- est range expansion in the west, but has a long extension eastward to southern Baffin Island. In the Hudson Bay country it overlaps the eastern S. cordifolia (sens. lat.), and there is a welter of intermediate forms (particularly in what has been called S. cordifolia var. callicarpaea). Salix niphoclada seems to be a northern arctic and subarctic segregate, overlapping S. glauca in much of interior Alaska, Yukon, northern British Columbia, and the District of Mackenzie. However, it extends farther into arctic Canada and the southern arctic islands than any other member of the S. glauca group. Salix brachycarpa (sens. lat.) appears to be a more southern segregate in the group, reaching northward not far beyond the timberline, and extending _ into the central Rockies of the United States. It is of inter- est to note that there is a region in the District of Keewa- tin where all four of the segregates in the S. glauca group come close together and probably overlap their ranges. An- other complex group is that related to S. arbutifolia (Map 23). The largest element here is S. pedicellaris var. hypo- _ glauca which ranges westward in the forested country _ from eastern Quebec to the Rocky Mountains and the cen- tral Mackenzie River valley. Salix arbutifolia, on the other _ hand, is a more arctic species with its largest devel ; THE WILLOWS OF BOREAL WESTERN AMERICA 11 ey, Maem 7 Sak KC ie ee fins rhe diet District of Keewatin. Present knowledge of the ranges provides no overlap of these two species, though they are close together in the District of Mackenzie. However, vind appear to be connected through S. athabascensis in Y and eastern Alaska. The fourth et heen S hebecarpa, is known only in two isolated localities: one on the Gaspé Peninsula, where it is surrounded by the S. pedi- —* — and the other on the Seward Peninsula = HUGH M. RAUP aol a ‘i i ‘ 3 aoe 2 ris Ph oi ae LY dlr : | Cas aly e- =< ae Far ye ‘Ni A Ie a D = ‘ 5 ‘ oe ra ot ae val SR iin tae aN bie Lei ' Li, er ae . a : of hia within the range of S. arbutifolia. Salix plani- . ee folia is a wide-ranging eastern element in the S. phylicifolia a faced buer'y sear ee thai peri westward in the forested 7 THE WILLOWS OF BOREAL WESTERN AMERICA 13 fgets Wie - mG Ye y : \ v _ . ’ 2 ae Jong ily ae: { Ne OS 3 a2 y i” " ot BS MC he! range of its far nortl S. pulchra. Salix phylicifolia, — Eurasian species included’ in our flora with some hesitati known only i in ene eon Yukon and along the iow Kusk 14 HUGH M. RAUP The second category of ranges is represented only by the two species in the S. pellita group (Map 27). Salix pellita is a species of the forested country, ranging from New- foundland to central Saskatchewan. Salix subcoerulea is characteristic of the northern Rocky Mountain region an reaches northwestward into the upper Liard River valley. These two species are so closely related as almost to defy taxonomic separation, but apparently their ranges do not overlap in Alberta or Saskatchewan. Further collection in the region between them may fill in the gap. A third category is made up of the S. ovalifolia group (Map 8), with relatively narrow, overlapping ranges con- fined to the northwestern coastal strip. The most extensive of these ranges is that of S. flagellaris (incl. S. arctolitor- alis), along the Arctic and Bering Sea coasts from the Mac- kenzie delta to the Aleutians, and southeastward along the Alaska panhandle. Salix ovalifolia extends along the coast from Cape Lisburne to the Aleutians, and has an isolated locality at White Pass. Salix stolonifera is primarily south- ern in Alaska, reaching from the Aleutians to the southern The latter extends from central British Columbia eastward _ to Hudson and James Bays. Saliz MacCalliana and S. Tyr- THE WILLOWS OF BOREAL WESTERN AMERICA 15 rellii are related to the S. glauca group, but their vegetative habit is so distinctive that I have maintained them in a small group by themselves. HYBRID WILLOWS There has been a tendency among some modern students of willows to recognize a multitude of natural hybrids. Of necessity such interpretations must be based upon the in- spection and comparison of specimens; and in most cases the comparisons are based upon a small number of speci- mens, so that modern methods for the statistical study of variation in form, and of introgressive hybridization, are not possible. Under these circumstances it seems to me less confusing and more useful for further study to attach intermediate specimens to the species they most nearly resemble, than to set them aside as “inspection hybrids” _— essentially nameless waifs that are apt to be missed by monographic or floristic students, and that may or may not actually be hybrids. The recognition of hybrids among the willows by simple comparison of a few specimens, if carried to its logical conclusion, can lead to utter taxonomic confusion where this is not warranted. From seeing two species as parents of a given specimen, it is a short step to seeing three, or even four or five; and if this is done con- sistently, many of the species boundaries in Salix can be broken down, so that a large proportion of the plants be- come hybrids. But this submerges the realities, for most of the great species complexes in Salix are real. For the pres- ent it seems best to preserve as much of this order as possible. NOTES ON THE GEOGRAPHIC DISTRIBUTION OF THE SPECIES Within the area covered by the present treatment, and with present knowledge of species ranges, the Territory of Yukon has the largest number of species (39). Northern British Columbia has 37, Alaska 37 and District of Mackenzie 35. Eastward and northeastward in the North- west Territories the numbers decline: Keewatin 18, and the western arctic islands (in the District of Franklin) 12. wai 27, northern Manitoba 28, and northern Ontario 24. Lists of species for each of these geographic divisions are : el ‘ oe. dae, 16 HUGH M. RAUP It should be emphasized that these lists, as well as those on species ranges that follow them, represent only present knowledge of the ranges so far as I have been able to gather it together. They are by no means definitive, and will with- out doubt be added to as more collections accumulate, or subtracted from as our concepts of the species change. There are still huge areas in the Northwest from which we have no plant collections at all; and even in those from which we have a great many the willow flora has been gathered critically in only a few widely separated localities. Some examples will illustrate. It is probable that as the Mackenzie Mountain system is further explored, more species of the Rocky Mountains will be found there, as well as more of the species that we now think of as peculiar to Alaska or neighboring Yukon. A few records suggest that more eastern arctic plants will be found there also. At the same time, further study of such difficult species complexes as the S. rotundifolia, S. ovalifolia, S. glauca, and S. arbu- tifolia groups may well alter our definitions of the species used in the present geographic arrangement. Any of these things can alter the definitions of ranges, and change the apportionment of species among the territories and provin- ces. What has been said about the Mackenzie Mountains applies also to the great area of overlap between the eastern and western species of the forested country, in the north central plain of the continent. It applies as well to many smaller areas from which critically made collections are scanty. ALASKA: S. lasiandra, interior, reticulata, Setchelliana, polaris udopolaris, rotundifolia, phlebophylla, arctica, arctophila, ovalifolia, stolonifera, flagellaris, glacialis, glauca, niphoclada, Farrae, Barclay, myrtillifolia, padophylla, commutata, Chamissonis, Richard- sonii, Barrattiana, amplifolia, alaxensis, candida, Bebbiana, fallax, arbutifolia, pedicellaris, athabascensis, hebecarpa, Scouleriana, phy- licifolia, pulchra, arbusculoides, sitchensis. YUKON: => lasiandra, interior, reticulata, Setchelliana, polaris. subsp. pseudopolaris, rotundifolia, phlebophylla, arctica, aretophila, See stolonifera, flagellaris, glauca, niphoclada, hy : cordifolia, mackenzieana, Farrae, Barelayi, myrtillifolia, padophylla, commutata, Chamissonis, Richardsonii, Barrattiana, alaxensis, can- : dida, Bebbiana, fallax, arbutifolia, pedicellaris, athabascensis, Scoul- eriana, — planifolia, pulchra, subcoerulea, arbusculoides, S58 MACKENZIE: S. lasiandra, se serissima, interior, reticulata, polaris Hs =e onstage Llgonts Dodgeana, arctica, arctophila, flagel- THE WILLOWS OF BOREAL WESTERN AMERICA 17 ie aa ie athabascensis, gracilis, Scouleriana, planifolia, pulchra, KEEWATIN: S. reticulata, herbacea, arctica, arctophila, n iphoclada, brachycarpa, cordifolia, pacha myrtillifo lia, a ia, Richardsonii, ealeicola, alaxensis, candida, arbutifolia, planifoli arbusculoi ides WESTERN FRANKLIN: 8S. reticulata, polaris hice, pseudopolaris, herbacea, arctica, arctophila, aN niphoclada, cordifolia, Richard- sonii, calcicola, alaxe ee eee ia. NORTHERN BRITISH COLUMBIA: S. lasiandra, interior, reticulata polaris, subsp. pseudopolaris, phlebophyla aretica, saahiolia: stolonif- era, flagellaris, glauca, ntphoclada, brachycarpa, MacCalliana, lutea, mackenzieana, Farrae, Barclayi, mri padophylla, commutata, pyrifolia, Richardsonii, Barrattiana, alaxensis, candida, Bebbiana, fallax, pedicellaris, éthabas ascensis, acilis, discolor, Scouleriana, planifolia, pulchra, subcoerulea, iano itchensis. RTHERN ALB gilis tita, arctica, glauca, brachycarpa, MacCalliana, lutea, ‘cseheaa Farrae, Barclayi, myrtillifolia — commutata, pyrifolia, cal- cicola, agente alaxensis, candida, Bebbiana, pedice llaris, atha- gracilis, discolor, Scouleriana, siecfela subcoerulea, arbus- culoides. NORTHERN SASKATCHEWAN: S. lucida, lasiandra, serissima, interior, or, dida, Bebbiana, pedicellaris, athabascensis, gracilis, discolor, Scouler- iana, planifolia, pellita, arbusculoides. Hinlgipsarsaidte MANITOBA: S. lucida, serissima, interior, reticulata, ves- herbacea, arctophila, glauca, brachycarpa, cordifolia, MacCal- oes lutea, Farrae, myrtillifolia, padophylla, cordata, oo cicola, candida, Bebbiana, pedicellaris, atha , gracilis, dis- color, Scouleriana, planifolia, pellita, ortusoulontes. Raison age ONTARIO: S. lucida, serissima, interior, reticulata, vestita, brachycarpa, coniifetia, ronan” be rigida, lutea, myr- ealcicola, ris, gracilis, discolor, F slasiolia, pellita, peariee 8 on organizing what knowledge we have of geographic di distribution einai the willows is in terms of actual species ranges. This could d be carried to somewhat greater detail than will be found in the following pages, but for present purposes some general categories should prove useful. sitio, _glaucophltoies, padophylia, la, cordata, pyrifolia, candida, B. na, pedicella RARE OR HIGHLY LOCALIZED SPECIES : gilis: an introduced species collected in our region thus far only sina yea ac in Alberta. S. vestita: common at Hudson Bay, i i in the M 18 HUGH M. RAUP but ‘with a single western locality in the vicinity of Banff. S. ampli- folia: known only from Yakutat Bay in southern Alaska. S. hebe- carpe: own in our region only at Teller, Alaska. S. phylicifolia: collected thus far only in two localities, in southwestern Yukon, an in the lower Kuskokwim valley of Alas SPECIES WITH DISTINCT THOUGH LIMITED RANGES IN OUR REGION In ie or arctic tundra: S. Setchelliana: central spa Kernel a 4 or to parts of these coasts. S. Chamissonis: western and et Alaska, northern Yukon, and the western border of Mackenzie. In the forested country: S. rigida: northern Ontario. S. glaucophyl- loides: northern Ontario. S. cordata: southwestern shores of Hudson Bay. S. fallax: apparently with a narrow range from the Rocky rit ; : Rocky Mts. and penta: of Alberta, gat ede British Columbia and the southeastern corner of Yukon. S. sitchensis: coastal strip and neighboring wiuintatna in northern and content British Columbia and in southern Alaska. WIDESPREAD SPECIES OF THE WESTERN MOUNTAINS, SOME WITH EXTENSIONS INTO ALASKA AND THE MACKENZIE BASIN S. lasiandra; S. mackenzieana; S. Barclayi; S. commutata; Barrattiana. a SPECIES WITH THEIR PRINCIPAL RANGES IN THE WEST BUT WITH EXTENSIONS EASTWARD S. polaris subsp. pseudopolaris: to the western arctic islands. S. glauca: to southern Baffin Island and Greenland. S. niphoclada: to western Manitoba. S. pulehra: to Great Bear L. and ‘Coppermine S. arbusculoides: to Churchill. SPECIES WITH THEIR PRINCIPAL RANGES IN THE BAST BUT WITH EXTENSIONS WESTWARD In the aretic or alpine tundra: S. herbacea: to Great Bear L. S. vestita: with a small isolated population in the Rocky Mts. S. arcto- ne rnd ees Rieke. 5 _cordifo lia: to Baker L. in Keewatin, with an isolated population in southwestern Yukon. S. calcicola: ‘Tin aml ltd poplin th the Rocky Mts. ee THE WILLOWS OF BOREAL WESTERN AMERICA 19 Rocky Mts. S. discolor: to northeastern British Columbia. S. plani- folia: to northwestern Mackenzie and southwestern Yukon. S. pellita: to eastern Saskatchewan. WIDESPREAD IN THE NORTHERN CENTRAL PLAIN S. lutea. WIDESPREAD THROUGHOUT MOST OF THE TUNDRA OR FOREST Species primarily of the tundra: S. reticulata; S. arctica. Species of the forested country: S. interior; S. myrtillifolia; S. candida; S. Bebbiana. NOTES FOR AN ARRANGEMENT OF SPECIES IN TERMS OF FORM, SIZE, AND HABITAT PREFERENCE A combination of form, size, and habitat groupings among the species may be of some assistance to collectors or to students of vegetational geography. As in the case of species ranges, the arrangement given below is tentative and incomplete. It can be used only as a point of departure, because observations of the behavior of the species of Salix with relation to habitat preference and the flexibility in their growth habits are too scanty to justify good generali- zation. I have made no attempt to classify the tundra species as to habitat. This is due in part to my lack of field experience with some of them, in part to the extraordinary plasticity of many of them in their capacity to occupy more than one kind of site, and in part to the extreme local varia- bility and instability of the tundra habitat itself. The willows of this region are most abundant in species and individuals on the most “open” lands, i.e., on lands with few or no trees, and least abundant in forests. Fur- ther, they are more frequent on the wetter soils than on the drier ones. In these circumstances, willows are most abun- dant in the arctic and alpine tundra, where there are no trees and the soils are nearly always damp; and they are least common in dry upland woods. Within the forested regions, in terms of moisture and the absence of trees, conditions analogous to those of tundra are most nearly met with at the margins of muskegs and wet meadows, and at the ~ margins of floodplain forests. There are nearly as many species of willows in this marginal vegetation as there are in the tundra. Next in a descending order of abundance are the species found on the gravelly and sandy shores of lakes, and on gravel and sand bars along the larger streams in mountains and foothills. Then come the willows of - shrubby or partially forested muskegs; of moist, rich up- 20 HUGH M. RAUP land forests; of sand dunes; of the dense forests of river floodplains; and finally of the drier forests of upland pla- teaus and hills. e forms and sizes of willows may be arranged in two main divisions: species that are consistently prostrate, sometimes with creeping branches that root at the nodes, or with decumbent branches that ascend above the ground only 10-20 cm.; and those with upright or strongly ascending growth habit. The latter group can be divided roughly into those that are of “medium” height (usually 1.5 meters or less) and those that are taller. These form categories are not sharply defined, for there are some species that are ordinarily upright and often as much as several meters tall which are capable of living as depressed shrubs on exposed sites. However, there are not many of these, and their characteristics are fairly well known. By far the greater proportion of the prostrate willows grow in the arctic or alpine tundra. Only two or three of them are to be found within the forest, and these only a short distance below or south of the timberline. Most of the medium-sized species are to be found in swales or along streams in the tundra, or on relatively open land in the forested country. Most of the species in the forests or immediately marginal to them are the taller ones. SPECIES OF ARCTIC OR ALPINE TUNDRA Stems prostrate or trailing, or the branchlets ascending only 1-2 dm.: S. reticulata; S. vestita; S. Setchelliana; S. polaris subsp. pseu 7a. herbacea: S. rotundifolia oa Dodgeana ; io. qui lebopieylin: S. arctica; S. arctophila; S. ovalifolia; 8. stolonifera; S. flagellaris ; S. glacialis; S. rag tirsnsdg S. cordifolia; S. Chamissonis; S. calcicola; S. arbutifolia; S. hebeca Stems upright or eis of medium height, usually 2 m. tall or less: S, interior ; S. glauca; S. niphoclada; §. brachycarpa; S. Par Dj -§ ealcicola; S. Barrattiana; S. alaxensis; S. Beb bbiana; S. planifolia; + pu - arbusculoides. SPECIES COMMONLY FOUND AT THE FORESTED MARGINS OF MUSKEGS, _ WET MEADOWS, PRAIRIES, AND OF ACTIVE RIVER FLOODPLAINS Upright or ascending shrubs, or small trees, 1-10 m. tall: S. lucida; x sia ; S. serissima; S. fragilis; S. interior; S. MacCalliana; oe Ss. rigida; S. lutea; S. mackenzieana; S. clayi L- I Bar ; S. glaucophy — Ss. eae S. cordata; S. pyrifolia; rt Richardsonii; S. ttiana; S. alazensis; S. Bebbiana; S. discolor; S. subcoerulea; : S. gracilis; S. Senulerana: SS. piglet: S- S. pellita; S. - lanifotia; pe ? THE WILLOWS OF BOREAL WESTERN AMERICA 21 SPECIES FOUND ON GRAVELLY OR SANDY LAKE SHORES, OR ON GRAVEL AND SAND BARS ALONG THE LARGER STREAMS IN MOUNTAINS AND FOOTHILLS rr otages prostrate, with branchlets ascending to 1-2 dm.: S. Setchel- “Upright or ascending shrubs, 0.5-5 m. tall: S. lasiandra; S. interior; clad S. glauca; S. nipho Se: 3: brachycarpa; S. lutea; S. mackenzie Ss. By S. myrtillifolia; . poe. S. aggre? S. Rioh- . Barrattiana; S. alaxensis; S. candida; S. : a Savulertiene. S. planifolia; s. pulchra; S. subcoerulea; S. arbusceu loides SPECIES OF SHRUBBY OR PARTIALLY FORESTED MUSKEGS Stems prostrate or trailing, or the branchlets ascending only 1-2 ifolia. Upright or ascending shrubs, usually 2 m. tall or less: S. serisstma; S. glauca; S. niphoclada; S. brachycarpa; S. Farrae; S. Barclayi; oe peg arovnltgg S. commatnts ; Ss. ao ape te S. Richardsonii; S. candid SPECIES FOUND IN MOIST, RICH, UPLAND FORESTS Stems prostrate or trailing, or the branchlets ascending 1-2 dm.: S. reti puree S. myrtillifolia. Upri ascending shrubs, or small trees, 1-10 m. tall: S. glauca; S. myreifota; S. Bo cm yee S. Richardsonii; S. alaxensis; S. Beb- biana; S. : S. arbusculoides. SPECIES FOUND AMONG ACTIVE SAND DUNES Upright or ascending shrubs, usually 0.5-3 (-8) m. ta : S. glauca; S. brachycarpa; S. Tyrrellii; S. Turnorii; S. amplifolia; 's silecsbobe: S. Bebbiana; wares Scouleriana. gee? FOUND IN RIVER FLOODPLAIN FORESTS Upright or ng shrubs, or small trees, 1.5-10 m. tall: S. glauca; S. ae rs. Bebbiana; S. Scouleriana; S. w rbusculoides. SPECIES OF DRY UPLAND FORESTS Badeig or ascending shrubs, or small trees, 1.5-10 m, tall: S. Beb- arbusculoides; S. Scouleriana. THE COLLECTIONS OF SPECIMENS The willows of all this region are badly i in need of further collection and study. Many of the species are known only from fragmentary an and infrequent collections, so that their true taxonomic Oe ia be Be eT ie a whether they make 22 HUGH M. RAUP Sterile material should be avoided, however, whenever possible; but if it must be resorted to, care should be taken that it is typical of the plants from which it comes. Young, rank-growing root suckers and sprouts, partitularly, should date of collection, and on the form, coloring, and size of the bush or tree from which they come. Good specimens will consist of typical leafy twigs and branchlets with their catkins attached. any species of Salix show a great deal of seasonal vari- ation in the size, Shape, color, texture and pubescence of their flowering habits; that is, the flowers appear at about e same i flower buds are the first to break in the spring, and the lows : S. Richardsonii, Barrattiana, candida, discolor, phyl- _teifolia, pulchra, planifolia, subcoerulea, pellita. In addi- are intermediate between precocious and coetaneous: THE WILLOWS OF BOREAL WESTERN AMERICA 23 leaves. This bloom (twigs that have it are said to be “pru- inose,” while on leaves it is most commonly called “glauces- cence” and the leaves are said to be “glaucous”) is caused by a waxy substance that is easily driven off by heat. Con- sequently, if artificial heat is used for drying specimens, notes on the amount and distribution of “glaucescence” on them should be made while the material is fresh. Willow catkins are particularly acceptable to certain small insects which commonly are collected with the specimens and go on eating the catkins while the latter are drying. A light sprinkling of ordinary naptha flake will drive off the pests and at the same time prevent mould. THE USE OF THIS PAPER The student who already has some knowledge of willows, and is able to “place” a specimen near its probable relatives, will turn at once to that part of the description of species that includes these relatives. He will narrow his decisions as to the identity of the specimen by comparing descrip- tions and by using the group keys if they occur; or he may turn back to the general keys to find definitive characters. He will use the notes on geographic distribution and habitat preference as a partial check on his decisions. The student who has no general knowledge of species or relationships among the willows must gain this knowledge or substitute something for it in order to reach the level of decision with which the above student began. There is no real substitute for the experience upon which such knowl- edge rests — experience in the observation of finely detailed differences among the plants, or in the learning of species “by acquaintance” in much the same way we learn to know people and are able to call them by name. _Nonetheless, it is possible to narrow, more or less mechanically, the fields within which decisions toward identification have to be made. The keys, and the arrangements of species based upon geographic distribution, form, and site preference, may be used as contrivances for narrowing the fields of decision. The use of the keys will be greatly facilitated if the stu- dent learns to make a few simple observations on a given plant at the outset. The more effective of the main keys is that involving female flowering or fruiting material. Here each of the species or species groups is placed in one or an- other of three series. Some appear in more than one series 24 HUGH M. RAUP to take care of variation within species or groups. Series I (5 species) is composed of trees or large shrubs, with cat- kins whose bracts drop off soon after the flowers expand. In all the willows of Series II and III the bracts remain on to the maturity of the fruit. Most of the species in Series I are also rather easily distinguished by vegetative charac- ters. Series II (19 species) is composed of low, mostly prostrate shrubs, the branches in some species trailing on the ground and sometimes rooting at the nodes. All of these species occur in the arctic or alpine tundra, where most of them have their main ranges. Series III (40 species) in- cludes only upright or ascending shrubs, though some are of low stature. Major subdivisions in both Series II and III are based upon whether the ovaries and their pedicels are glabrous or pubescent, whether the leaves are regularly serrate or entire, whether the leaves are green on both sides or glaucous beneath, whether the ovaries and cap- sules are essentially sessile or pedicelled, and if the latter, whether the pedicels are long or short. With a little practice the collector will make these simple observations quickly, and as quickly place his plant in its proper series and get it into a small number of species or species groups. Within this small number he must then make a few more detailed observations for further identi- fication. The use of information given in the preceding pages on geographic distribution, form, and habitat preference prob- ably can be explained to best advantage by some examples. Let us suppose that a student is working above timberline in the Brooks Range of northern Alaska. He may be making as complete a collection of the flora as possible, or he may be making detailed descriptions of vegetative cover in con- nection with studies of wildlife or of geomorphic processes. In any case, we will assume that he has no prior knowledge of willows except perhaps that they are “shrubby” plants with woody stems, that they have their flowers and fruits in seta and that they have “willow-like’” leaves. chances are excellent that any willow he picks up wit “si one of the 37 species listed under “Alaska” on p. 16. Because he is sean in the tundra above timberline he will find by comparison of the Alaska list with that for the pence that only 27 of the species are common to : both lists, and he has narrowed the field by 10. If he now . ‘compares his list of ye Alaska tundra a Species with the lists THE WILLOWS OF BOREAL WESTERN AMERICA 25 of those whose ranges are highly localized or of limited ex- tent (pp. 17-18), he will find that one (S. hebecarpa) is extremely rare and known only from Teller, on the Seward Peninsula. Four others (S. ovalifolia, stolonifera, flagel- laris, glacialis) apparently are restricted to coastal Alaska. Although he should not eliminate these five species entirely, the chances of his finding them in the Brooks Range are remote. He has reduced his field to 22 species and has not yet had to look at the willow he collected. At this point he must make an initial decision based upon the structural characteristics of the plant in question. One of the least equivocal is whether or not the stems are trail- ing or prostrate on the ground. Alternatively they could be upright or strongly ascending from their roots. We will assume that they are of the prostrate or trailing kind. Turning again to the list of tundra species on p. 20, he finds that this decision has narrowed the field from 22 to 11 spe- cies. Eight of these are strictly prostrate in habit, while the other three (S. niphoclada, arbutifolia, and myrtillifo- lia) may be upright or spreading shrubs and have the ap- pearance of prostrate ones. The 11 species are as follows: reticulata, Setchelliana, polaris subsp. pseudopolaris, rotun- difolia, phlebophylla, arctica, arctophila, niphoclada, myrtil- lifolia, Chamissonis, and arbutifolia. All of these species are in Series II of the key to pistillate flowering or fruiting material (pp. 28-30), or they may be separated rather easily with only sterile specimens. The second example will be drawn from the forested country. We will assume that a student is working along the banks of the Peace River in the great bend between Car- cajou and Vermilion, where the river valley lies between the Buffalo Head Hills and the Caribou Mountain Plateau. This area is all in northwestern Alberta, and the list on western Alberta, and thus far have never been collected so far east as the lower Peace River valley: Barclayi, caleicola, commutata, Barrattiana, alaxensis, subcoerulea. The sev- enth, S. fragilis, is an introduced species for which I can find no record in our region except along the Athabaska River east of Lesser Slave Lake. Two more species, vestita and 26 HUGH M. RAUP arctica, can be eliminated on this basis, and also because they are known only in the alpine or arctic tundra, and no tundra exists on the banks of the lower Peace River. Most of the habitats on the banks of the lower Peace are fiood- plains and semi-open grasslands, both with forested mar- gins. Most of the willows along the river will be found among those listed for such marginal habitats on p. 20. However, small areas of shrubby muskeg are to be expected, and three species more or less restricted to this kind of site (see p. 17) are listed for northern Alberta: Farrae, pedi- cellaris, and athabascensis. It is possible that these will turn up on the floodplains, but as yet none of them has been found in the region of the lower Peace except on the neigh- boring uplands. Thus we have reduced the number of species probably to be found on the banks of the river between Carcajou and Vermilion to 18; or if we include the muskeg species, to 21. Three th ra, serissima, and interior) are in Series I of the key (p. 28). Seven of the remaining 18 are in that group of Series III species that have glabrous ova- ries and capsules (pp. 30-31) (Farrae, lutea, macken- zieana, myrtillifolia, padophylla, pyrifolia, pedicellaris) . The other eleven, also in Series III, have pubescent ovaries (pp. 31-32) (glauca, brachycarpa, MacCalliana, can- dida, Bebbiana, athabascensis, gracilis, discolor, Scouleri- ana, planifolia, arbusculoides) . Further examples could be postulated for other regions and other kinds of sites, but the above will be sufficient to suggest the method. The site-wise subdivisions that I have made for the willows of the forested country should be used guardedly, and with allowance for a somewhat greater spread in habitat tolerance than I may have given to some of the species. This is particularly true for species that are _ situated at or near the margins of their geographic ranges, where they sometimes occupy unusual sites. Further, the concept of the “forest margin” will be conditioned by the definition the student is using for “forest.” The term “forest” is used here to mean a vegetation of trees whose crowns form a closed or nearly closed canopy. Open, park- like forest, such as occurs at timberline or in partially tim- bered 1 muskegs, will show a great variety in the density of Por pending upon the extent of closure by "he poem of “in : sediade” coecivens 6 mute to wtine the ic tifice on of willows. ee eee ent THE WILLOWS OF BOREAL WESTERN AMERICA 27 to be patient with these refractory plants, bringing them as close as he can to recognized species, and then holding them until he has gained a wider acquaintance with the general nature of species in Salix. When he has this experi- ence he will find that many of the supposed intermediates will come within the range of variation that seems to be allowable in one or another of the major species. There will, of course, be a residue that cannot be taken care of in this way. Some of the residue might be natural F: hybrids, for which there may be evidence in imperfectly or incomplete- ly developed catkins and leaves, or they may be the result of free interbreeding among closely related species or vari- eties, with recombinations of the characters by which these entities usually are distinguished. Again, it is entirely pos- sible that they are undescribed species in their own right; for in a region as vast as this, in which so little willow collecting has been done, new species are almost certain to found. ACKNOWLEDGMENTS me i ch of his Flora of Alaska a Yu N based. Dr. ee re. _ kK. W. Baldwin of the National Herbarium _ critical drawn from his OT cs peels indebted to several aomemeg i that have a my own field work in the ‘can Phi hical Society, the American : : scoak Ged 3 Society of the Sigma Xi, and t ’ au the Highway were made during t agible without the aid of the Northwest wervice — 28 HUGH M. RAUP Command of the Ss. Army. Later studies, in 1948, in southwestern Yukon were supported in part by the Wenner-Gren peor aic eid Anthropological png the Peabody Foundati for reha i i most helpful with the loan of field equipment. My studies in Eur herbaria were financed by a further grant a the Milton far ye Harvard. KEY TO THE SPECIES AND GROUPS, BASED UPON WELL-DEVELOPED FEMALE FLOWERING OR FRUITING SPECIMENS A. Bracts of the flowers not persistent in fruit, straw-colored; catkins us or serotinous, borne on leafy peduncles; usually tall shrubs of alluvial soils, or occasionally of muskegs ........ Series I black, rarely straw-colored; catkins appearing before, with, B. Low, prostrate shrubs, some species with rooting branches; cat- kins appearing with the leaves (coetaneous) or after the seg (serotinous) Series II. B. Erect shrubs or small trees, with branches never prostrate or gui: ing (though many are of low and spreading form) ; catkins usual- ly appearing before or with the leaves Series III. SERIES I ES bee provided, toward the upper end or at the base of the blade, distinct, often irregularly lobed, glands; leaves lanceolate, dis- tinetly and Agel closely glandular-serrate; catkins borne singly on leafy S. lucida group. i Petioles and hanes of the blades without prominent glands, or the linear, entire or variously toothed, but not finely and closely gland- aa ; catkins borne singly or 2-3 eT °§. fragilis. 2. Leaves nearly sessile or with very short, non-viscid petioles, some- eg pekriy | entire but usually with rather distant, often spiny S. interior. SERIES II 1. Pistillate fiver with 2 glands, on the inner and outer sides of the base of the pedicels; catkins Pg to be terminal on the branch- lets; leaves strongly reticulate-veined ................ S. reticulata group. _ Pistillate flowers with only 1 peti on the inner side of the base of the pedicel; catkins usually on leafy lateral branchlets. 2. Ovaries and pedicels glabrous, even when beg oo 8 Leaves more or less distinctly serrate on the margins. 4. Leaves regularl; oer te Prveed the whole of the een on both sides, not aaa cal _ 5. Tiny aretie plants with sane eaibbes vane branches, the aerial twigs a. with Agta 2-4 rounded, thin, shiny leaves — em. long ....... ~ Be herbacea. THE WILLOWS OF BOREAL WESTERN AMERICA 29 5. Larger plants with copiously leafy twigs, the leaves ~— to ovate or oblong-lanceolate; the more prostrate form the usually erect ........0... S. myrtillifolia — S. pias pales 4, Leaves irregularly serrate or entire, or with minute, more or ess regular teeth around the lower half of the blades; green or glaucous beneath. 6. Leaves somewhat fleshy or leathery in texture, obovate or oblanceolate and tapering —. to a petiole-like base, rounded, acute, or even retuse at the apex, entire or irregular- ly serrate; twigs commonly pruinose; poienien shorter than the glands S. Setchelliana. 6. Leaves not fleshy; obovate or elliptic, rounded or obtuse at the apex, more or less re rly and finely serrate around the lower _ twigs not pruinose; pedicels usually longer than the PIBNES ooo cccccsscensi css S. arbutifolia (See S. arbutifolia group). 3. Margins = the inuwes entire, or at most with only occasional small teeth. 7. Leaves somewhat fleshy or leathery in texture, obovate or ob- lanceolate and tapering gradually to a petiole-like base, acute. rounded or retuse at the apex; twigs commonly pruinose ............ S. Setchelliana. 7. Leaves not fleshy or leathery; twigs not pruinose. 8. Densely matted plants of arctic and alpine tundra, formed by frequent branching of the stems; leaves 2 cm. long or less . S. iehaudsfelie group. 8. Arctic-alpine plants with elongated prostrate branches; leaves 1-4 cm. long S. ovalifolia group. 2. Ovaries and pedicels more or less densely pubescent (sometimes only the lower parts of the ovaries). 9, Leaves finely though distinctly serrate all around the margin, obovate, up to 4-5 em. long and half aS Wide OF MOTE .........:ccccceeereee 9. Leaves entire or nearly so, or serrate only around the newer half. 10. Bracts of the flowe rs 2-3 times longer than wide, uniformly straw-colored, yellowish, or pale brown, or sometimes darkened a little toward the apex, rather short-hairy, with the hairs much shorter the bracts _. S. cordifolia or forms of S. niphoclada (See S. glauca group). 10. Bracts of the flowers broader in proportion to their length, the ‘width commonly more than half the length, variously colored but usually with long hairs or merely cilia ate at the gant ted shru ¥ sp: 12. Small, aaa 1 cg (no I than 1.5 em.), with an abundance of marcescent, skeletonized leaves from preceding seasons clothing the stems and form- ing a conspicuous part of the mats .. S. phlebophylla (See S. rotundifolia group). 12 Marcescent leaves not abundant, and not forming a con- of the mats. 13. Styles none or very short, at most 0.1-0.3 mm. long; small prostrate plants with slender branches; Scares euake or poe ee ee a ee 30 HUGH M. RAUP 13. Styles 0.5-1 mm. long or a little more; branches matted, much divided; leaves 1-2 cm. long, ovate or obovate g : sateen er eeeeenee ? . long 14. Leaves up to 8 em. long (commonly 2-6 cm.), pe or ob- scurely and irregularly toothed; catkins upright, sometimes as much as 10 em. long S. arctica group. SERIES III 1. Ovaries and pedicels — even when young. 2. Pedicels distinct, 2-6 times as long as the ies ds. 3. Leaves with entire Seasns or guts only around the lower half, smooth, green above and glaucous beneath S. arbutifolia group. 3. Margins of the leaves distinctly serr ate. 4. Leaves green on both surfaces, the lower surface sometimes of a slightly lighter shade. 5. aap acute to acuminate at the apex; slender, erect shrubs to 3 m. high, with pale grayish-green bark ........ S. Turnorii. 5 Leaves rounded or obtuse at the apex; lower shrubs, 2 dm. to 2 m. high, with dark-colored branches ........ S. Barclayi group. 4. Leaves green above and glaucous beneath. 6. Leaves giving off a balsamic fragrance (even Jong after they are dried), broadly pi elliptic, or elliptie-oblong; pedicels of the capsules 2-4 mm. long pyrifolia. 6. Leaves not giving off a balsamic fragrance. 7. Leaves usually broadly lanceolate, ovate or cbevate. the width commonly 1/3 to 3 ii 5 the length .... S. glaucophylloides ee: 7. Leaves usually lanceolate, oblan ceolate, oblong-lanceolate, 0 narrowly pra AP the width commonly 1/6 to 1/3 the eneth or occasionally a little broader . Tig group. 2. Pedicels none or very short, at most and even in fruit not more than twice the length of the glands. oe aes of preceding years persistent on the twigs, conspicuous, _ jinear-to era yer oaepatos or reniform, glandular-serrate on the margins or entire; catkins sessile on the twigs of the pre- ceding year, eats on leafy peduncles ............S. Richardsonii group. 8. Stipules not persistent after ie growing season; catkins borne m leafy peduncles. 9. ia of the leaves distinctly and er serrate, at least on the lower part and eti all < 16. Pedicels usually less than 1 mm. long; styles 1-2 mm. long. a Leaves glabrous beneath and becoming glabrous above as y mature .0... . Barelayi (See S. Barelayi group). 11. eanee densely grayish-hairy and somewhat silky on both S. cordata. el: 218 mm. lone: — 05-1 ims one 8. commutata. oceasional teeth at ir- — THE WILLOWS OF BOREAL WESTERN AMERICA 31 12. Styles 2 mm. long or less. 13. Pedicels about 0.5 mm. long; leaves usually serrate, but etimes entire ........ S. Barelayi (See S. Barclayi group). 13. Pedicels 1-1.8 mm. lon. 14. Leaves glaucous benea th, all entire or oe with minute teeth, the blades completely glabrous .... S. Farrae. 14, Leaves green beneath, serrate or entire, schsaeent on both sides at least on the veins S. commutata. 1. Ovaries pubescent (rarely only on the base 15. Leaves green on both sides, though sometimes of a lighter shade beneath, not glaucous, glabrous except when very young, 3-7 cm. long and 1/4 to 2/5 as wide, glandular-serrate on the margins S. MacCalliana group. 15. Leaves glaucous with a bluish bloom on the lower surfaces. 16. Pedicels distinct, 2-6 times as long as the glands. 17. Pedicels short, 0.5-2.4 mm. long; catkins sessile and precocious, or on very short leafy peduncles and appearing — the early iscolor — 17. Pedicels conspicuously longer, 2.5-5 mm. uns caikins born on well-developed leafy peduncles, and appearing with the leaves. 18. preaeser prrantay nar acuminate, 4-5 times as long as wide, en- tire or toothed, pubescence if present giving a some- what si eid appearance to the under - .. _ 1-2 mm. lon acilis. 18. Leaves ovate, oblong, obovate, or ovate-lanceolate, Sa width 3 to % the length, entire or dentate, pubescence if present not silky in appearance; bracts commonly 2.5-3.5 mm. long S. Bebbiana. 16. Pedicels — or very short, even in fruit scarcely more than twice the length of the glands. 19. Flowering twigs more or less distinctly pruinose with a bluish bloom. 20. Lower surfaces of the leaves with a permanent cov ering . whitish, felt-like, — ‘pu ADEREEREE ; styles 2 mm. long more; syed usacsil alaxensis an gf ane SRE (See S. alucousis group). 20. Pubescence on the lower surfaces of the leaves, if _Dresent, shining oa not felt-like or — styles usually less than mm. long; catkins precoc 21. Lower surfaces of the lanes s with a silvery, shining pubesc- S. pellita group. - Leaves ‘labrous “beneath except on the midribs and main «the phakics} obs group. 19. Siete twigs not prul 22. Bracts of the flowers more or less oe carga yellow- ish-straw-colored or yellowish-brown ; pubescent h_ short hairs: that are much shorter than the bracts ila bosks o . glauca group. 22. nieces on a flowers : oF two colors, crag beter to black toward the apex, or if nearly all of one color, grayish-brown | ae ee 32 HUGH M. RAUP 23. Lower surfaces of the leaves, even in age, densely white- or light gray-tomentose or silky. 24. Leaves linear-lanceolate or narrowly oblong, commonly 5-7 times longer than wide, with entire or undulate e margins; young twigs, | ower leaf-surfaces, and capsules covered ae. giving the whole shrub a hoary appear- S. candida. 24. ks broader in proportion to their length, usually only 3-4 times as long 25. Pubescence of the lower leat-surfaces sea them a sil- very, shining, or satin-like appearan 26. Catkins precocious; bracts of the eas about 1-1.5 mm. long; twigs usually pruinose but sometimes with the bluish bloom lost S. pellita group. 26. Catkins coetaneous ; bracts of the flowers 1.5-2 mm. long; twigs never pruinose S. sitchensis. 25. Pubescence of the lower surfaces of the leaves doll white or dull grayish, not shining. 27. Leaves thickly felted beneath or on both sides Bhi a white tomentum, the mid-veins on the lower aces where visible through the tomentum, yellowish ; “stipules rather distantly glandular on the margins S. xensis group. 27. Leaves whitish or light grayish hairy-pubescent on both sides, somewhat silky in appearance; midribs on the surfaces of th ves not yellowish; stipules densely on the margins S. Barrattiana. 23. Lower surfaces of the leaves glabrous or somewhat pubesc- ent, but not with a dense whitish or grayish pubescence or tomentum. 28. Leaves distinctly, said: and often conspicuously glan- dular-serrate on the 29. Styles 0.2-0.8 mm. “i bracts about 0.5-1 mm. long S. reer 29. Styles 0.5-2 mm. long; bracts about 2 mm. long. . phylicifolia (See S. phylicifolia group). 28. Leaves pera or the margins with occasional smal]! glan- dular teeth a ' | t irregular intervals, or with serrations only around the lower half. borne on leafy peduncles; bracts ioles 5-13 mm. long (mostly 8-10 WORD icc Oe 31. Leaves obovate to a 4.5 em. long or jess, usually ' in proportion to length, commonly less than twice as long as wide; petioles a oni mm. ne fmonty i. mm.) ols S. arbutifolia bracts oc he . _ and the styles 08-2 mm. Wiehe sae THE WILLOWS OF BOREAL WESTERN AMERICA 33 KEY TO SPECIES AND GROUPS, BASED UPON STERILE LEAFY PLANTS EXCLUS OF RANK-GROWING SPROUTS 1. Leaves regularly crenate-serrate or aries . around the margins. 2. Petioles provided, at the distal end o the base of the leaf blade, with distinct, often irregularly teak glands; leaves dis- tinctly and rather closely glandular-serrate ........ S. lucida a group. 2. Petioles and bases of the blades without glands. . Low, prostrate shrubs with ee trailing; or depressed shrub rubs with twigs ascending 4. Leaves rather thick and bs or \nathecs. bluish-green on both sid h s or green, or glaucous only beneath; twigs commonly i S. Setchelliana. 4. Leaves areca thin, not fleshy or leathery, not bluish-green above; twigs never pruinose. 5. taken usually “Glathod beneath with long, white, asd ap- pressed hairs, though sometimes glabrate in age — on the veins; low _—- of cae tita. 5. Leaves glabrou 6. Leaves eouaea or broadly obovate, at most not more than twice as long as wide and usually much less. i & — rounded at the base, scarcely if at all longer than rb a epee wedge-shaped at the base, commonly somewhat longer an hamissonis. 6. Leaves oblong, oblong-ovate, or ohlone-inacepiate, usu ally 2-3 times as long as wide; low forms of ........ S. myrtillifolia. 3. Upright shrubs, a few dm. to a few m. hi 8. Leaves linear or narrowly lanceolate, 8-10 times longer than wide, nearly sessile or with very short petioles, sometimes near- ly _— on the margins, but usually with rather distant spiny tee 8. es usually broader in proportion to their length, rarely linear, and ectiy serrate or crenate-serrate. — 9. Leaves permanently silky or hairy on both si 10. Leaves entire or finely and rather inconspicuous serrate, in habit of growth; alpine shrubs beeen mie on gi gravels at lower altitudes) of the northwestern ; S. Bivatsioen: 10. Leaves conspicuously glandular dentate-serrate, green and densely grayish-hairy and somewhat silky on both sides, more spreading in habit; eastern subarctic and north tem- . cordata. shrubs: 9. Leaves not permanently silky or hairy on both sides. 11. Leaves — acuminate at the i usually with long lanceolate or ca ips. 12. Branchlets brittle, easily broken off at the base; introduced with leaves rather coarsely un undulate-serra ‘ate . Soh eS Wisachiets: not brittle; native, often tall. , spreading shrubs. with closely glandular-serrate se Jones Soe are! group. 2 es E: te, but not conspicu- ee secoreeanccnared at the apex. 34 HUGH M. RAUP 13. Dried stipules persistent on the twigs for 1-5 years ...........+ S. Ri ne group. 13. Stipules not persistent. 14. sia elliptic to er aeets or oblanceolate, usually 3-6 mes longer than Se Leaves green on both piilen: rather firm and leathery in texture. 16. Tall, upright and somewhat pyramidal shrubs with pale grayish-green bark except on the young twigs which are red; leaves pale green ...........00+ S. Turnorii. 16. More spreading shrubs with dark reddish or brown rk; leaves bright green ............ S. MacCalliana group. ath. scence on the under surfaces of the leaves long, appressed, and slightly tangled ............ S. phylicifolia. 17. Pubescence on the under surfaces of the aiacees minute, straight and appressed, giving a slightly satiny appear- 18. Pubescence of mixed white and tawny or rust-colored hairs; petioles 6-10 mm. long or more ........ S. gracilis. 18. Pubeseence almost entirely of white hairs; petioles ging somewhat shorter, 4-8 mm. S. arbusculoides. 14. Leaves anal oblong, ovate. or obovate, Bagi iaga broad- in proportion to their length than in the above five poe or if serene. fag glaucous foment 19. Shrubs usually of mossy muskegs or mossy tundra. 20. Leaves green on both sides ................ S. Barclayi group. 20. Leaves glaucous beneath. . Leaves giving off a balsamic f ong ragrance. lo: after drying, broadly ovate, elliptic, or Be is S. pyrifolia. Tes Leaves without a — fragrance. 22. Leaf margins conspicuously se ef. S. Barclayi group. 22. Leaf margins said and eS serrate eeveeree athabascensis. 19. Shrubs usually found on alluvial soils or in upland woods. 23. Leaves glabrous, even when yo’ y. glaucophylloides group. 23. Young leaves gusta glabrescent in age or the pubescence ort decir . Leave both sides, not glaucous beneath. 26. Dead, dry ee persistent through one or many years, some- x er as Se RNS vi Ui een a lee a alee THE WILLOWS OF BOREAL WESTERN AMERICA 35 27. Leaves larger, up to 5 cm. long, the veins not prominently ove or raised beneath. 28. Leaves usually not over 2.5 cm. long and commonly smaller, the biades thin)... S. polaris subsp. pseudopolaris. 28. tae rather thick and fleshy, commonly more than 2.5 cm. lon S. Setchelliana. 25; Feats glaucous beneath. 29. Leaves strongly reticulate-veined, which shows conspicuously eath, rounded, obovate, or oval in outline; stems trailing. 30. Leaves prominently Se as e on the upper surface, 1.5 to 4 em. long, borne on reddish petioles. S. reticulata. 30. say not prominently in all above, usually 1 cm. fom or ss, borne on relatively short petioles ...........-...+ alis. 29. paaehe not conspicuously reticulate-veined beneath ; is various shapes St: Twigs pruinose, and the leaves glaucous on both surfaces; leaves rather thick and fleshy or leathery, entire or ~~ and irregularly serrate; a depressed shrub A lep ascending branches Setchelliana. 31. Twigs not 32. Leaves facie eniulareeeene only around the lower part of the blade; trailing shrubs with elliptic to obovate leaves, em. long or less, the width usually more than half the S. arbutifolia. on. peck entire around the whole margin or, at most, obscure- irregularly serrate. 33. Adult leaves averaging less than twice as long as broad. 34. Prostrate shrubs with trailing stems . ovalifolia group. 34. Prostrate shrubs ‘with ascending branches which are spreading and often gn arled. 35. — rounded to heart-shaped at the Ls aus soon ene h at least on the veins; low form heat 2 cordifolia. — Adult scihior averaging more than twice as ek as broad. . Shrubs with trailing branches, often forming presage 36. Prostrate or ascending shrube, with gnacied, cpio Mt branches Si, Petinles 2 mm. long or less; prostrate forms OE eas s. niphoclada. 37. Petioles longer, more th ; prostrate forms of .. ... S. glauca or S. ardifebe ‘(See S. glauca group). 24. Upright shrubs with acral athy not or ascending from 38. peasy green nee a both sides, sometimes paler beneath, but not —— linear or narrowly jaxiceciate, 8-10 times longer than wide, nearly sessile or with very short petioles .......- S. interior. 89. Leaves usually broader in Baga to their length, not linear, borne on well-develo petioles. 40. ‘Leaves commonly rounded or somewhat coo eto at the tae mostly shrubs * a or swampy tundra. 36 HUGH M. RAUP 41. Leaves thinly tomentose above when young, glabrous be- neath, commonly becoming blackened in drying .........-+-++ Barclayi. 41. Leaves densely grayish-tomentose on both sides when young, not blackening in . commutata. 40. Leaves more frequently obtuse or wedge-shaped ‘at the eras shrubs of dry woods, prairie margins, or floodplain forests . S. Bebbia: 38. Leaves glaucous beneath, or the lower surfaces obscured by thick tomentum. 42. Lower surfaces of the leaves covered thickly with permanent pubescence or tomentum. 43. Pubescence on the under surfaces of the leaves shining, sil- ike. very, satin-l 44. Twigs concn species” bi the Rocky Mts. and of north temperate eastern Ameri S. pellita group. 44. Twigs not sooo a maniac slope species .... S. sitchensis 43. Pubescence on the under surfaces of the caves opaque, nek shining. 45. Leaves linear-lanceolate or narrowly oblong, commonly 5-7 times as long as wide, the upper surfaces impressed-reticu- late and grayish pubescent at least when young; the whole plant having a hoary appearance S. candida. 45. Leaves broader in proportion to length, usually not more i — times —— than wide, not conspicuously impressed- cula 46. Stipules ILE or with a few long hairs . seseee . Barrattiana. 46. Stipules densely tomentose or hairy-tomentose .............:++ S. coe group. _ 42. Lower surfaces of the leaves glabrous, hairy, or pubescent, but not densely covered with a thick, tL ageket tomentum or pubescence. 47. gp polar sessile or on petioles usually less than 2 mm. long; shru edie Mstag y less 1 m. tall, upright or depressed, with -green leaves. 48. poten oblong-ovate to rounded, usually much less than twice as long as wide S. calcicola. 48. Leaves longer in = to their breadth, usually twice as long as wide or mo 49. gy rses elliptic to narrowly obovate, 2-4.5 em. Jone eecsieinen - 49. Leaves ovate or obovate to obl: late, averaging smaller than in the last (15-3 cm. lene .. S. brachycarpa. is Leaves borne on well-developed petioles more . than 2 mm. long. «50. Stipules evident, and — through 2 or more years on old wood. eee —— > though sometimes gia brate in Phan, al S. glenea var. stenolepis. or ns older than ¢ one > year, = por ee ae NST a et eh Oe OE ee ite Lee ee Ee aoeey Sm Pree eee THE WILLOWS OF BOREAL WESTERN AMERICA 37 52. Twigs older than one year glabrous or thinly pubescent, commonly pruinose; stipules linear, narrow to the base without prominent glands on the teeth ............ = 50. Stipules not persistent beyond the growing seaso 53. Leaves linear-lanceolate or narrowly oblong, ‘ocasake 5-7 times longer than wide, the upper surfaces impressed- reticulate and grayish-pubescent at least when young; the whole plant having a hoary appearance S. candida. 53. Leaves not as above, shorter in proportion to breadth. 54. Twigs yellowish, or yellowish with reddish streaks or blotches; tall shrubs with pale, yellowish-green, acumina ate leaves that are usually serrate but sometimes ent Pe eeeeeeneee eseeeeee . lutea. 54, Twigs gray, brown, or reddish, not yellow. 55. Leaves s oblong-ovate to rounded, usually much less than twice ong as wide, up to 5 cm. long ........ S. calcicola. 55. Leaves oe broadly oblong-ovate to rounded, on longer in proportion to breadt th than the above, © relatively nape obovate to broadly elliptic and pct ing shorter (1-3 56. Pubescence on ex under surfaces of the leaves of aera straight, appressed hairs, giving a slightly pearance. io take eae of mixed white and tawny or Sp ygone hairs; petioles 6-10 mm, long or more ........ 57. Pubescence almost entirely of white hairs; Leer averaging somewhat shorter, 4-8 mm. epcacareesieceaccdseneoesseenneseseery ere) satiny. Low, erect or spreading shrubs, usually less than 1 m. tall; leaves 1-5 cm. long. 59. Erect or spreading sre sometimes nearly 1 m. tall; leaves 1-5 em. long. 60. Leaves rather stiff, leathery, entire, dark bluish-green above, obovate, & glabrous, 2-4.5 em. long usually 2.5-4 S. pedicellari. ieike as long as broad, the margins co minutely — around the lower half ... oie sewvastonebns oo arbutifolia. 59. Low, spreading ing shrubs, commonly a over 3 or 4 dm. ; leaves usually 3 > mg or less. blin: S. icella var. hypoglauca, except Rotem a ; smaller leaves (3 long at most); known in our region in the Straits area .-- S. hebecarpa. 61. Leaves broadly ovate, oblong, ‘or elliptic 1. 5-3 cm. ¢ tuse ~green, t or glabrate -...---------reeecssesee s. oriole: : er shrubs or | usually an 1 m. 58. val, or if smaller the leaves differing from HUGH M. RAUP 62. Tall shrubs or small trees, commonly 3-8 m. high. ve aa. veins beneath; common in lowland and moun- tain slope woods and thickets throughout the region — S. Bebbiana. 63. Leaves larger at maturity, commonly up to 8 cm. long, distinctly obovate. 64, satis commonly about twice as long as wide; a s known only from the Yakutat ai she T of ka plifolia. 64. Leaves usually about 3-4 times as lone as nas sir S. discolor group. 62. Medium ona: usually 1-3 m. tall. eaf-shape spicing elliptic-oblong, or elliptic, _ e., broadest near or below the middle ye some lanceolate leaves will be found. in these plants, erou 66. Twigs reddish or reddish-brown, emails glabrous and shining. 7. Leaves dark green and shining above oe Leaves entirely glabrous, the margins entire or with occasional, irregularly-sp small teeth; on in damp thickets at the margins of wet meadows and muskegs, and extending northwest- ward to the upper Yukon valley where it merges with the closely related S. pulchr s. planifotia. 68. Leaves finely pubescent when young, but becom is) : d along the lower K . S. phylicifoha. 67. Leaves yellowish-green and rather dull above; a species of the northern acon Mts. and Alaska .... Farra 66. Twigs gray, glabrous or pubescent; leaves dull grayish-green above, usually with rather prominent bbiana. or Shianenclate. i. e eedeat abawe the sais ge ovate lanceolate, or elliptic forms will be found on these plants, but they will be in the minority). _ 69. Leaves rather dull grayish-or bluish-green 3-8 cm long, about 2.5-4 times longer than wide, commonly . _ Dubescent — < on oth sides usually _ grayish-puh maaeierings an abundant ead highly vari- 2 ae species in in muskegs, damp meadows and open ae — below Sbentions 8 — in oto age ve THE WILLOWS OF BOREAL WESTERN AMERICA 39 flowers late, and the dead aments often persist on the branchlets into the following season ..........+..++++++ S . gla 69. Leaves bright green and glabrous above when ma 70. Leaves 2.5-5 em. long (av. about 3 cm.), usually about half as wide as long, thinly appressed- hairy beneath, though finally becoming glabrous when old, shrubs usually less than 1.5 m. tall; rare or occasional in muskegs and wet ‘aeasowe Lebsee athabas 70. Leaves 2-7 em. long (av. about 4 cm.), usually 173-1/2 as wide as ie slightly silky when young but soon becoming glabrous; somewhat taller than the last, up to 2 m. (cf. also S. Barelayi which sometimes has nearly entire leaves) .......- S. fallax. DESCRIPTION OF THE SPECIES SALIx L. WILLOW Shrubs or small trees, ranging in size from tiny alpine plants only 2 or 3 em. high to trees up to 10 m., and oce occupying almost the entire spread of percEee habitats. Buds with a single scale; bia borne in catkins or aments, each of which is a simple spike of flowers on a stigmas each of which may be shes ovary sessi e or aa on a splits down two sides from the apex, freeing large numbers of small each of which is supplied with a mass of silky _ that is sanity blown by the wind; a= ng atten of | except that the ovary is reple 12 eae ee upon the species. KEY TO THE SALIX LUCIDA GROUP A. Leaves green on both surfaces, not glaucous beneath though sometimes paler green B. Young branchlets — or soon glabrate C. Leaf blades more less _ abraptly caudate-acuminate, - C. Leaf bla =. cote - gradually pierre sual 45 soe r than wide; a species es of 1 western merica. ......- 40 HUGH M. RAUP D. — mostly acute at the apex, scarcely acuminate, the width % much less than 1/3 of the length. ................08 S, serissima. js beater poner! Lg ere at the apex, width more commonly 1/4 the r les E. vent testis pee or soon glabrate S. ee (typ.). E. Young branchlets usually densely pubesce S. lasi aude var. lancifolia. Salix lucida Muhl. in Neue Schr. Ges. Naturf. Fr. Berlin 4:239 (1803) rge shrubs or small trees, in our region up to 3 m. high with slender trunks and brown, shiny, glabrous twigs; leaves lanceolate to ovate-lanceolate, up to 1.5 dm. long and 5 em. wide, usually long- caudate, serrate at the apex, wedge-shaped to rounded or even cordate at the base, finely glandular-toothed, glabrous, green on both sides but somewhat paler green beneath, shiny above, on petioles up to 15 mm. long; summit of the petiole at base of blade with large glands; fy pedunc up to 5 cm. long; capsules mm, long, coor Saas on pedicels up to 1 mm. long; bracts yellowish, "thinly hairy, falling off soon after the expanding of the catkins; styles 0.5 mm. long; eaistiuibe catkins up to 5 em. long: stamens 3-5 or more; filaments pubescent below An eastern species of temperate America (Map 1) which has heen collected at Moose Factory, York Factory, and in the valleys of the F naeieute lanceolate and seoee aus to eiiiie te tips, while in the typical species they are more abruptly narrowed to the long, cau- date-acuminate tips. Salix lucida appears to be the principal eastern element in a series of ov overlapping species that make up what I have called the “S. lucida group,” in which the more western elements are S. serissima and S. ira. «Salix lasiandra Benth. Pl. Hartweg. 335 (1857) 7 shrubs, or stnall trees up to 6 or 8 m. high, with reddish, shiny twigs; vented 6-12 cm. long, 1.5-3 em. wide, mostly lanceolate t thou gh : ommon. or and mud bars eine the tonic sama in the central an southern paris of the Macken ackenzie basin, THE WILLOWS OF BOREAL WESTERN AMERICA 41 and common locally in the Alaska Highway region from Lesser Slave L. to the Tanana valley. It oceurs sporadically in the lower Yukon Mats and in i Endicott Mountains of Alaska, and down the Mac- kenzie at least to Simpson and possibly as far down as Norman (Raup, 1947: a. “ors ( 1951: 138) found it along the Canol Rd. as far up as mi. 127, along the Lower Lapie R. Th ternmost collections I have seen are from mes = Pt., on os north shore of L. Athabaska (Raup, can 228-9). yi. Salix lasiandra is highly Hee oe species, masher ed in the shape and coloring of th leaves, and in the pubescence of the branch- lets. Some students have regarded these varatioas as separate spe- our s p ae hav den region. The following key, modified from a treatment by Ball (1926: 146), will distinguish them. Still other en are likely to appear as more collections accumulate. A. Leaves pale or somewhat glaucous benea’ B. Young branchlets glabrous or soon labvate S. lasiandra (typ.). B. Young sbesgeseiie: usually densely pubescent ...... var. lancifolia. A. Leaves green eath C. Young Peaehicg: sowie: or soon gilabrate ........ var. caudata. C. Young branchlets pubescen var. recomponens. Specimens determined as typi a S. lasiandra were collected by (Raup, lc.), and the latter was collected in the tdsgorst of Summit L. by Taylor et al, 4 (V); by our field parties along south of Watson L., 10967, 10968, 10969; and by Setchell at Fairbanks, 433 (GH). This variety pe also been found along the Yukon at Dawson by Eastwood 112, 113, 128, 304, 466 (K). Variety caudata (Nutt.) Sudw. in Bull. Torr. Bot. Cl. 20: 43 (1893) (S. caudata (Nutt.) Heller) was collected by our party along the Alaska Highway at the Sikanni R., 10402. Variety recomponens Raup in Sargentia 6: 149 (1947) is known thus far only from its type caulsetion: on a sand bar along the Mackenzie R. near Simpson, Raup 9073, 9076 (A). alix lasiandra is closely related to S. serisstma and S. lucida. It is the westernmost phase in a continuous population of lucida-like willows that extends across most of the nent forested part of the continent. Salix serissima (Bailey) Fern. in Rhod. 6: 6 (1904) S. lucida Muhl. var. serissima L, L. H. Bailey ex Arthur in Bull. Geol. Nat. Hist. Surv. Minn. ws 19 Leow a ‘ae up to 4 m. high, wi th glabrous, shiny, pelweh: -brown 1-3.5 em. wide, elliptic to SS he base, finely late, short-acuminate at the apex, acute to rounded at t and ar 42 HUGH M. RAUP leafy peduncles; capsules glabrous, up te 10 mm. long, on pedicels 1-2 mm. long; bracts up to 2.5 mm. long, yellowish, iach hd ; styles less than 0.5 mm. long; staminate catkins 1-2 cm. long; stamens usually 5, the filaments thinly hairy Salix serissima has been collected in the vicinity of Lesser Slave L., Brinkman 4648 (BH) and northward to the upper Mackenzie valley where it appears to reach its northwestern limit (Hay River, Lindsey 115, CAN; Mackenzie R. above Providence, Porsild 16554, CAN; lower Keele R., Lindsey 341, 343, CAN). It is 2 related to the western shorter catkins, longer pedicels, less eee Dee leaves, and by its later flowering time (mid-to late summer). I have seen specimens from the north pd ag of L. Athabaska, Rae 6353 a, and from the Cypress Hills, pia 5104 (K). Its northern limit in the Hudson Bay region seems to be west of James Bay, and I have seen specimens from the eo seg coast of that Bay, and from the upper Severn R. (1943: 92- 3). See note under S. lucida. Salix fragilis L. Sp. Pl. 1017 (1753) . long; catkins ap pearing with the leaves, borne on leafy peduncles, ih istillate up to 7 em. long; capsules about 5 mm. long, glabrous us, on pedicels 1 mm. long or less; bracts oblong, ciliate on the margins, falling off soon after the catkins expand; styles 0.3-0.7 mm. long; staminate esoncey 3-5 em. long; stamens usually 2; filaments hairy at the ike _ Salix fragilis is an introduced species, brought from Europe an widely spread from cultivation in temperate eastern America. I hae seen a single collection of it from our region, in the Mackenzie drain- _ age basin (Ball 116493, (CAN), on the bank of tl the Athabaska R., east of Lesser Slave L., at Athab thabaska Landing). _ Salix interior Rowlee in Bull. Torr. Bot. Cl. 27: 253 (1900) Salix longifolia Muhl., non = — shrubs, sometimes as uch 1 as 3-4 m. sigan _— — sharp, rather her widely spaced teeth, a or 2 bide t petioles, some of them 2m nearly opposite on the twigs; with the ‘ | leafy peduncles, the piatitlate upto 8 usually THE WILLOWS OF BOREAL WESTERN AMERICA 43 Common on sandbars along the = igh rivers (and occasional on Good Ho lake shores) northward to York F Te ag the vicinity of aring our region repre r. pedicellata (And ay tall in Can. Field- cas 40: 175 pes which has very ane leaves faite not over 6 mm. wide). The exceptions came from along the Liard R. south of Watson L., 10965, and from the delta of the Slave R., Raup, 561 sats Both of these have the broader leaves of the typical species i heeleri nm reported by Scoggan qd 1957: pe es the Nelson R. about 30 mi. from York Factory. It is distinguished by the more or less permanent silvery- silky pubescence on its leaves. KEY TO THE SALIX RETICULATA GROUP A. Prostrate shrubs, with leaves glabrous or nearly so, borne on elongate petioles (up to 3 cm.) ; lateral veins of the leaves few, the lower ones rising from near the base ........------- S. reticula A. Shrubs with prostrate or erect branchlets; leaves usually con- spicuously s silky-hairy on both sides (or occasionally glabrate), the lateral veins well scattered along the midrib; petioles not much longer than the buds ......-.--. aS tita. Salix reticulata L. Sp. Pl. 1018 (1753) Prostrate shrubs, trailing along the ground and — only a few centimeters above the surface; twigs smooth, purpl ish-tinged impressed to form @ reticulate pattern, conspicuously glaucous sometimes silky when young but soon glabrous, borne on long, reddish petioles, entire on the margins or nearly so; catkins slender, reddish, up to 3 em. long, appearing terminal on the branch- lets; capsules 3-4 mm. long, sessile, densely P ; bracts about 2 mm. long, wine greenish with dark tips, hairy; ” styles nearly in damp alpine tundra, @ , t region fag rate in Alaska to the region of Gener throughou re eo and in the ! also found ©: nalts Bays gene 1957: 15, map 118). Easily me other prostrate alpine willows by = io ish ly smaller _ Map 28. Salix — is closely related to S. vestita, and ove! te - northern part of -ealled the the “S. See a one MeKinle Park in 1932. Later coll 44 HUGH M. RAUP Salix vestita Pursh, Fl. Amer. Sept. 2: 610 (1814) Depressed or sii ne shrubs, usually less than 6 dm. high but sometimes as much as 1 m., with stout, spreading branches and pubes- cent or glabrous twigs; leaves rather thick, rounded to obovate, ellip- tie or oblong, 1-8 cm. long (usually 2-4), closely crenate on the mar- 1.5 long, blackish toward the tips; styles short or obsolete; stamens 1 or rarely 2; filaments glabrous A predominantly eastern arctic species (Map 3), of somewhat more southern range than S. reticulata, that has been collected as far south as northern Manitoba and has an apparently isolated population in the Rocky Mts. I have seen specimens from Churchill and Fort Severn, and Scoggan (1957: 232) reports collections from York Factory and from the Hayes R. at various points u up to 100 mi. 8S. W. Hudson Bay. The material a Churchill and from the Rocky Mts. represents var. erecta Anders. ex DC. Prod. 16°: 300 (1868), which may be distinguished from the eben by the following characters. A. Capsules narrowly ovoid, definitely tapering to the blunt tip; pistillate catkins 0.5-3 em. long; staminate catkins 0.6-2 cm long; leaves, obovate or orbicular, usually somewhat retuse pa reticulately veined S. vestita (typ.). A. Capsules only slightly tapering to the broad rounded summit; istillate catkins 2-5 em. long; staminate catkins 1-3 cm. long: ack elliptic or oblong, often subacute, usually plane above .. var. erecta. _— vestita is closely related to S. reticulata, and the two may be dered as segregates within an S. reticulata group. Salix Setchelliana Ball, Univ. Calif. Publ. Bot. 17: 410-12 (1934) Prostrate. shrubs; branchlets sometimes ascending to 15 or 20 em. above the ground, pruinose, pubescent when very young, but hat ; sig or bluish-green and shiny above, green or glaucous beneath, glabrous when full-grown; catkins appearing With the leaves oo on shank leafy peduncles, up to 2 em. long; capsules glabrous, up to ‘7 mm. long, borne on very short pedicels; bracts up to 4 mm. long, oo yale , ciliate on Margins, sometimes notched at the apex; ee ow 02-03 foes — stamens 2: Glaments hai ii originally described a eigkerial collected in é 1732) ee helt col- R. bridge, an wad ae THE WILLOWS OF BOREAL WESTERN AMERICA 45 between Kluane L. and the Donjek R. Argus (personal communica- tion) reports a eee in the University of Alaska Herbarium col- lected at Pa Vy erson; and Spetzman cites a specimen collected by Gudin at the Itkillik Lakes in the Brooks Range. The species is locally abundant in the Kluane L. area, and is conspicuous on gently sloping, wave-washed shores near Burwash. p 4. Salix polaris Wahl. Fl. Lapp, 261 (1812) Represented in our flora entirely by subsp. pseudopolaris (Flod.) Hultén in Fl. Alaska & Yuk. 510 (1942) (S. pseudopolaris Flod. in hors ysis _ Bot. 20A, No. 6, p. 8, 1926; S. polaris var. selwyn- 1 n Contr. Arn. Arb. 6: 44. 1934), to which the following ees at Prostrate, small, trailing shrubs, often forming ense s bro e g with leafy peduncles, short (usually less ees 3 ecm.) and thick, few-flow- ered; capsules pubescent, 5-7 mm. long, nearly sessile; bracts up to 2 mm. long, obovate, mostly dark brown to black, glabrous or thinly hairy; styles 0.5-1 mm. long ; stamens 2; filaments glabrous. Comm abundant in the alpine and arctic tundra of western and ages - Alaska, Yukon, and to the eastern slopes of the Macken- zie Mts. the Arctic Coast east of the Mackenzie delta (Hultén, is common alpine species in the vicinity of the Alaska Highway. Thus al it has = been collected within the continental borders east of ackenzie Mts. except on the coast just east of the Mackenzie delta (Porsild, 1957: 175, map 119). ie 6. Salix herbacea L. Sp. Pl. 2: 1918 (1753) glabrous and shiny, : peti tioles, usually ely 2-4 leaves on each twig projecting above moss in which it grows; catkins appearing with “the leaves or later, 5-8 : "8 : glabrous, 5-6 mm _ long, n early sessile; bracts very short, obovate, nearly glabrous on the outer surface, but with long white hairs on the inner side; styles 0.5 mm. long or less; stamens 2; filaments glabrous. bese ks ee nals faa 2 ae Great Bea ea te (1957: 232) reports collections of it at Nejan- jlini, Baralzon, and Nueltin Lakes in northern Manitoba. Map 6. KEY TO THE SALIX ROTUNDIFOLIA GROUP = a a” Small, matted shrubs with glabrous, ovate or elliptic ves 1-2 oe em. long, snd with an abundance of er skeleton wonined 46 HUGH M. RAUP dead leaves from several preceding seasons clothing the stems S. and forming a large part of the mats .........-.... phlebophylla. A. Marcescent leaves not abundant though a few may persist through one year, not forming conspicuous parts of mats; leaves very small, rounded or ovate, 4-10 mm. long ppd 4-7 mm.); tiny matted plants of alpine ‘tandra B. Lateral veins of the tice more or less raised on — sides, even the veinlets sige nent S. rotundifolia. B. Lateral veins 0: Skee slightly incised oily ieee beneath, the a6 not visible S. Dodgeana, Salix rotundifolia Trautv. in Nouv. Mém. Acad. Mose. 2: 304 (1832) A tiny species that forms mats of thin branchlets in rather dry alpine tundra, the whole plant often not more than 2-3 cm. high. Acaxeniy Cid in its " eneeien nee: to Alaska (Hultén, 1942: 508-9) and neighboring Yukon, where it is an arctic and alpine species. It has been collected in only one place in the vicinity of the Alaska Highway, on rock ledges and scree slopes above op apg on near the southeastern end of Kluane L. at m 12167, 12168, 12294. Map 5. Salix rotundifolia is one of three closely related species that I have re whose ranges overlap, and among which intermediates are commonly found. Salix Dodgeana Rydb. in Bull. N. Y. Bot. Gard. 1: 277 (1899) Tiny mat-forming shrubs, searcely over 2 em. high, wi ort, = underground s , the aerial parts densely leafy, glabro obsolete; staminate pores usually 3-4 flowered; gp i Salix Dodgeana was ein ueiekunty described from setetial coll _ Electric Peak in Yellowstone Park, and and has since Co had c seine Places in the Rocky Mts. of Wyoming and south- his origin: escription - also cited a THE WILLOWS OF BOREAL WESTERN AMERICA 47 (1942: 511) agrees with Schneider’s diagnosis. Porsild (1945: 18) reported three collections of S. sige be hg along the Canol Rd. east of the Yukon-Mackenzie divide. Later dy of this material con- vineed him that it represented Ss. pokes aa ana, an se with which I concur, for I have seen the specimens and they a good match for the type of S. Dodgeana. Otherwise the i a is pl in our region. It is closely related to S. retundifolia and S. phlebophylla. Map 5. Salix phlebophylla Anders. Gfvers. Vet. Akad. Férh. (Stockh.) 15: 131 (1858) very small willow of the arctic and alpine tundra, forming dense mats of branching stems; leaves glabrous, ovate or elliptic, 1-2 cm. long, rather prominently few-veined, entire, green on both sides; cat- kins commonly 1.5-3 cm. long; capsules glabrous or slightly pubescent when young, on very short pedicels; bracts up to 1.5 mm. long, black with white hairs; styles distinct, but short; stamens 2; filaments glabrous; dry, p artially skeletonized remains of dead leaves marces- cent, remaining on the twigs for several years and clothing them dense Widespread in the tundra of the Arctic Coast and interior moun- tains of Alaska (Hultén, 1942: 511-12). Porsild could find no record of it in Yukon except on the Arctic Coast (1951: 140), and it has not been found in the immediate vicinity of the Alaska Highway though it is to be expec orsild’s record for it on the eastern alpine slopes of the Mackenzie Mts. (1945: 18) he now believes should be col 5. Salix phlebophylla resembles S. rotundifolia, but is easily distin- guished from that species by the masses marcescent leaves that clothe its stems. I regard it as one of a closely related series including rotundifolia and S. Dodgeana. KEY TO THE SALIX ARCTICA GROUP ume? d extending out from the midribs at sometimes nearly dicularly; glands short and beond. as long as thick, b: truncate at the apex, y ¥ 7 half the length of the pedicels of the capsules ........ S. a ves usuall e green above, opaque or slightly lustrous, the ja, re * ou t from midribs at much ane angles; 2-4 age i fa TOE rter than the glands or rarely a ® little longer .. ... S. arctica Salix arctica Pall. Fl. Ross. 1, 2: 86 (1788) An extremely — and widespread arctic and northern alpine ailing ranches often forming dense mats; leaves i ially comet 0 and glaucous beneath, obovate, oval, oblang, or elliptic in outline, borne on well-defined petioles, acute, obtuse to rounded or retuse at the apex, som: sometimes villous when young, but e at naturi oS — 48 HUGH M. RAUP catkins up to 9 cm. long or more, much longer than thick; capsules 8-10 mm. long, pubescent, or in some forms becoming glabrate in age, sessile or on very short pedicels; bracts about 2 mm. long, dark brown or black, long-hairy; styles up to 1.5 mm. long; staminate catkins up o 5 em. long; stamens 2; filaments glabrous. In this treatment I am making of S. arctica a broad and inclusive species. It is so variable, and has been so commonly collected and m recen i Schneider G : (1918); Hultén, 1942: arene: Porsild, 1951: 140-141; 1957 70; Raup, 1943: 98-100; 1947: 150-51; Polunin, 1940: 157-9; Ball, 1950. baa 1s Eastern arctic forms of S. arctica have been variously treated, by some students as a se te species y others as a group of vari Single species rt my fidence i of the hears of the Hudson. Bay region and the L S. arctica population in the eastern mmonest of these rm forms is the plant I then called S arhca: var. Br - ders. in DC. P 16°: 286 (1868) (S. angl ). It is distin- with great difficulty, as being generally less robust than the typical species, and in having leaves that are, on the oo some- what smaller. Similar forms are to be found in the western range of e species, however, and I do not now consider hee bo to have any tax- onomic oie Fis © pica ta tec eset ton . Ball has ob- 41) and p 1at S. hudsonensis is iden THE WILLOWS OF BOREAL WESTERN AMERICA 49 Some of the specimens cited by Schneider under S. hudsonensis, however, are not in agreement with his description or the type mate- rial. They have broadly elliptic-ovate to somewhat rounded leaves that are mostly blunt or rounded at the apex and wedge-shaped to subcor- ate at the base. The elude collections — these characters: J. M. Ma udson Bay, lat. 55°-56°, acoun 18822 (CAN); Aoeerig Hudson Bay, J. M. Macoun 79148 (CAN) ; ei of the Alban: any | anglorum var. kophophylla Schn. in Bot. Gaz. 66: _— 1918). Hultén has maintained, sooner pic ae. in his treatment of the arctica-group in Alaska and Yukon: S. areti ca Pall., S. crassi- well-defined geographic segregation: S. arctica on the Arctic Coast and in the Bering Straits region; S. er assijulis on the southern coastal mountains of Alaska, the Aleutians, and in the Bering Straits region; . torulosa on the interior mountains of Alaska and Yukon, but also appearing at Bering Straits. The key characters by which he distin- guishes them are as follows: (1942: 502): A. Seales light brown, oe or pointed, styles about 1-1.5 mm. long, poses? = bro A. Seales round, oo beesae black from the tip, thus Gften bi- colored or eatinely rhlace: rounded or orbicular, styles long B. Leaves orbicular to obovate, often on long petioles, reatkion — and thick on leafy peduncles, styles long and stout (¢ ~~ 2 mm.), capsules short-pedicelled .........--.-++-+-+ erasst B. a smaller, on shorter peduncles, styles shorter, caps capsules essile It prs be noted that all of these So . ioe ley pase gi difference within narrow ranges of color, shape ze, When they are applied to a large number of specimens pevevaphic segrega- tion proposed by Hultén immediately breaks rate Porsild evidently had this experience when he was unable to see S. torulosa in any of his many collections of S. arctica along the Canol Rd., though he cites a number of specimens along the more southern parts of it that ap- proach S. crassijulis. I have been unable to segregate from our Alaska Highway collections any wl idespread torulosa-form, and I have seen plants of the crassijulis-form in the mountains of the in- ther study may bring out a more realistic definition of es, but for the present I prefer to consider the existing ones only as in- termingled varietal phases of a single species. As such I pro a the torulo. . ) r e é : v7 . biCA ; . Be hn.) Soc. Nat. Mosc. 2: 309, 1832) ; S. arctica var. araioclada (Se Raup in Sargentia 4: 100 (1943) (S. anglorum var. araioclada Schn. in Bot. Gaz. 66: 133, 1918; Jour. Arn. a S: 67, 1 1921; S- crassijulis Trautv. in Mém. Soc. Nat. Mosc. 2: 308, 1832, at least as to American plants). te the iene Saher relatively large, bi I am unable ogee od ot i the Wank: ‘The affinities of S. paisa among the American forms of S. arctica tén says that it differs “from S. erasst, areemiahe 10 50 HUGH M. RAUP the long and narrow eri aaeeee light green leaves, glabrous on both sides except in the margin and sometimes along the median nerv e be- low” (1942: 520). In leaf-form this suggests var. antiplasta, cise it is of interest to note that Ball (1.c.) has given the latter a wide range which has broadly obovate leaves, obtuse to rounded at the apex and wedge-shaped to nearly round at the base. These suggest var. kopho- phylla. Within the region covered by the present Hartigan therefore, five phases of S. arctica Pallas may be recognized: S. oe (typ.) ; var. antiplasta; var. kophophylla; var. sie cards and var. torulosa. It should be emphasized that these forms are poorly Aafoed, and "that the following key will do little more than. indicate trends. A. Leaves obovate, obovate-oblong, ovate, ovate-rounded, or broadly elliptic, rounded to subacute at the apex, taperin rounded or nearly cordate at the base, the width usually one-half to four-fifths se ron be scales of the flowers black or with black tips, usually B. Wil decals ae usually 3-6 em. long and 2-4 em. wide, y obovate thin-papery, u in gene outline, obtuse to rounded at the apex, and tapering to the base; pistillate cat- kins up to 9 cm. long at maturity .. S. ar (ty B. Well-developed leaves more broadly proportioned, commonly roadly obtuse to rounded or subcordate at the base, thin- papery or thick and firm in texture; fruiting catkins usually not over 5.5 cm. long Cc. Sayers leaves hairy above though glabrous, thick and firm at maturity, 1.5-3.5 em. long, 1.2-3 em. wide, some- what plieate at the apn twigs purplish; fruiting catkins y 3.5 em. long or les kophophylla. C. Young leaves glabrous sae thin-papery when mature, up to g and 3 em. wide; twigs yellowish; fruiting cat- kins up to 5.5 em. long var. araioclada. A. Leaves shee pore Brom maprate or -obovate, narrowly oval, or elliptic, the width commonly less than half the length; scales of se Ga tee ar. D. Leaves firm in texture, thick, usually acute at both ends, com- _ Saliz arctien (sens. lat.) is common in the arctic and alpine tundra _ from Hudson Bay to ee ee eee ioe _ Porsild, 1957: 176, wre ore: It extends southward in the ee to the Puget Sound region, and in the Rocky Mts. to the RT ee i Pe ee ee 0.5 mm. long or less) ; bracts about 1.5 mm. long, reddish catkins w THE WILLOWS OF BOREAL WESTERN AMERICA 5 have included only two species: the multiform S. arctica Pall., and the closely related, primarily eastern arctic S. arctophila, Map se Salix arctophila Cock. ex Heller, Cat. N. Amer. Pl. (ed. 3) 89 (1910) Low, trailing shrubs with glabrous twigs; leaves elliptic to obovate, 2.5-4 em. long, 1.5-3 em. wide, rounded to acutish at the apex, rounded to deg nal a at the base, entire on the margins or nearly so, glab- rous, dark green and shiny above, glaucous beneath, the lateral veins usually eames: parallel, rather prominent, and leaving the midrib at a some agitate wider mnale than —— of S. arctica; sgpagt F gard ed in eastern American arctic-alpine situations (Raup, 1943: 100-101), gone" extending westward in the tundra to the Arctic Coast west of the Mackenzie delta (Porsild, 1943: 27; 1957: 176, map 123). Specimens — collected by Cantlon & Gillis i in the Jago L. region of northeastern Alaska (57-1456, 57-1078, 57-946) extend the species a little farther westward along the coast. Map 7. Hultén’s record of it in Alaska is based entirely upon a sterile specimen col- at KEY TO THE SALIX OVALIFOLIA GROUP A. Leaves bimeges ovate to rounded, the width commonly 4/5 the length or mor B. Styles iced usually 0.5 mm. long or less; gre not aaa stoloniferous S. ovalifolia. B. Styles mm. long; branches often. with eauer subterranean stolons S. stolonifera. A. Leaves obovate or elliptic, commonly not more than %2 as wide as long a tepstar: Salix ovalifolia Trautv. in Mém. Soe. Nat. Mose. 2: 306 (1832) Stems prostrate, trailing; leaves broadly meats to rounded, 1-2.5 cm. long, four-fifths as wide as lon. ong or more, entire at the margins, ad at eve apex and base or tapering rather abruptly at the base kins borne on leafy peduncles, the pistillate up to 2 or 3 em. long; cap- ‘sules glabrous and often glaucous, 5-7 mm. long; styles short (usually brown, hairy; > : c scabs about 0.5 mm. ail staminate p to 15 cm. long: 52 HUGH M. RAUP 138-9, 1918) under S. ovalifolia var. pubescens and var. subarctica. Both of these varieties have slightly pubescent capsules. Hultén (1.c. Salix stolonifera Cov. in Proc. Wash. Acad. Sci. 3: 333 (1901) A prostrate, trailing, alpine species that is very closely related to S. ovalifolia. Its leaves are slightly less broad in proportion to their length, but hardly enough to be outside the range of variation in the tter. The principal distinguishing character is in the length of the styles which in S. stolonifera are 1-2 mm. long, while in S. ovalifolia they are 0.5 mm. long or less. This character appears to be consistent in all the herbarium material I have seen, but whether it justifies the Separation of the two as species is questionable. Hultén (1942: 522) suggests that these two species and S. glacialis Anders. may be merged eventually as geographic races of a single entity. This prob- ably is true for S. stolonifera and S. ovalifolia, but S. glacialis ap- pears to me worthy of its own specific rank. Salix stolonifera as here defined seems to be limited to coastal Alaska, and to southwestern Yukon. I have seen the following speci- mens from our region: White Pass, Cowles 979 (GH); same, Malte, Aug. 24, 1916 (CAN); mountains south of Burwash, Clarke 217 (CAN); Haines Rd. near Yukon-Pacifie divide, Taylor et al. 820, 1179, 1194 (V); Jago L., Cantlon & Gillis 57-1305; about 40 mi. east of Cape Lisburne, Cantlon & Gillis 57-149, 57-209. Map 8. Salix flagellaris Hultén in Sv. Bot. Tidskr. 34: 376 (1940) _ Maintained as a specific entity with much hesitation. The type was aerate Walpole 16 Us) on the Kruzg tén ‘ fa 4 ; insula, and Hultén cites specimens from Muir Glacier, the Katmai to rds it as most nearly 1 era, THE WILLOWS OF BOREAL WESTERN AMERICA 53 species by its shorter styles, larger capsules, and by its larger and narrower leaves. In his key he separates it from S. ovalifolia and S. on the capsules are: Deering, Anderson 4806; Pt. Lay, Anderson 4417; same, Mason 6425; Mackenzie Delta, Porsild 7363. Other speci- mens showing some distinctly obovate or oblanceolate leaves are: Porsild 7363; Anderson 4806; Kivelina, Anderson 4626; Nome, Jones 9038, 9039. Map 8. In view of the preceding discussions, I propose that we have in the tundra of Alaska, Yukon and northwestern Mackenzie an S. i- glaucous. A sub-group with long styles (1-2 mm.) and broad to rounded leaves (width commonly 4/5 the length or more) has been called S. ovalifolia; and those with narrower, elliptic to obovate or oblanceolate leaves, called S. flagellaris (incl. S. arctolitoralis). Salix glacialis Anders. in Ofvers. Vet. Akad, Férh. (Stockh.) 15: 131 (1858) Known thus far only on the arctic and western coasts of Alaska, with one station on Akutan in the Aleutian Islands, and one on St. Lawrence Island (Hultén, 1942: 523-4). Map 9. Hultén characterizes it, from his studies of this material, as a decumbent, dwarf shrub i slender, reddish-or yellowish-brown branches, small, obo- vate, ciliate leaves, distinctly and often densely gray-hairy capsules, and short thick styles. The catkins are borne on leafy peduncles, and Staminate flowers have 2 stamens with aments; of the material cited by Hultén at Stockholm, and found the species a rather distinctive one ubescent capsules. Its leaves are strongly glaucous and promin- ently reticulate-veined beneath, suggesting S. reticula: re- e es are glabrous except for cill te Schneider (Bot. Gaz. 66: 139, 3 camdensis, based upon material collected by F. Johansen at Cam A. Petioles well developed, more than 2 mm. long B. Leaves usually much longer than broad, those of the peduncles about 2.5-4 times longer than wide; mostly upright (though often low) shrubs arene S. glauca. Seegraensecesseernee 54 HUGH M. RAUP B. Leaves usually short-oblong or short-ovate to rounded, those of he peduncles usually less than 2.5 times oo than wide; very low, spreading or somewhat matted shru - cordifolia. A. Petioles of the leaves short, none, or at most 1-2 mm. long C. ome short-oblong to spherical; leaves averaging 1.5-3 cm S. brae yearpa. Cc. Ses cylindrical, song more than twice as long as thick; leaves averaging 2-5 cm. long S. niphoclada. Salix glauca L. Sp. Pl. 1019 (1753) or ——- shrubs which, due to pubescence or glauces- cence or both, have a dull grayish-green appearance when seen in the aggregate. They are commonly a meter or so tall, but may reach 2 m. on suitable sites, or be depressed, almost trailing, shrubs in exposed places; twigs grayish-pubescent when young, glabrate in age; leaves borne on well-developed petioles, occasionally elliptic, but nearly aways broadest above the middle to make them obovate or broadly oblanceolate, entire at the margins or with a few scattered, obscure, glandular teeth toward the base, obtuse or acute at the base, obtuse (or almost rounded) to acute at the apex, dull green above, glaucous beneath, pubescent or hairy on both sides when young but commonly 0} age), commonly varying from as much as 10 cm. in length; cntkins appearing with the leaves or after them, on leafy peduncles, usually more than twice as long as thick, the pistillate commonly 5 > long or more; capsules densely grayish pubescent, styles eat ess than 1 mm. a apes syed or very short, even in fruit scarcely oleae twice the length of the glands; bracts oblong to obovate, rounded or occasionally sestralg me the tip, pangs yellowish or yllowish-brown; staminate catkins usually no e than 3. em. stamens 2; filaments hairy at the base or edkeaed glabrous. “‘Salie glauca is the principal Pipe in a huge complex of northern _ througho orthern of the bore Rocky Mts. This —- which I shall call the “S. hos group,’ _ variations, and cee which there are so man Geccadion ns, that anything more than an approximation of their true relationships is, knowledge. I-nigh im many that could be constructed. My only sefonse Sor it is aa it comes closest, for the ae being, to a rational interpretation of my own personal experience with these ‘etlions in the field and in her- : baria. The nomenclature of the group is voluminous and tangled. I have threaded my way through some parts of it, but even if I were : able to include dias all, the present work snail ont be — by "it. THE WILLOWS OF BOREAL WESTERN AMERICA 55 other species, S. MacCalliana and S. Tyrrellii, could be included in the group on the basis of their catkin characters, but their vegetative characters are so distinctive that I prefer to place them for the time being in a separate group. arctic treeline. Southward it reaches to the northern edges of the Great Plains, and into the central Rocky Mts. (Raup, 1947: map, PI. 21). It grows in muskegs, open woods, prairie margins, wet meadows, t a, Th the damp, mossy types where it is associated with dwarf birch, to ry and scree slopes. It is apt to loom large in any collec ion of willows from this region, for its abundance and extreme variability in form, size and habitat selection continually tempt the collector. They also tempt the student who is looking for describable novelties, particularly if he has only limited collections from a relatively small area. Four phases are here recognized within the mass of willows that answer to the above description of S. glauca. I am at present inclined to assign them a somewhat greater validity than the various western forms of S. arctica, because they show a little better geographic seg- regation, and in some cases a better segregation in terms of site. However, none of them is sharply defined, and intermediates among them are frequent. t. yin, variety of habitats, is var. acutifolia (Hook.) Schn. in Bot. Gaz. 66: 327 (1918). Most of the extensions of the species into the alpine and of this variety. \ Univ. Calif. Publ. Bot. 17: 416 (1934) (var. poliophylla (Schn.) Raup in Jour, Arn. Arb. 17: 233, (1936) has been found at Great and at Good an. : oth southwestern Yukon and central Alaska westward to the coast. The fourth phase is var. stenolepis (Floderus) Polunin in Nat. Mus. an. Bull. 92: 163-4 (1940). In 1947 I described an S. ea Vi perstipula (1947: 154-5, Fig. 13) which was distinguished by having linear-lanceolate, accuminate stipules that were persistent for as much nzie ce it has bee collected several times by Porsild along the Canol Rd. (1951: 142) and eastern slopes ackenzie Mts. In the vicinity of the Alaska Highway I have collected it only in south- ever, I have seen a specimen of it the n Highway (near Eureka Roadhouse in the mountains of southern Alaska) collected by nderson 8467 (GH) and it is to be ¢ ere in the interior of Alaska. Polunin’s var. stenolepis (9. lepis Floderus in Herb.) fale Tit ( : = VE I described var. ‘Baffin Island (Type: Malte 118812, CAN). When * des€ var. Bef olan’ cree seen this type material, but because Polonin’s — 56 HUGH M. RAUP description said nothing about the persistance of the stipules, I pre- sumed that they were not persistent, Sin nce then I have seen the Malte ac sistent stipules, all or most of var. stenolepis would be placed in S. glauca var. Serwctone and seems to parallel this variety in having a wide range of habitats which — far into the cd tundra. Whether it should be maintained as a separate variety or only as a form of var. acutifolia is sia to aaa belay but for the Leni I shall leave it with a varietal sta Present knowledge eae therefore, oes of the two most abund- ant and ee distributed fo orms, var. cutifolia has the greater ranges of the first two, are more limited geographically. The following key will serve to indicate the four phases noted above. As. iach persistent on the twigs, sometimes for as long as 5 years, appearing as narrowly lanceolate, dried appendages, often 1cm long or more, their margins glandular .................... var. stenolepis. A. oo not persistent on old wood, falling off within the first B taeies densely grayish-hairy or tomentose on both sides, even in age, averaging shorter and broader than in the other varieties ar. Aliceae. B. Leaves grayish-pubescent on both sides when young, but lack- ing the long-hairy appearance of var. Aliceae, and usually glabrate in age, at least on the upper sides C. Most of the leaves with an obtuse angle at the apex, or almost rounded . S. glauca (typ.). C. Most of the Ladiean with an acute angle at the “ee var. acutifolia. _ Hultén’s eitnent t of the various forms of S. glauca differs con- y from the above, and deserves some discussion here. I have omar published some notes on the “S, glance sv aiageco (1943: 102-4; 1947: 152-3), and will now only summarize Most of the confusion surrounding the gout of the forms of S. glauca in America seems to stem from varying interpretations of three questions and the observations concerning them. All of these interpretations have been conditioned historically by the gradually enlarging collections of the group, by the geographic spread of the collections, and by the increasingly critical faculties of the collectors. First is the question as to whether true S. glauea occurs in America at all. It was originally described by Linnaeus (1753) from specimens collected in —— oo earliest peg ee of any ine in the glauca-grou THE WILLOWS OF BOREAL WESTERN AMERICA 57 tions to the northern interior of the continent (1819-27), and in a few early Alaskan collections. From study of these collections came a confusion of names that are now difficult and in some cases impos- sible to typify (see Schneider, Bot. Gaz. 66: 318 et seq. for a discussion of this tangle). The great Swedish salicologist, Andersson, studying : : of ; ce that a comerer ae part of it represen ted a _variety Me he true S. Fl. Ror! Am. 2: 144 (1838). At about t the same tim an a (1899: 271) thought that if true S. glauca were present in America it was very rare, and that all or nearly all American ee of it represented one or another segregate. Coville, reporting on the willows of the Harriman Alaska Expedition (1901: 321), wus unable to distinguish much of the Alaskan material from the typical species, a vali ies in Alaskan NG pongirttcs ) Schn. My own studies of northwestern willows e led me agree with Coville, and both Hultén and Porsild are — pe Res “this trend. The second question centers in the concept of Salix desertorum Richardson. Of all the American willows that are closely allied to the type collection in the herbaria at Harvard and nea New York Botanical Garden, at the National Herbarium of Canada, in the Barratt Herbarium at Wesleyan University, Middletown, Conn., and at Kew. The specimens all appear to have come from very low plan many of them with a somewhat rsstes Sr — of growth so that the catkins (immature though well formed) are numerous in rather close pee s. Its name has been applied, over the years, to various gates in the S. glauca group, such as S. brachycarpa, and one or or another of the varieties of S. coraiieiia: or to some forms of S. glauca itself. I ‘ave seen oi fe two other collections that I could match with Richardson’s type material. One of these is my no. 534, from Taltheilei Narrows in the eastern arm peng Great a L. Here ae oe lants -on an exposed rocky headland in ughty soil, an hardly ‘more man exposed Joh (Raup, 1981: 241-4, Pl. 218). ‘They differences are found even on a single plant. The second i was made by Lindsey (71, CAN) on Et-then Island in Great Slave ei and Sees resembles Richardson’s type. From study 0 of these speci- mens I have concluded that S. desertorum is best interpreted as an aberrant form of S. Here — by local site factors, and as S sorts no Sepia taxonomi ee Sass 58 HUGH M. RAUP However, Hultén has chosen to maintain a concept of S. desertorum; and in doing so he revives for it the combination S. glauca subsp. desertorum (Richardson) Andersson in Ofvers. Vet. Akad. Forh. Stockh., 15: 129 (1858). It is difficult to know what forms he pro- i ertorum, w. The pl dently a low shrub” (1.c., p. 526). This brief description would cover a great deal of the material here called S glauc d var acutifolia, particularly specimens from low-grown shrubs xposed places. M the material that has been called var acutifolia, on low). His meaning is further confused by the following state- ents: “There seems to be no doubt that this Alaskan plant is identical with the eastern American called S. cordifolia var. calli- carpaea ...S. desertorum seems to me to be the arctic American glauca representative distributed from Alaska to Newfoundland...” Even if the dubious S. desertorum were allowed some taxonomic sig- nificance, it would be sufficiently difficult to crowd into it all the varied stern ro : i As with several other species of Salix, Hultén found himself with a residuum of specimens which he accounts for by hybridization. glauca x Farrae Walpolei.” 1 have seen none of these specimens and ean have no opinion on them. Hultén (1.c., p. 528) says that they “agree a Fi alpolei, except that they have pubescent capsules and leaves hai Ily on the lower side.” He cites a . a3 a € specimens most nea closest to S. glauca he says form “the bulk of specimens named by Schneider S. glauca acutifolia.” He cites approximately 80 specimens ig Lr “we both sexes in the same catkins ; I have one other record of this phe- iomenon in our region, in our no. 12217 from the shore of Kluane L. _ Another of the citations, Eastwood 620, is a sterile specimen of __S. Scouleriana. I had no difficulty in putting all except the last, and _ the two with imperfect catkins, in $. glauca or its var. acutifolia, _ about equal number ;in each. I am unable to see the entirely different _ S. padophylla (= S. pseudomo teola) as an influence in any of this THE WILLOWS OF BOREAL WESTERN AMERICA 59 The third confusing element in the interpretation of S. what I shall call its “apparently inherent variability.” The ¢ Set of this could become known only with the deedaal accumulation, over many years, of enough specimens and field studies to bring it out. For many parts of the S. glauca group there still is not nearly enough material with which to wor a Not only is there wide variation in form, but also in the capacity to occupy apogee sites. I do not think that the taxonomic realities in S. glauca or in the group of entities related to it will be understood until the full range of their variability is better known, and until we have some light on its causes. Salix niphoclada Rydb. in Bull. N. Y. Bot. Gard. 1: 272 (1899) Upright or spreading shrubs, sometimes decumbent and less 2 dm. high, or sometimes upright and 1-1.5 m. tall, with grayish- pubescent branchlets; leaves 2-4.5 ecm. long, sessile or with petioles only 1-2 mm age elliptic to narrowly obovate, entire, grayish-pube- a even in or sometimes glabrate above; catkins appearing ties on leafy peduncles; apeae as much as 6 or 7 mm. long, though commonly shorter, nearly sessile or on very short pedicels; styles short or nearly obsolete; bracts of one iets yellowish-strawcolored or yellowish-brown, pubescent with short hairs that are much shorter than the bracts themselves; staminate sre up to 2.2 cm. nepda sta- mens 2; filaments glabrous, or sometimes hairy toward the b Occasional in sa meadows, crevices, and on sandy lake ac e northwest coast of Hudson Bay westward to Alaska (Por- places along the Canol Rd., and Hultén (1942: 530-1) gives it a wide though somewhat scatte Pe: in northern, western, interior Alaska. It was originally described from material collected along the Mackenzie R. about 30 ae north of the Arctic rinioh: and has been collected southward in the Mackenzie basin as far as Great Slave L. ad 10. t will be noted from the keys and from the above description that S. ‘nkodiile differs from S. ycarpa (see below) chiefly by its to glabrescence in the leaves. It is like the latter species in having sessile or nearly sessile a: in this character the two species aga from others of the S. glauca group found in our region. Hultén (1.c., pp. 529-32) recognizes three other willows in Alaska and Yukon that should be be discussed here, for I believe them to be ‘scarcely if at all separable from S. niphoclada. The first has been called carpa var. Mexiae Ball in Univ. Calif. Publ. Bot. Li: 412-14 Ga), which differs from S. brachycarpa chiefly by its d pedune neles. poe growth, and larger leaves which tend to glabrescence in age. But these are f gens 2th the ag eri _ by which S. niphoclada is se aR from S. brachycarpa. ct -Ball’s rather po seinen at Mezxiae a yt as oak te taal for = or a have seen 2 Spite se) of var. _ Mexiae ae 60 HUGH M. RAUP separate it from S. niphoclada except by its glabrous leaves, the character emphasized by Hultén. i S. niphoclada toward glabrescence in the leaves, S. Muriei can be The third “species” is S. fullertonensis, described by Schneider (Bot. Gaz. 66: 340-3, 1918) from specimens collected by J. M. Macoun 79164 lected by Jones 9045 at Nome, on the Seward Peninsula. In Stock- holm I saw a fragment of the type of the species, and the Jones speci- men from Nome. The label of the latter states that the plant grew “prostrate on tundra.” Both of these specimens (photographs of the type were also at hand) show whip-like branches that must have been at least decumbent. Otherwise I could not have distinguished them from S. ni a. no more than 10-15 em. high, while in less exposed places in the im- mediate vicinity they may be 1 m. tall or more, with many transition phases between these two extremes. In my field notes on collections » “depressed shrubs”; no. 11899, from margin of prairie near mi. 1021, “low shrubs, 15-30 em. high, semi-trailing”; no, 12358, from gravelly-sandy shore of Kluane L., “shrub 60 cm. high”; no. 12403, dry bluff near shore of Kluane L., “shrubs 60-120 em. high.” Like- wise I have well-developed flowering or fruiting specimens of S. glauca from shrubs ranging on 15 to 90 cm i 0-60 cm. high, sometimes decumbent.” In a group with such flexi- bility as this, and in the absence of any experimental evidence that genetic segregation has occurred, it seems unjustifiable to give specific THE WILLOWS OF BOREAL WESTERN AMERICA 61 S. lingulata Anders. I have seen no authentic material of this poorly known plant, but judging by what descriptions and discussions I have at hand (see also Schneider, Bot. Gaz. 66: 353), I agree with Hultén that it is probably identical with S. niphoclada. Finally, it should be emphasized that S. niphoclada is closely related to S. glauca. It is distinguished from the latter by its very short or obsolete petioles (a character that appears to be consistent through large series of specimens), by a somewhat lower habit of growth, and by catkins which average shorter than those of S. glauca. In these northern British Columbia and southern Yukon. It may well be that S. niphoclada is not worthy of specific rank, and should be considered only as a subspecies of S. glauca. Salix brachycarpa Nutt. N. Am. Sylva 1: 69 (1842) short ones (at most 1-2 mm. long) ; catkins appearing with the leaves, on leafy peduncles, short-oblong to nearly spherical in shape, usually not more than 2 em. long; capsules grayish-wooly, up to 5 or long, sessile or nearly so, with short or almost no styles; bracts of the flowers of one color, yellowish-strawcolored or yellowish-brown, pube- scent with hairs that are much shorter than the bracts themselves; stamens 2; filaments hairy toward the base. Wynne-Edwards 8459 (CAN); bank of Henry Cr., Henry 146 (PH); near Norman R., Henry 105 (PH); near Akie Pass, Henry 555 (PH); is s i H Pa bd 4 p meadow in t Pass, 10665; gravel fan on east shore of Muncho L., 10354, 10855; Alaska Highway, mi. 472, near Muncho L., Taylor et al. 1595 (v); High- — way ard R., mi. 497, Taylor et al. 337 (V); Ww ow: muskegs along Liard R. south of Watson L., 11010, 11011, 11012; Highway near Cracker Cr., mi 988, Anderson s2 (s). (1957: 233) reports the typical species also from and York Factory in northern Manitoba. Map 10. A form with glabrescent leaves, called var. antimima (Schn.) Raup in Rhod. 33: 241 (1931) appears to be southern in the western and central parts of our area, but farther eastward it extends into the arctic tundra. It has been collected several times in the neighbor- ill and at k in oo) Bae | ; ‘ict of Macke and I have noted it in the Wood Buffalo Park of northern Albe 62 HUGH M. RAUP (1935: 122-3). It also occurs in the joes Rocky Mts.: Aylard Summit, Henry 439 (PH); Graham R., Henry 506, 507B, 509 (PH); Nelson R., Henry 549A, 550 (PH). Still eae variant is var. Deaiminophale Raup in Jour. Arn. Arb. 17: 230-1 (1936), known thus sk erect branches, broadly ovate (rarely obovate) leaves that are rounded to heart-shaped at the base and densely white silky-pubescent on both sides Hultén (1942:529-30) probably is correct in his opinion that typical S. brachycarpa has little to do with Alaskan willows that have been attached to it. Forms described by Ball as var. Mexiae of this species (1934: 412-14) seem more closely related to, if ar identical with, S. niphoclada Rydb. (see above). Salix cordifolia Pursh Fl. Am. Sept. 2: 611 (1814) Probably ee in our flora by its var. callicarpaea (Trautv.) Fern. in Rhod. 184 (1926), which is here a depressed or decum- bent shrub, with are ovate or obovate, entire or nearly so, petioled leaves that are 1.2-5 em. long, rounded or broadly obtuse at the apex and acute or obtuse at the base, hairy-pubescent on both sides though less so and glabrescent above, pale green above and glaucous beneath; catkins on leafy peduncles, the pistillate 2 cm. long or more, much longer than bee aise 6-10 mm. long, light grayish-pubescent, on very short ; bracts wlliwni ean short-hairy; staminate catkins up to 25 cm. long; stamens 2; filaments glabrous or hairy toward the base. Salix cordifolia is the principal eastern American segregate in the S. glauca group. It has been collected at the southern end of James Bay and along the Kazan R. in the vicinity of Yathkyed L., Porsild 5788, 6024 (CAN). Porsild (1957: map 122) places its western limits ; urst et and northeast of Great ot L, Chae of Hudson oe anal the more ates. erie ‘nlands carn Consequently there is an area of overlap between S. glauca and S. _ eordifolia in the Hudson Bay. region. In the region treated in this =: inet e seen also a collection that is identified as S. cordifolia ie Kluane L., #228t. The leaves and aetkinn are young, and perhaps not fully ex- panded (collected July 7). Salix cordifolia is not recorded by either Hulten or Porsild for Alaska, Yukon, _< northern British Columbia. Fi issi lia and its eastern — see Fernald oe or ions of S. cordif. ee (1928: 1818), 2 and bstind bones 105-8). ior ane nail, — about 3.5 cm. longs ponies “sessile or on cels less than 2 mm. long; Styles 2-2.5 mm. ee Re A i THE WILLOWS OF BOREAL WESTERN AMERICA 63 A. Leaves commonly 7-8 cm. long; capsules on pedicels up to 2 m long; styles 1.3 mm. long or j Rare Rcaeaeeware S. MacCa ey Salix MacCalliana Rowlee in Bull. Torr. Bot. Cl. 34: 158 (1907) oe upright shrubs, up to 2 m. or more in height, with an oe wn branchlets that << acat t when young but soon glabro wiakaés leaves rather firm and leathery in texture, elliptie lanceolate long by 2.5 ¢ or narrowly —— as much as 7 or 8 cm. wide though more commonly smaller, obtuse or acute at the base acute or acuminate * the apex, pubescent w. youn g¢ but i apt ' t green are a to ong, and with styles up to 13 1 mm. long or less; bracts Rowate abane: yellowish, short-hairy or glabrescent toward the tips; staminate catkins 2-2.5 em. long; stamens 2; filaments hairy at the base. Common in the Wood Buffalo Park in northern Alberta (Raup, 1935: 123) and occasional in the Peace R. region; Dixonville, north of Grimshaw, Moss 6143 (ALTA, GH); Deep Cr., B.C., Henry 675 the western shores of Hudson and James Bays and L. Winnipeg: oose R. estuary, Baldwin 1463 (CAN AN); Attawapiskat, James Bay, Dutilly & Lepage 15473 ene York Factory, Scoggan 6028 (CAN); Nelson R. 20 mi. northeast of Norway House, Scoggan 3003 (CAN); Hayes R., 130 nu peaseccesl pe L. Winkiges. Scoggan 5318 (CAN); Riverton, 75 mi. north of Winnipeg, Scoggan 9023 (cAN). Map 11. alix MacCalliana is a distinctive species which, superficially, has little resemblance to the S. glawea group, parti because of its serrate, non-glaucous, bright green leaves. It is related to this group, however, because of its clearly frogger catkins. I prefer to — rincipal species: ; Calliana group” which wi will sichide: ome closely related but poorly known S. Tyrrellii. It should be looked for throughout the ae part of our region, where it probably is more common collections indicate. It grows in damp sai at the OE Br prairies, lakes, or sluggish streams. Salix Tyrrellii Raup in Jour. Arn. Arb. 17: 231-2 (1936) Spreading shrubs, up to 2 2 m. high, with h reddish bark and reddish twigs Sak are glabrous even when young; leaves 2-3.5 cm. long, 4-8 mm. wide, jenrealate to elliptic, rarely oblanceolate, acute to acuminate at the apex, acute at the base, green and shiny on both sides, some ewhat marwuwts in texture, irregularly and minutely though dular-serrate_ on the margins, the teeth usually 2-3 2. . ; tkins 64 HUGH M. RAUP narrowly ormanannstd ies yellowish in the upper part, glabrate to- ward the apex; sty -) mm. long; staminate catkins unknown formed but immature catkins. ong, narrow, yellowish bracts, the coetaneous catkins borne on fonts peduncles, and the short-pedi- celled, pubescent ovaries relate it to S. glauca but it is closest to the unique S. MacCalliana. From the latter it differs in having leaves that are neh smaller and have fewer, more salient teeth. Its leaves are acute to acuminate, and ad elliptic or seme ei while in S. MacCalliana they are only subacute and com only oblanceolate. Its styles are 2-2.5 mm. long while in vhs oes species they are 0.8-1.5 mm. long KEY TO THE SALIX RIGIDA GROUP A. Twigs yellowish, though sometimes with reddish blotches or streaks; pedicels 1-3 mm. long - tea. A. Twigs reddish to brown, or yellowish only when very you B. Pedicels of the fruiting oy aga 3-4 mm. long; gestae glabrous rom the first, or nearl ackenzieana. B. Pedicels of the fruiting ‘capsules 1-1.5 mm. vase “leaves: pub- escent, at least when y S. rigida. Salix rigida Muhl. in Neue Schrift. Ges. Nat. Fr. Berlin 4: 236, t. 6, f. 4 (1803) Salix cordata Muhl., not Michx. See Fern ald, 1946: 27-38. Seeds up to about 3 m. high, the twigs reddish and pubescent when g but soon pipe leaves oblong-lanceolate, rounded to some- ea | ee at the base, short- acuminate at the Pig glabrous . broa a ery ie glabrous, ‘matched by sterile specimens from Albany and fe Dutilly & Lepage 16206, 15529, (GH). Map 12. Salix rigida is the easternmost element in a series of ovenapnine ‘arther west a THE WILLOWS OF BOREAL WESTERN AMERICA 65 to 7 em. long, one-fifth to one-fourth as wide as long, glabrous when mature, acuminate at the apex, rounded to somewhat cordate at the base, petioled, closely glandular-serrate or sometimes nearly entire on the margins, yellowish-green above and glaucous beneath; catkins 0. ments glabrous. Common on the higher parts of sand and mud bars in the Atha- ka-Peace delta, and northward to the lower Slave R. Reported ? katchewan. It is occasional on river sand bars in the upper Peace R. region, and I have collected it northward along the Alaska Highway to the Sikanni R. Map 12. Salix lutea is closely related to S. mackenzieana and S. rigida, from which it differs mainly in its yellowish twigs and yellowish- green leaves. Intermediates are common. Salix mackenzieana Barr. apud Anders. in Ofvers. Sv. Akad. Forh. 15: 125 (1858) Tall spreading shrubs, usually 2-3 m. high in our region; twigs brown, pubescent when young but glabrate in age; leaves lanceolate or oblanceolate, up to 1 dm. long, commonly one-third as wide as long, inate at the apex, more or less cordate at the base, glabrous, rather finely glandular-serrate, bright green above, glaucous beneath, borne on prominent petioles; catkins appearing with the leaves, the pistillate 4-6 cm. long, on leafy peduncles; capsules glabrous, 5-7 mm. long, on pedicels 3-4 mm. long; bracts of two colors, about 1 mm. long; styles about catki bout . stamens 2; filaments glabrous. Occasional to common along the Mackenzie R., northward prob- and it is likely that the Mackenzie material noted by Hooker in Fil. Bor.-Amer. 2: 149, “Between Fort Franklin and Cumberland House” belongs here. The species is common in the upper Peace R. region, and apparently occasional along the Alaska Highway northwestward ‘hitehorse, growing in p thickets on along the larger st _ Porsild (1951:48) made a single collection of it on the Canol Rd. west of the Mackenzie Mts., along the Pelly R. near mi. 136. Apparently unknown in Alaska. Map 12. It should be noted that S. mackenzieana is a northwestern phase in ‘a complex of closely related willows involving, from east to west, Se rigida, S. lutea, and S. mackenzieana. A more southerly phase in the _ Rocky Mts. is S. monochroma Ball (1921a: 431-5). 66 HUGH M. RAUP Salix Turnorii Raup in Jour. Arn. Arb. 17: 234-5 (1936) Erect shrubs narrowly pyramidal in form up to 3 m. high, with pale rayish-green bark except the young twigs which are red, the young em. wide, lanceolate, elliptic, or oblanceolate, acute to acuminate at the apex, acute to acunierat rounded at the base, on petioles 2-5 mm. long, pale poor on both sides, but a little paler beneath, somewhat leathery in textu finely and regularly glandular-serrate on the margins, sth beneath hd young but soon glabrous; pistillate cat- kins 1-3 em. long, 1-1.5 em. thick, borne on short leafy peduncles, appearing “with the Hou ae NE capsules about 5 mm. long, glabrous, reddish-brown, on pedicels 2-3 mm, long; bracts narrowly linear or linear-lanceolate, page a third the length of the pedicels, glabrous or thinly pubescent, reddish-brown; styles about 0.5 mm. long; stam- inate catkine un a own Known thus far only from its type locality among inland shifting Ath sand dunes south of William Pt., L. At ka, where it is common (Map 13). Salix Turnorii is unique among northern willows, and as yet I can only suggest a place for it among the usually recognized sections of the genus. Other novelties in Salix that were described from the same locality — S. Tyrrellii, S. stlicicola, and S. brachycarpa var. psammophila — can be related to other species or species groups, but S. Turnorii seems to be as unique in its way as S. Setchelliana, S. pyrifolia, S. arbuseuloides, S. argyro carpa, and others that stand alone. About the most that can be said is that with its glabrous, long- pedicelled capsules, and with its regularly glandular-toothed leaves that are sometimes rounded at the base, it suggests the Section Cor- datae. But beyond this eg likeness does not go, for the shrubs have a narrowly pyramidal e bark — one the red young twigs is conspicuously pale pen irigtily and the leaves are much smaller than those of anything in the Cordatae a S. myrtillifolia, to which S. Turnorii bears no resemblance otherwise Salix Farrae Ball, in Contr. U. S. Nat. Herb. 22: 321 (1921) Spreading shrubs up to 1.5 m. high, with past ap shiny branches, the ec shoots yellowish, finely hairy but soon glab- rate; lenves gfe lanceolate, or oblanceolate, up to 6-7 em. long and a third to a half as Wide, acute at the apex, acute or somewhat sil “sith — on | the cae or baoeiacagesd so, glabrous, at the base and brown at the apex, fe I mm or thinly hairy; styles 0.8 mm. long or less; staminate catkins about 1.5 em. long; stamens 2; filaments glabrous. Typical S. ares Se anaes Ye to common in swampy ground in ies ; fave refered fo Lao THE WILLOWS OF BOREAL WESTERN AMERICA 67 in Bot. Gaz. 71: 485 (1921). It differs from the above description in having more pubescent branchlets, brown seasonal] shoots, so ewhat larger and broader leaves with more conspicuous serration, and longer catkins. Both the species and the variety are to be exp in northern B. C. and Yukon, and should be looked for in the boreal h larg n the known distribution of S. Farrae s.1. that I have preferred to map it as a group of isolated populations (Map 13). Apparently it has not been collected in the Rocky Mts. of northern British Columbia, or anywhere along the Alaska Highway in British Columbia and Yukon, or along the Canol Road. It has been found in the Mackenzie basin only in the Caribou Mts., and eastward only in the neighbor- Ball, in a later publication (1942: 230-32), raised var. Walpolei to specific rank, took it out of the Section Cordatae, and related it to S. pyrifolia in the Section Balsamiferae of Schneider. Salix pyrifolia is a distinctive species, so much so that Schneider described a new sec- tion of the genus for it. The close relationship of var. Walpolei to S. Farrae is to me so obvious, and its lack of resemblance to S. pyrifolia so clear, that I prefer to retain it for the time being in its original position (see also Hultén, 1942: 532-3, under S. Farrae subsp. Wal- polei). KEY TO THE SALIX BARCLAYI GROUP leaves much longer than broad, the margins making an acute angle toward the apex but with the apex itself blunt ....... aosseecennees co Ceewease coal yrt A. Pedicels about 0.5 mm. long; shrubs commonly 1-2 m. tall; leaves 2-4 times as long as wide, ovate or obovate, acute at the apex -...- cae - : S. Barclayi. teeeeeee segeresewsouetesesneeeseeeeseenrer ers Salix Barclayi Anders. in Ofvers. Sv. Vet. Akad. Férh. 15: 125 (1858) ovate or obovate, acute at the a : rounded at the , petioled, crenate-serra r al entire on the margins, thinly entose above when young, but glabrate in age, glabrous th, but es green Or nearly so; catkins ap ing with the leaves, the pistillate up to 6 em. long and about 1.5 em. thick, borne on leafy peduncles; capsules 6-8 mm. long, catkins 1-3 cm. long; stamens ; filaments ae vie It will be seen from the above description that S. Barctayt 1s an ety eatebie species in growth habit, leaf structure and color, 68 HUGH M. RAUP and in its wide selection of habitats, Like S. myrtillifolia, its leaves have a tendency to blacken in drying, making their natural color often difficult to define. It is common in the northern Rocky Mts., occasional reported. Hultén (l.c.) has noted them with S. erassijulis (arctica), S. glauca, and S. d icola (padophylla). Suggests one with S. planifolia. On an alpine slope just above timber- line in Summit Pass o i lap in the northern Cordillera and in eastern Alaska, and the species i ed geographic segre- gates in a continuous population of Barclayi- or myrtillifolia-like willows that extends from the Aleutians to Newfoundland. Salix myrtillifolia Anders. in Gfvers, Sv. Vet. Akad. Férh. 15: 132 (1858) Low shrubs, usually less than 1 m. high and commonly only 2-3 dm., ba upright or spreading branches that are brown or grayish- 6- i - long; bracts 0.7-0.8 mm. long, gray-hairy, brown or grayish-brown; ri up ms 0.8 mm. long; staminate catkins 1.5-2.5 em. long; stamens Common to abundant in damp woods and thickets, particularly in vera, Muskeg thickets, throughout most of the forested parts of our Tegion, up to timberline in the ins, and northward to the arctic treeline (Raup, 1947: map, Pl. 21). Map 14, It is extremely vari- Ps stile and in the size and mossy muskegs it is a 1-3 dm. hi i i oo ‘igh, with small leaves, while on ves. and prairie margins it grows to 3 m. high THE WILLOWS OF BOREAL WESTERN AMERICA 69 and he leaves as much as 5 em. long. In ot plants there is a series of forms transitional to var. sinites (Anders.) Ball ex Hultén, Fl. Alaska & Yuk. 538 *(i9aa), “distinguished nd having leaves much longer than broad, the margins in; angle toward the apex but with the apex itself often blunt, potent n an the typical species, and by having slightly longer styles. I have found these forms especially common in the region along the a Highway between Dawson Creek an on, 537-8) cites them from central Alaska, the Alaska Range district, the central Pacific Coast district of Alaska, and from the upper Yukon (Anders.) Rydb. An eastern variant of the species is var. brachy- da Fern. in Rhod. 16: 172 (1914), distinguished mainly by Lehi of this variety from our region, but Dutilly, Lepage, and Dum: (1953: map, fig. mae indicate a record for it at the mouth of ie Moose Rs James unknown in western ‘arew n, ower Colville R. It is closely related to pecies — in the northern betwee! re KEY TO THE SALIX GLAUCOPHYLLOIDES GROUP A. Leaves somewhat leathery and thickened when full-grown, lanceolate, ovate, or obovate, the width commonly 1/3 to % e length eae «Se Lloides. A. Lesy ves not cer’ end? Phickened, broadly ovate, elliptic, or idth common se ease oe obovate, the pe with ton 0) pied Tegan Salix glaucophylloides Fern. in Rhod. 16: 173 (1914) Shrubs or small trees up to 5 m. high, the twigs usually brown, shiny, glabrous or soon glabrate; leaves oblong, lanceolate, or narrowly ovate, green ani and shi beneath, povietead leathery in rt 6 cm. long; capsules 7-10 mm. long, peduncles, the pistillate up to ee or mini tonite glabrous, on pedicels up to 1.5 mm. long; 70 HUGH M. RAUP ish, long-hairy; styles up to 1.5 mm. long; staminate catkins 2-4 cm. long; stamens 2; filaments glabrous. A species of north temperate eastern America, included here be- cause it has been collected in the Moose R. estuary on the southwestern shore of James Bay (see Raup, 1943: 111), and at Attawapiskat on the western shore of that bay, Dutilly & Lepage 15479 (GH). It should be looked for farther westward in Ontario, and in northern Manitoba and Saskatchewan. Map 15. Salix glaucophylloides appears to be closely related to S. pado- phylla, and I have therefore placed them together in a group. Their ranges overlap in northern Ontario. These things suggest that they are easte and western segregates in a continuous population, further understanding of which will require more detailed study. See note under S. padophylla. Salix padophylla Rydb. in Bull. Torr. Bot. Cl. 28: 499 (1901) Salix padifolia Rydb., ibid. 28: 272-3 (1901). Salix pseudomonti- cola Ball in Contr. U. S. Nat. Herb. 22: 321 (1921). Salix pseudo- monticola Ball var. padophylla (Rydb.) Ball in Jour. Wash. Acad. Sei. 28: 450-52 (1938). Shrubs usually 1-3 m. tall, but sometimes up to 4 or 5 m., with reddish-brown, shiny twigs that are glabrous except when young; leaves up to 6 em. long and commonly one-half to three-fifths as 8 g i & yah s 4 B 5 g J S 2 s é g 3 g e twigs or nearly so, 3-7 em. long; capsules glabrous, 6-8 mm. long, on pedicels 1-1.5 mm. long; bracts of two colors, commonly 1.5-2 mm. long, long-hairy; styles 1 mm. long or less; stamens 2; filaments glabrous. Oceasional in lowland thickets, at stream borders, and in aspen- willow woods in the Wood Buffalo Park (Raup, 1935: 124), in the reported at many places in the forested coun: west of Hudson and James Bays, in northern Ontario and Manitoba (Scog- gan, 1957: 235). I have seen most of the material upon which these records are and can find no real difference between them and the western specimens of the species. They constitute still another has h THE WILLOWS OF BOREAL WESTERN AMERICA 71 Salix padophylla appears to be a close western relative of S. glauco- phylloides, the ranges of the two species overlapping in the region of James Bay. The eaves of S. glaucophylloides are a little narrower to are born ile in S. padophylla ey ee sessile or nearly However, this is a variable chatactes in species. In his Seiginal description of S. pseudomonticola Ball te) eiatal that the catkins were “nearly ele ”” Later he reduced S. padophylla Rydb. to varietal status and attached it to his S. pseudomonticola, and in doing so he said that var. padophylla differed “from the species, S. pseudomonticola Ball, chiefly in having the aments borne on short leafy peduncles, where- base.” ¥: found the catkins sessile, or on short peduncles with or without 1-3 small leafy bracts. I have seen specimens of S. glaucophylloides, on the other hand, with very short peduncles (1 em. or less) petite only commonly have leafy peduncles, and I have seen a specimen from the Wiachuan R. on the east coast of the Bay, Dutilly & Lepage 14302 determined by Ball as S. pseudomonticola var. padophylla. Consider- ing these intergradations, therefore, I regard the character of the peduncles as scarcely definitive. The same may be said of the pubes- cence on the twigs. Some forms of S. glaucophylloides have a heavy pigtail on the young twigs, some of which persists for two or in some of the more southern forms the leaves are also cophylloides, in which the pubescence pe! ne yeal Pubescence on the twigs shows a similar “unilateral” variation in pulchra. In this pair of species it is the northwestern one, that has a variant with dense, persistent pubescence (var. sis). Salix cordata Michx. Fl. Bor.-Amer. 2: 225 (1803) Salix et ase Hook., Fl. Bor-Amer. 2: 146 (1839); not S. cordata Muh Shrubs up fi 1.5 m. high, with twigs densely pa er broadly lanceolate-oblong to pea, long-acumina\ ; Oe apex, rounded or heart-shaped a’ sre eg ge ers cen 9 cm. wide, glandular-dentate-serrate a e searioe re bine ee densely grayish-hairy and somewhat silky on hi nee Ree 8 white hairs; styles cels up to 1 mm. long; bi with long up to 15 mm. long; staminate catkins 2 em. long or less; stamens 2; fil Z : laments doar pee ie See wi ha ten collected on nig western and southwestern shores james a 72 HUGH M. RAUP along the lower Severn R., and at Weenusk on the shore of Hudson Bay at lat. 55° 15’, Dutilly "& Lepage 16894 (GH; cf. Raup, 1943: 111- 12). There is also a Churchill record based on a sterile specimen re- ported by Grgntved (1936: 32). The species should be looked for farther westward in northern Manitoba and Saskatchewan. Map 16. Salix commutata Bebb in Bot. Gaz. 13: 110 (1888) Shrubs 1-3 m. high, in our region ages 1-2 m., the ioe gray- hairy when young but glabrate in age; lea 8 cm. long and mesons 4 half ech i eles or nearly so, elliptic to iesadle oblanceo- egy veh rr obtuse and abruptly pointed at the apex, pata at the ae mitts or with a few inconspicuous glandular — on pedicels about 1 mm. long; bracts densely woolly, about 1 mm. long; styles 1-1.5 mm. long; staminate catkins 2-5 cm. long; stamens 2; filaments glabrous. T pare seen the following specimens of S. commutata from our open spruce woods ey small lake, Summit Pass, 10668; L. ‘Atlin, “Rasinor 665 (s); L. Bennett, Macoun, July 8, 1902 (s); White Pass, Eastwood 920, ae (CAN); same, Eastwood 879 (K); ee Eastwood 912 (Ss); Whitehorse, Eastwood 616 (K); Haines Rd., B.C.-Yuk. boundary, Taylor et al. 1079 (v); Haines Rd., mi. 75- 83, Taylor et al. 817, 928, 1381 (v); Alaska Range (southern slope), Viereoky 1053 (Hb. Argus). Porsild (1951: 144) n notes it as common meadows along the Canol Rd., and I have collected it at Brintnell L. in the Mackenzie Mts. (1947: 156). In Alaska it appears to be restricted to the southern coastal districts re 1942: 536-7; 1949: 1732; see also Raup, 1947: map, Pl. 21). It is = found in the Rocky Mts. of southern B. C. and Alberta. Map Forms with less conspicuous pubescence on the ee erat and more serration on their margins, have been referred by Ball aga 498; 1930: 333) to var. denudata Bebb in Bot. Gaz. 13: 111 (1888). Hultén (Le.) is inclined to disregard this variety, however, and ap- parently assigns forms with more leaf serration to a hybrid with "Atl S. myrtillifolia, Specimens from L. agway, narrow- leaved and he leaves somewhat glaucous ben he Is hybrids S. commutata and S. Barelayi. One of these, L. Atlin, Eastwood, 665, I have not hesitated to call S. com id maui rs (L.c.) notes specimens from the Canol Rd. that resemble the S. myrtil- apes hybrid, and an extreme form which he suspects is a hybrid of commu S. Chamissonis. i the time being I prefer to cover them with Bebb’s var. agggenentt ate allowing for ample variation in the characters that define it. Salix Chamissonis Anders. in DC. Prod. 167: 290 (1868) ‘ish twigs that are —— Prostrate shrubs with or brown: or slabrate; leaves up to 45 em. long and 2.5 em. wide, but us smaller than this, broadly obovate or sometimes nearly round, sean THE WILLOWS OF BOREAL WESTERN AMERICA 73 to rounded at the apex, wedge-shaped at the base, mag closely glandular-serrate on the margins, green on both sid s but paler beneath; catkins appearing with the leaves, the pistillate raboat 4 cm. Jong, on leafy peduncles; capsules 4 mm. long or more aragenced on very short pedicels or nearly sessile; bracts w to 2 mm blackish, hairy; bey about 1 mm. sageyi ee, catkins rey alin Chamissohee appears to be rare in our region, and —— to the western Arctic (Map 16). Porsild a 27) reports a tion of it from the Richardson Mts. west of the Mackenzie delta, poe ere are also records from the Rat and Porcupine Rivers. Hultén 1942: .509-10) cites specimens only from St. Lawrence Isl. in Bering from the Seward Peninsula, from the L. Iliamna region, and from the upland tundra between Fairbanks and Circle. Porsild did not find it along the Canol Rd., but cites material collected in Macmillan Pass on the Yukon Mackenzie divide (1951: 139) which he thinks may be a hybrid of S. C/ ‘hamissonis and S. arctica. I have seen a part of this collection, and am inclined to relate it most closely to S. arctica, though it has low, crenate teeth on the leaves. Salix pyrifolia Anders. in Svensk Vetensk. Handl. 6: 162 (1867) Upright shrubs, in our region 1-3 m. high, with shiny, reddish- brown or greenish branches; leaves gollceih only 3-5 em. long and more than half that in width, ovate to oblong-lanceolate, acute at the apex, rounded to cordate at the base, thin, delicate and nearly transparent when young, showing a purplish tinge at one stage of development, thin but rather firm when mature, green above, glaucous beneath, prominently Begeainta veined? beneath, glabrous, finely gland- ular-serrate on the margins, > pearing with the leaves, the pistillate up to 9 em. long and 2 cm. thick, borne on leafy peduncles; See tat widely spreading or refiexed, glabrous, 6-9 mm. long, on -4 mm. hairy, 1.5 mm. long or more; sles: i 5-1 mm. long; staminate catkins 2-4 em. long; stamens 2; filaments Lara noe plant giving off -like fragrance, which in specimens persists long and sloughs northward in ck the Wood Buffalo Park (1936: pes aca ag (1957: 235) reports i ani 3934 (BH); wet mossy abou’ e Beatton R., 10121, 10122, 10270, 10871; thicket on bluff ae Beaton R., 10271. The species ould be looked for in muskeg thickets throughout the upper Peace R. country, and northward at least to Nelson. Map 17. KEY TO THE SALIX RICHARDSONII GROUP linear- to half-cordate-lanceolate, acuminate, glandular- = paso nine 5e15 mim. omg --sseorveeesereeeer S. Richardsonii. 74 HUGH M. RAUP A. Stipules half-cordate to reniform, entire or glandular-dentate: petioles usually not more than 2 mm. long and rarely over 4 mm. long S. calcicolu. Salix Richardsonii Hook. Fl. Bor.-Am. 2: 147 (1839) Upright shrubs commonly 1-2 m. high, with stout, densely hairy a obtusish at the base, green above and glaucous beneath, petioled; stipules prominent and persistent for several years, long and narrow, with prominent glandular teeth around their margins; cat- kins appearing before the leaves, sessile on the twigs; capsules glab- rous, sessile or on very short pedicels; bracts hairy, about 2mm. long; styles 1.5-2.5 mm, ody staminate catkins up to 3.5 cm. long; stamens 2; filaments glabro _ Essentially an arctic- aaa species, though it is occasionally found in the edge of the timber or on stream banks and lake shores at low altitudes. It seems to be most sender in the tundra of northwestern ackenzie, Yukon, and Alaska, but has been found as far east as Hudson Bay and Baffin Isl. (Raup, 1947: map, Pl. 21). Hultén (1942: 538.9; 1949: 1732) gives it a wide range in northern and interior ‘Alaska, but apparently it avoids the southern coastal districts. Porsild (1951: 145) found it it common fag sd the cane} Rd. I collected it at several places in the mountains betwee ap Pass and central Alaska (see Porsild, Oey: 115, map 117). As above, S. Rich oni is. extremely variable in, leat Shape and oe but it is easily di b: of hairy twigs, smooth leaves and nasil and persistent ck stipules. It is one of a small group of willows in our region that have persistent depuis — S. pulchra, S. glauca var. stenolepis, S. var. S. sage ae S. Richard- sonii — all of which are easily told apart on other characters. Salix Richardsonii a close relationship to S. pag and I i “S. Richard. they are geographically segregated. I have not seen enou gh specimens from the area of ee to determine whether intermediates occur there. Salix calcicola Fern. & Wieg. in Rhod. 13: 251 (1911) nic or prostrate shrubs, up to 5 dm. high, with spreading, young twigs densely hairy, the older ones glabrous; above and glaucous beneath, th, acute ae Me ton short-acuminate at apex, rounded to heart-shaped at the base, borne on short, stout on margins, ie es ees ee gate in pecrwre Bog aie long; capsules glabrous, 7-9 mm. long, on very short pedicels; bracts THE WILLOWS OF BOREAL WESTERN AMERICA 75 blackish, ig hairy; styles 2-3 mm. long; staminate catkins up to 3.5 em. and sometimes almost as thick; stamens 2; filaments ea A predominently eastern American species that has been collected on the western coast of Hudson Bay at Cape Eskimo, Churchill (Raup, 1943: 113-14), and at the mouth of the Nelson R. (Porsild, 1957: 175, map 116). In more recent years it on been found on river gravels along the upper Saskatchewan R. in the vicinity of Banff (Porsild, personal communication), and should be looked for elsewhere in the northern Rocky Mts. It appears to be an eastern segregate in a small group of willows in which S. Richardsonii is its western counterpart. Map 18. Salix Barrattiana Hook. Fl. Bor.-Am. 2: 146 (1839) Low, upright oe usually 1 m. high or less, with pubescent twigs; leaves commonly 4-7 cm. long and one fo urth to one third that in width, ee? HA the apex, acutish to rounded at the elliptic to obovate, serrate on the margins or nearly entire, densely whitish or light grayish silky on both sides, petioled, usually with an upright habit of growth; catkins appearing before the i debtor es, sessile on the twigs, the pistillate 3-5 cm. long, usually upright in habit; capsules white-silky, about 6mm. long, on pedicels 1-2 long; bracts about 3mm. long, acute, black, silky-hairy; styles 1.2-2.5 mm. eres staminate catkins up to 4 cm. long; stamens 2; filaments glabrous. amp alpine meadows from the northern and spotty in Alaska, limited to the central and eastern = of a ieageets and the north slope. Cantlon & Gillis collected it in 1957 opie astern tisha (911, 1557, 1558). Porsild ( (4951. 147-48) found it long the ed at Yellowknife, on Great Slave L., by Cody ( 1956: 110). Map 17. i ws, and rather ly ed, ‘even at a ‘2 distance. a prsaa tip forms low, dense thickets, ligh’ h in color, with twigs, leaves and catkins scales, stipules and young sa ae oo the papers in which they dry are stained Va: es ul af shape were described by Andersson (1868: 2a): : var. sere and var. angustifolia, and Hultén on thi bel tifolia. sae mi coal ane ene hae : 216-17, : 1920), gave these variations little or no taxonomi ic ee if cance. "In 1947 I described a var. in Sargentia 6: en tBT-8 (1947), from the Mackenz‘e Mts., ipul cens, ere ae having persistent sti distinguished pacers by nted in the Northwest Territories, ggests that ies is re] Ye sO pico However, the specimens coll 76 HUGH M. RAUP by our field parties in the vicinity of the Alaska Highway, with one sect exception, do not show persistent stipules, and I am inclined the variety as somewhat more restricted than Porsild proposes The single exception is no. 11396 from a meadow just below timberline on Mt. White. There is an occasional 1-year-old stipule on this material. Salix amplifolia Cov. in Proc, Wash. Acad. Sci. 2: 282 (1900) Tall shrubs or ec trees, 3-8 m. high, with densely hairy young twigs that become glabrate after a year or two; leaves commonly 4-5 cm. long but coisa as much as 8 cm., usually at least half as wide as ee rn or obovate, obtuse or rounded at the ag acute or obtuse at e base, pubescent when young but glabrate in age except on a bens beneath, entire on the margins or serrate toward the base, short-petioled; catkins appearing with the leaves, 4-6 cm. long, about 1.5 em. thick, borne on rather bre — peduncles ; cap: aul glabrous, 8-9 mm. long, on pedicels long; bracts up to 3 mm. long, hairy, daviccplored at least jail ay the EDs: — slender, 2-4 mm. long; stamens 2; filaments hairy at the glabrous. Salix amplifolia is a northern Pacific Coast species that has been collected only in the region around Yakutat Bay (Hultén, 1942: 543), where it is conspicious on coastal dunes and on the lower slopes of the coastal mountains. Map 19. KEY TO THE SALIX ALAXENSIS GROUP A. Leaves glabrous or glabrate and bright green vee pe upper ensis. A. Leaves thickly covered on both sides with Enrees fomentum S. silicicola. Salix alaxensis (Anders.) Cov. in Proc. Wash. Acad. Sci. 2: 280 (1900) Shrubs varying from straggling alpine plants _ than 1 m. high, to lowland tree-like forms up to 6 or 8 m. tall, with t gs densely white woolly, or glabrous, yellowish or reddish, and Brie except w! very young; leaves 340 em. long, 2-4 em. wide, obovate or oblanceolate, acute or rounded at the grea acute at the base, entire at the margins, glabrous or glabrate and deep green above, corer —— with : cence, e inea: ceolate, 1-2 — ahsee.? or more, distantly Siemiolan on the margins, seovered heise! like the leaves with a dense white tomentum; a et appearing — leaves, sessile on the twigs, the pistillate . long; capsules densely Wwhite-hairy, 6-8 mm. long, sessile or on ae tae short pedicels; bracts blackish iaeard the the tips, 3 mm. long or more, with long white hairs; styles about 2 mm. long; staminate catkins 5 em. long or less; stamens 2; filaments glabrous. pi regenance a lat.) is one of the commonest and most m edge timber ‘ar saan ge oro and south in the alpine tundra of the Rocky THE WILLOWS OF BOREAL WESTERN AMERICA 17 Mts. to the Athabaska Pass. On the coastal mountains it reaches nearly to the southern end of the Alaska panhandle (Raup, 1947: map Pl. 21). The matted white pubescence on the under sides of its leaves makes it stand out sharply among most of the shrubs with which it grows. ree variants have been described for our area, to which the following key applies. A. Petioles expanded in the lower part, to form sheathes around the buds var. obovalifolia. A. Petioles not expanded to form sheathes around the buds ........+.+++0 Twigs, even when several years old, densely white lly S. ala is (typ.) woo! zensis B. Twigs soon glabrate, usually covered with a bluish waxy Dloom (pruimOse) «.....-sesescseessecsesesesesnsesnsnnneesnees var. longistylis. The typical form of S. alaxensis is the commonest one above timberline in the western mountains, where it grows in gullies, along watercourses, or in swales. Here it is usually a low shrub 0.5-2 m, tall. altitudes, where it is sometimes almost tree-like. The commonest Gard. 2: 163, 1901) which also grows to size on lake beaches ver bars. It also is found occasionally as a small shrub above timberline. Our field parties found it a itat at Fraser specimen of it from above timber on the Haines Rd., mi. 85, Clarke 536 (CAN). Intermediates between this variety and the typi Ss 0 up de mountains. Variety obovalifolia Ball in Jour. Wash. Acad. Sci. 28: 443 (1938) Wi Til material collected in Keewatin, and Ball gives ing that the variety does not have as a geograp segregation as Ball indicated, and that it may be only a minor variant. .3 above key, the following, noted by him, should be mentioned. He regards S. alaxensis as “tree-like, with gnarled trunk and branches, and longistylis as “bushlike, with straight branches”; young leaves of the species as “vi fragrant, fragrant”; pistillate bracts of the species as “Janceolate-oblanceolat acute, light brown,” and those of longistylis as obovate-obtuse, almost black.” Considering the amount and kind of variation I have seen in all of these characters among northwestern willows, and noting the strong resemblance between alazensis and longistylis, I 78 HUGH M. RAUP prefer ae maintain them, for the present at least, as a single species ple: Sake ‘alaconae is a unique ees except for a single, distinctive though closely related form that I described in 1936 as S. silicicola. ape aie I suggest that until more is known of the latter specie . alaxensis be considered the principal entity in a small group which will juehide both species. Salix silicicola Raup in Jour. Arn. Arb. 17: 246-7 (1936) Erect shrubs with spreading branches and reddish-brown bark, up to 3 m. high, the twigs pes ly white-hairy or White-tomentose; leaves 3-6 em. long, 2-3.5 em. , broadly elliptic to obovate, obtuse to rounded and slightly —— at the “apex, rounded or slightly un shore of L. Athabaska, but thus far unknown elsewhere (Map 20). It is a conspicuous willow, clearly related to S. alaxensis, trom Mich it differs by its short stipules (up to 7 mm. ths as agai that are broadly ovate to lanceolate rather than ies la eesti to filiform, and by having Ugloniee heavy, hairy-tomentose covering on both sides of the leaves instead of ee sores or glabrate and bright green on the nines surfaces. Also its leaves are consistently broader in arepertan to their a tik. ptow! of S. alaxensis, as well as more strongly obovate and obtuse. Salix candida Fliigge apud Willd. Sp. Pl. 4: 708 (1805) Upright shrubs, in our region usually 1 m. high or less, the twigs versie cuir in age, but covered with a white woolly pubescence ung; leaves 310" em. long, 0.5 to 15 cm, wie oblong or above. rather narrowly cylindric a form, usually sessile on “he capsules 6-7 mm. long, densely white-woolly, on apenraien 1 mm. oe or less; bracts pale brown toward the apex, a little over 1 mm. long; styles about 1 = long. staminate catkins 1-5 em. long; stamens 2; filaments glabro! Occasional Faget common in muskeg thickets and wet meadows in the more southerly parts of our region, and apparently sporadic north- ward in the Mackenzie basin. I have seen specimens from as far north as Churchill on the western coast coast of Hudson Bay (Raup, 1943: THE WILLOWS OF BOREAL WESTERN AMERICA 79 115), and Porsild cites material from Great Bear L. (1943: 27). My that the species is con: to r Ss in by Paleozoic or younger rocks in that area. Moss (1953: 212-52) reports S. at several poe in the Peace RL coun and just east of Lesse: an Salcha Slough near Fairbanks. Porsild notes it in Yukon only in the vicinity of Whitehorse (1951: 148). I have seen the following speci- mens from the vicinity of the Alaska Highway: Liard Hot Springs, 10908; same, A. E. Porsild 9065 (CAN); oe Mie 65 rae wet meadow along Liard R. south of Watson L., 11009; Whitehorse, A. se Porsild 9147, 10663 (CAN); Map 19. Strongly ieeenent teens be called forma denudata (Asine) Rouleau in Nat. Canad. 71: 266 (1944). Salix Bebbiana Sarg. in Garden & Forest 8: 463 (1895) Shrubs 2-6 m. high, sometimes becoming tree-like, the young twigs grayish with a pubescence that sometimes persists for several years; long and one fourth to one half that in margins, short-hairy or tomentose when young, sometimes persistently so id sometimes glabrescent, upper surfaces dull green and Pi rai appearing he levee usually on short leafy peduncles, the pistillate 2-5 on. long, often lax and nodding; capsules finely pubes- cent, up to 8 or 9 mm. long, slender, on pedicels 2-5 mm. long; bracts yellowish- ria sees 3 mm, long or a little more, sparingly hairy; styles none or ery: short; staminate catkins 1-2 cm. long; Common to abundant throughout most of the wooded parts of our region, in dry or Luigeega! well-drained eee (Map 21). It isa characteristic species of floodplain poplar and spru ce woods, prairie n aspen and 1 ey bobs and an bared over upland woods. In the West it reaches nearly or quite to the timberline and occasionally into see in the — Bay along the Canol Rd. northward to Sheldon L. (mi. 222). Hultén —— 544-5; 1949: 1733) gives it a wide range in the fe of Alaska, though i it appetrs to be abeent from most of the south- Several attempts have been made to ere from this species a form with thin glabrescent leaves, usually under the name S. perro- strata Rydb. in Bull. N. Y. Bot. Gard. 2: 163 (1901), or nes Bebbiana var. perrostrata (Rydb.) Schn. in Jour. Arn. Arb. 2: 7 (1920). Fernald has recognized it as a variety (1950: eee disregarding pubescence and using the following key characte: A. Mature leaves reticulate- beneath ........ - S. Bebiana (typ.). ves reti rugose * A. Mature leaves plane and scarcely reticulate pon ee 80 HUGH M. RAUP However, intermediates me these two forms are so abundant and widespread in our region, so common locally, that I am in- clined to agree with Hultén (1. = that with present knowledge they have but little taxonomic significance. (See also Raup 1948: 116-17; 1947: 159, for discussion of this problem.) In 1947 (pp. 159-60) I described S. Bebbiana var. depilis from specimens collected at Brintnell L. in the Mackenzie Mts. These plants are almost completely glabrous, even to the capsules, pedicels and axes of the catkins, and to the very a leaves and twigs. At most there are a few long ciliate hairs on the margins of the icon bracts. A few specimens collected by Mise. Alice Eastwood at Dawson and elsewhere in southern Yukon (nos. 238, 519, 566), found in the G Ep g Q 8 ey oi 3 an : g a 5 B 3 3 ® dency toward glabrescence that can be seen throughout the range of S. Bebbiana as a whole. Salix fallax Raup in Contr. Arn. Arb. 6: 149-50 (1934) Shrubs 1-1.5 m. high, the young twigs finely tomentose and reddish- brown, the older ones gray and glabrous; leaves 2-7 cm. long, 0.7-3.2 em. wide, hag 2 entire on the margins, acute or acuminate at the apex, acute at the base, silky when young but becoming glabrous, bright green ped and glaucous beneath, on petioles up to 1.3 long; pistillate catkins appearing with the leaves, 3-4 cm. long, 1-1. 2 thick, borne on leafy peduncles; capsules 4-6 mm. long, finely appressed-pubescent, on pedicels 1-2 mm. long; bracts pomadet ay: hairy, brown, about 1 mm. long; styles 0.5-0.8 mm. long; stamina’ catkins unknown. This species was first described from material collected in 1932 from open muskeg woods in the upper valley of the Peace R. a few miles east of Mt. Selwyn, Raup & Abbe 4312. It was not again recognized until Porsild collected ip a numbers of it from an open muskeg along the Canol Rd. at the lower Lapie R. crossing, near mi. 132: 9537, 9827, 9828 oer 149). Since then I have seen specimens by Cla in 1943 at the Liard Hot Springs (cAN), by Malte 122167, gy (cas) at Jasper, and by Argus at Fairbanks (413). Map 22. Saliz fallax seems most nearly related, judging by the character of = pistillate catkins, to S. pedicellaris, and I have therefore placed Pa the mountains of northern B, C. and southern Yukon. KEY TO THE SALIX ARBUTIFOLIA GROUP A. Prostrate or decumbent shrubs, the ascending branches usually not more than 3 dm. hi: B. Ovaries thinly pubescent or glabrous when young, glabrous a maturity; leaves more or less regularly and finely ccerase B. Ovaries pubescent to maturity or becoming glabrate in age; leaves entire all around the margins; repent or prostrate shrubs usually 1-3 dm. high S. hebe THE WILLOWS OF BOREAL WESTERN AMERICA 8L A. Upright or — shrubs 8-10 dm. high. C. Ovaries and mature capsules glabrous semess usually 2-4 mm. long; leaves glabrous, dark bluish-green above; entire Ss. licellaris. C. Ovaries short-appressed-hairy, remaining so in age or be- coming glabrous at maturity; pedicels 1-3 mm. long; leaves Leaves glabrous, more or less regularly and finely — around the lower half; pedicels 1.5-3 mm. ug ovarie: glabrous at maturity unto. D. Leaves thinly appressed- -hairy, entire or cinutely toothed around the whole margin; nee cels 1-1.5 mm. long; ovaries Sa thinly hairy, or glabrate in age thabascensis. Salix arbutifolia Pall. Fl. Ross. 1, 2: 79 (1788) Trailing shrubs only a few centimeters high, or ascending-spread- ing and up to 6-7 dm. high, with glabro us twigs; leaves 1-3 em. long and commonly more than half that j in gabe eh elliptic to obovate, glab- ae obtuse or rounded at the apex, obtuse or acute at the base, and glaucous beneath, petioled; catkins appearing with the leaves, the pistillate about 2 cm. long, on n leafy peduncles; capsules glabrous or sometimes pubescent en young, 5-8 mm. long, on pedicels 1.5-3 mm. long; bracts about 1 mm. long, hairy, dark-colored toward the apex; styles very nna or nearly obsolete; staminate catkins up to 1.5 em. long; stamens 2; filaments glabrous. A western arctic species whose range in Alaska is given by Hultén (1942: 545-6) as chiefly in the ‘west and north, with eastward exten- sions in the Alaska and Brooks Ranges. It has also been found in the tundra region between Fairbanks and Circle, and Argus has collected it in a muskeg at Fairbanks. It reaches the southern coast only from Kodiak Island westw: tward. Porsild (1943:27) cites material from i eas! a I have seen a specimen also from Back R., Tener 353 (CAN)- Porsild a not find it along the Canol Rd. Map agree with Hultén (l.c.) that this plant is the S. fuscescens of Anion Bhs 230) and of Coville (1901: 329). I — that he is correct in reducing to it the Pra “described all as Fiaaiboeess var. ‘chat (1935: oe Salix arbutifolia was the fi scribed in a series of willows that appear — aie related " ie called an S. arbuti- folia group. This group includes S. pedicellaris (var. hypoglauca and te seens in our Soa: 'S. athabascensis, and S. hebecarpa. Salix anada and to Tee an 82 HUGH M. RAUP Salix hebecarpa is a localized segregate known in the east only from its type locality on the Gaspé Peninsula, and reported by Hultén from the Seward Peninsula of Alaska. The large areas of overlap between S. pedicellaris and S. athabascensis suggest that the latter Ww of its range I prefer to maintain it as a species. It reaches consider- ably farther westward than S. pedicellaris, and has not yet appeared in the easternmost parts of the latter’s range. Salix pedicellaris Pursh, Fl]. Am. Sept. 2: 611 (1814) Represented in our region chiefly by its var. hypoglauca Fern. in Rhod, 11: (1909), to which the following description applies: Shrubs usually 1 m. high or less, with glabrous, blackish twigs; leaves 2-4, 5 em. long, 1-2 em. broad, obovate-oblong to broadly oblan- ceolate, usually obtuse or rounded at the ape. acute or obtuse at the lower surface, glabrous, green above nor glaucous beneath; catkins appearing with the leaves, on leafy peduncles, usually about 2 cm. long; capsules glabrous, 5-8 mm. long, on pedicels 2-4 mm. long; bracts about 1 mm. long, vile aah or nearly so; styles cari obsolete; stamens 2; ‘labro' southerly species in the feread aa. ce found in muskeg dace and having a range in our region somewhat similar to that sf ida. In the Hudson Bay country it is occasional or common northward to Churchill (Raup, 19438: 117-18) and to a lake ee on the Tha-anne R. northwest of Churchill in southern bce Sad sild, 1943: 28). In the Mackenzie basin it is common the sand. shores = ponds just south of L. Athabaska, at Secunia ul nacimwetd to Great t Bear L. (Raup, 1936: 238; 1947: 160). Apparently rare in upper Peace R. region of western Alberta. Hultén makes no mention A second variant of the species, though unknown elsewhere in our pm : be (1909), and was found in an upland muskeg (Raup, 1935: 125; 1936: 238). It is distinguished by having oblanceolate to linear- oblong leaves that are acute at both ends and not more than 1 cm. wide, and by having rather lax, loosely-flowered catkins with cap- sules up to 10 mm. long. i dicellaris is here regarded as closely related to a group of species including S. arbutifolta, S. athabas and S. hebecarpa. See S. arbutifolia for further discussion of the group. Salix athabaseensis Raup in Rhod, 32: 111-12 (1930) rect shrubs 3-6 dm. high, with finely gray-hairy young twigs; — 1-3 em. long, 0.5-1.5 em, wide, elliptic or obovate, entire or minutely glandular-dentate on the margins, acute or acuminate at the THE WILLOWS OF BOREAL WESTERN AMERICA 83 apex, obtuse to rounded at the base, green and thinly silky-pubescent above, becoming glabrous in age except on the mid-vein, glaucous and thinly appressed hairy beneath and glabrate in age, on petioles up to 6 mm. long; pistillate catkins 1.5-2 cm. long, borne on leafy peduncles; capsules 5-6 mm. long, covered with short appressed hairs on pedicels 1-1.5 mm. long; bracts 1-1.3 mm. long, obtuse or rounded at the apex, brown, silky-hairy; styles about 0.3 mm, long; staminate catkins unknown. Apparently common locally, though in widely scattered places. I have seen the following specimens from the western part of our area: wet meadow near Liard Hot Springs, 10907; same, Clarke 64 (CAN); margin of small pool in willow-spruce woods near Liard R. south of Watson L., 10998, 11001, 11002, 11005, 11007; thickets and wet meadows at the eastern end of Watson L., 10921, 11050, 11061, AN ‘ew the south of it are also referred to S. athabascensis: York Factory, Scoggan 6026 (CAN); Riverton, 75 mi. north of Winnipeg, Scoggan 9024 (CAN); north of Prince Albert, Sask., Macoun 13671 (CAN); Goldburn, Sask., Breitung 456, 458 (CAN); McKague, Sask., Breitung 461 (CAN). Map 23. The type material of S. athabascensis was dried over rather intense artificial heat which drove off the glaucescence of the leaves. This to the type. < okt In the S. arbutifolia group of willows, of which this species is a part, it is most nearly related to S. pedicellaris var. hypoglauca, and may prove to be no more than a varietal segregate of the latter. Salix hebecarpa Fern. in Rhod. 26: 123 (1924) Repent or prostrate shrubs 1-3 dm. high, with glabrous or glabrate twigs; leaves up to 3 cm. long and 1-4 cm. wide, elliptic-oblong to oblong-ovate, obtuse or rounded at the apex, mostly obtuse at the base, entire Seward Peninsula by Walpole (1826, US). Otherwise the species is i i M aspé Ss is a close Ball (1950: 10-16) considered that S. hebecarpa icellaris var. hypoglauca and S. Twi cies Salix hebecarpa appears to be closely related to S. arbutifolia and S. pedicellaris, and I have included it in what I have called the S. arbutifolia group. 84 HUGH M. RAUP Salix gracilis Anders. in Ofvers. Sy. Vet. Akad. Férh. 15: 127 (1858) S. petiolaris of auth., not ay var. rosmarinoides (Anders.) Schn. in Jour. Arn. Arb. 2: 19 (1920). Shrubs 1-3 m. high, with slender erect or ascending branches, the twigs glabrous or soon glabrate; leaves up to 6 or 7 cm. long when 2g. r or becoming glabrate in age, shiny green above, glaucous beneath; cat- kins appearing with the cone the pistillate up to 4 em. her in fruit but shorter when in flower, borne on leafy peduncles; capsules silvery-silky, 5-7 mm, long, on panes pedicels 2.5-4 mm. long; bracts narrow, 1-2 mm. long, yellowish or brown, thinly pubescent; styles obsolete or nearly so; stamens 2; filaments hairy toward the base. Saliz gracilis is Hd nad in the country west of James and Hudson Bays as far north as MacBride L., 56° | 52’ N., 99° 57’ W. (Scoggan, rl: zoic or younger rocks (Raup, 1936: 238; 1935: 125; 1947: 160). Farther west I have seen specimens from the Lesser Slave L, district, Brinkman 3870 (BH); Dawson Creek, Raup & Abbe 3503, 3532, 3534, 3539 (A); Fy and Cache Cr., north of the Peace R., Henry 676 n, how (S. petiolaris var. rosmarinoides in his treatment, l.c.) a re southern range in Manitoba, with its northernmost colleetion fame Norway House. He rega the form seen in the region west 0: Hudson Bay as var. textoris Fern., in Rhod. 48: 46 (1946) (S. petio- laris of auth., not Sm.). This variety is tarcnagincagig by having somewhat jonger capsules (up to 9 mm.); leaves up to 10 cm. long setae cm. bens glabrate or rarely silky when inwouee, strongly glan- a! In using et name S. gracilis Anders. I am following Fernald (1946: 46-8; 1950: 516-17). The species (sens. lat.) has long been called, in American studies of the willows, S. petiolaris J. E. Smith, Trans. Linn. Soe. 6: 122 (1802). Forms with more delicate, less serrulate leaves (here called S. gracilis) have been named, in 8. = tiolaris v: var. rosmarinoides poner) Schn. Fernald (Rhod. ot the common Amer but was ive ss the British Isles. Ball (1948: 178-87) has taken issue with = parr that Smith’s plant was from America, prob- ably introduced to Britain by persons unknown. In the absence of any firm evidence on psig geographic origin of asin plant, _argu- to S. arbus like and single-stemmed. Ball reports that G. H. Turner has found a few single-stemmed plants of the petiolaris type in in aspen woods near eal THE WILLOWS OF BOREAL WESTERN AMERICA 85 involves more assumptions than Fernald’s, which does not go beyond Smith’s simple statements that his S. petiolaris was found in Britain, and that it was a tree. KEY TO THE SALIX DISCOLOR GROUP i a long; leaves lanceolate, obovate, or elliptic, acute at the apex, more or less regularly alternate toward the ends of the twigs .... discolor. A. Catkins precocious or appearing with the early leaves, sessile or on very short leafy peduncles; pedicels 0.5-1.5 mm. long; leaves arranged so that they appear fan-like at the ends of the twigs S. Scouleriana. Salix discolor Muhl. in Neue Schr. Ges. Nat. Fr. Berlin 4: 234 (1803) Shrubs or small trees up to 6 m. high, with glabrous or glabrescent gs; leaves 3-10 cm. long, 1-3 em. wide, lanceolate, obovate, or sistent rusty pube: beneath; catkins appearing before the leaves, and nearly mature be- fore the leaves come out, the pistillate up to 12 cm. long and 3 cm. thick at maturity, sessile on the twigs; capsules 7-12 mm. long, finely pubescent, on pedicels 1.4-2.4 mm. long; bracts dark or brown to black, about 2 mm. long, long-hairy; styles distinct, about 0.8 mm. long; staminate catkins 1.5-5 cm. long; stamens 2; filaments An early-flowering precocious species of temperate eastern America, known in our region in Manitoba northward at least to the Hayes ae (Scoggan, 1957: 237), and in the Peace R. country east of the Roce! latter I can cite the following specimens and records with some degree of confidence: Lesser Slave L., Brinkman 3869; Grand Prairie, Groh 886 (DAO, A). Although I have seen no other specimens than these, Moss (1953: 212-52) reports the species from many localities in the ‘rom ave L. to Fort S. John, down the Peace to Vermilion, and northward on the Mackenzie Highway to Steen R. He notes it as occurring in a variety of forest types, from tamarack bogs to pine woods. Map 25. The eastern S. discolor and the western S. Scouleriana are closely related species, and I have placed them together in an “S. discolor 86 HUGH M. RAUP group.” Their ranges overlap in the forested country between north- ern Manitoba and the upper Peace R. region just east of the Rocky Mts. Saliz Scouleriana Barratt in Hook. Fl. Bor.-Am. 2: 145 (1839) ee in our region chiefly by its var. coetanea Ball in Jour. commonly one third that in width, oblanceolate or obovate, obtuse or rounded at the apex and sometimes abruptly acute, wedge-shaped at the base, entire or sometimes shallowly crenate-serrate, rather thick and firm in texture, dark green and glabrate above, glaucous and sometimes silvery or reddish pubescent beneath, petioled, commonly ey kins appearing with the leaves or a little before, sessile on the twigs or on very short leafy peduncles, the pistillate up to 5 em. long; cap- sules 7-9 mm. long, Romero on pedicels up to 1.5 mm. long; bracts black, obovate, long-hairy; styles about 0.5 mm, a staminate cat- 1-2 em. long; stamen: ns 2; filaments glabro Salix Scouleriana is a close western relative S. discolor. In the central Mackenzie basin it is known only around L. Athabaska, where it is common (Raup, 1936: 238-9). In the Alaska Highway region it is oceasional to common northwestward at least to about mi. 1381 in the Tanana valley, and Argus (personal communication) notes it as common in muskeg spruce woods in the Fairbanks area. However, the range does not seem to be continuous, for it has not been found in southwestern Yukon west of Whitehorse. Hultén (1942: Pees ran eh edie it a limited range in southern coastal Alaska ukon valley northward to the vicinity of Dawson. Fors epee 151) found it common along the Canol Rd., mostly in oN a has peat it also along a moe tributary of the Godlin R. g the Canol Rd. east of the Yukon-Mackenzie divide (1945: = sae I found % in the Brintnell a tall willow growing in the river floodplain. The catkins are normal and were maturing in the second week of July, and they are typical of S. Scouleriana; but the leaves are elliptic to elliptic-ovate and sharply acute at the apex. They can be matched in the herbarium, but they Alaska Highway are mainly from dry pine or aspen woods. The specimens from near mi. 1381 came from well-formed, single-stemmed trees about 10 m. high with trunk diameters up to 13 cm. Seoggan (1957: 238) reports several specimens from central Manitoba, and one collected by Baldwin. at Reindeer L., lat. 57° “37 N. I have not seen this material, but in view of similar eastern range cccictaigns OF thas ‘weltern’ ypecien, 5. Sensleriane tu. un te expected in this region. — THE WILLOWS OF BOREAL WESTERN AMERICA 87 Plants whose leaves have a dense, persistent pubescence beneath have been called forma poikila by Schneider in Jour. Arn. Arb. 3: 76 (1921) (see also Jour. Arn. Arb, 2: 12, 1920). Our field parties on woods along Lewes R. at Whitehorse, 11175. I have also seen speci- mens of it (GH) from Dawson, and from L. Athabaska. Salix Scouleriana appears to be the western vicariad in a group of willows in which S. discolor is the eastern element. In spite of close relationship the two species are rather easily distinguished on leaf and other characters. KEY TO THE SALIX PHYLICIFOLIA GROUP A. Leaves finely and more or less regularly serrate on the margin, though some of them occasionally nearly entire; catkins usually taneous (sometimes subprecocious), sessile or on promin- ently leafy peduncles . S. phylicifolia. A. Leaves entire or with a few small, egularly spaced teeth; cat- kins precocious, sessile or with a few small leaves at the base. B. Stipules persistent for 3 or more years as dried, linear or linear-lanceolate appendages on old wood S. pulchra. B. Stipules not persistent after one growing season . Se .. S. planifolia. Salix phylicifolia L. Sp. Pl. 2: 1016 (1753) above and glaucous beneath, finely serrate on the margins or some- This speci i . § phylicifolia has not been recognized on the North American Continent i rsh demonstrated that its closest relative in eastern America was a distinct species, which he named S. planifolia. —— a deal Herbarium that I have no other course at present than to identify it as such. The shrub was about 2 m. high, and when seen in mid-June had well-formed leaves, with catkins approaching maturity. Since then I have seen a second collection, made by Dr. William Drury along the lower Kuskokwim R. and identified by him as Ss. phylicifolia. It came from the hills 2 mi. north of Aniak (1507, GH, pistillate shrub about 11 dm. high). Although its leaves are more in our no. 11785, the two collections otherwise match each other closely. Map 26. ‘p 88 HUGH M. RAUP Salix phylicifolia belongs to a group of three closely related species in America, of which the other two are S. planifolia, and S. pulehra. I have called this series the “S. phylicifolia group,” but if S. phylici- folia proves untenable in the American flora, the group should be named after S. planifolia, the next older name. See note under S. pulchra for further discussion of the group. Salix planifolia Pursh, Fl. Am. Sept. 2: 611 (1814) Shrubs up to 3 m. high, with purplish twigs that are glabrous and sometimes pruinose; leaves 2.5-7 cm. long and one-fourth to one- third that in width, elliptic or elliptic-lanceolate to oblong or oblanceo- late, usually acute at both apex and base, glabrous, firm and some- ommon to abundant on stream banks and at wet slough margins westward in the forested regions to the Rocky and Mackenzie Mts.; cceasional in the upper Liard R. valley and westward over the Con- tinental Divide to Whitehorse ( Map 26). In its eastern range it ex- tends northward in the tundra to the Chesterfield Inlet-Dubawnt L. region west of Hudson Bay, and to southern Baffin Island (Raup, the lower Mackenzie R. valley. Hultén does not mention it in his Flora of Alaska and Yukon, but Porsild (1951: 150) reports it Forms with narrowly oblanceolate, narrowly lanceolate, or narrowly elliptic leaves have been called var. Nelsonii (Ball) Ball in Amer. Mid. Nat. 45: 747-9 (1951). These forms occur throughout the range of the species, and like the varieties of S. pulchra based on leaf-shape, shi at margins of wet meadows to dwarfed spreading forms only a few cen’ TS in exposed rocky places. The form of its bi ies along with these differences, twigs commonly pruinose, pubescent when young but soon glabrate; leaves usually 3-6 em. long, and from one-third to one-half that in THE WILLOWS OF BOREAL WESTERN AMERICA 89 width though sometimes narrower, elliptic or in some forms oblong or obovate, usually acute at both apex and base though sometimes (mostly in obovate forms) rounded or abruptly acute at the apex, glabrous, reen above and glaucous beneath, entire at the margins, or with a cels; bracts about 2 mm. long, blackish in the upper part, hairy; styles 1.2-2 mm. long, staminate catkins 2.5-3.5 em. long; stamens 2; filaments glabrous. Salix pulchra is common in Alaska and Yukon, with minor exten- sions southward into northern B. C., and eastward into the District of Mackenzie. Along the Alaska Highway it is common northwest- ward from Summit Pass in the Rocky Mts. Hultén (1942: 547-9; 1949: 1733) gives it almost universal distribution in Alaska except for the southeastern panhandle. Porsild (1951: 149-50) found it common along the Canol Rd. up to the Yukon-Mackenzie divide, and I collected it near Brintnell L. in the Mackenzie Mts. (1947: 160-1). Eastward along the Arctic Coast it reaches beyond the Mackenzie delta, where it is noted by Porsild as common, to Berna! Coppermine (1943: 28). I have seen a single spe Mackenzie valley, labeled “Fort Franklin, Mackenzie River,” Richard- son (NY). is species is a characteristic tundra plant of the far Northwest, and occurs below timberline in wet meadows, muskegs, woods. Map 26. Three varieties of S. pulchra have been described from Alaska and will indicate the trends. A. Twigs densely pubescent, and remaining so for at least 2-3 var. yukonensis. year: e A. Twigs glabrous or essentially so, even when very young Leaves rather broadly rhomboid or elliptic, geet be both ends Leaves narrowly to bi ly obovate, or obovate-oval, ror to abruptly acute at the apex, or the newest ones acute ........ pce .. var. Looffiae. C. Leaves narrowly oblong, elliptic-oblong, or narrowly tl bl late a Henke Aa meri. (Lc.) f var. yukonensis i Arn. Arb. 1: 72 (1919) to the interior of Alaska, and to the Lewes- Yukon valley in Yukon. Porsild (Le.) states that along the Canol Rd. it is limited to below timberline. thinks it is a dis- tinctive variety, perhaps worthy of ific rank. Our field parties on the Highway collected it the mountains north of Kluane L., where it was growing in alpine tundra. Variety Looffiae Ball in Madrojio 6: 228-9 (1942) was described from Kodiak Island material, but Ball cites specimens from several places in 90 "- HUGH M. RAUP Alaska, the islands in Bering Sea, and from the interior between the Kobuk R. and Circle. Variety Palmeri Ball in Madrofo 6: 229-30 (1942) came originally from the Matanuska Valley, though specimens are cited from many places in central southern coastal Alaska, and from central and west-central interior Alas! Considerably more significant, for the moment, than this attempted definiti tion of varietal segregates within S. pulchra is recognition of to the eased of Whitehorse. Thus the two species Crate 4 in north- B. C. and sow i i i all of which occur in both species, the principal character by which pulchra may be distin: tinguished is the persistence of its stipules. An additional character not found in S. planifolia is the occasional pres- f dense pubescence on young twigs of S. pulchra, persisting for 23 years or ae but this is limited to that part of the population that has been called var. yukonensis. If our record of S. pulser in S. W. Yukon proves to be correct, this species should added t the S. planifolia-S. pulchra complex, to form the “S. phyliesfolia up.” KEY TO THE SALIX PELLITA GROUP A. Lower surfaces of the leaves silvery-tomentose with ee straight, appressed hairs; a western Cordilleran species ... A. Lower surfaces of the leaves silky-tomentose with laneed scarcely straight hairs that are slightly appressed; an eastern boreal forest species, closely related to S. subcoerulea ............ S. pellita. Salix pellita Anders. in Sv. Vet-Akad. Handl. 6: 139 (1867) Shrubs or small trees up to 5 m. high, with greenish or — 4-12 cm. ite, acuminate at the apex, acute or obtuse at the base, rather thick and firm in texture, entire on the margins or nearly so, green and glabrous above, glaucous and usually densely silky-pubes- cent beneath, producing a silvery appearance; catkins appearing be- fore the leaves, nearly sessile or subtended by a few small leaves, the pistillate up to 5 cm. long; capsules 4-5 mm. long, densely whitish pubescent, on very short pedicels; bracts 1-2 mm. long, blackish, long- hairy; styles up to 1.2 mm. long; staminate catkins unkn Salix pellita is an eastern counterpart of the western S. subcoeru- lea, to which it is closely related. Both should be considered members THE WILLOWS OF BOREAL WESTERN AMERICA 91 of an “S. pellita group,” to which the central Cordilleran S. bella Piper can be added. Salix subcoerulea Piper in Bull. Torr. Bot. Cl. 27: 400 (1900) Shrubs 2-3 m. high with brown to blackish, glabrous, pruinose twigs; leaves usually oblanceolate though sometimes oblong-oblanceo- late or oblong-lanceolate, 3-6 or 7 cm. long, 0.8-2.5 em. wide, acute or cence, petioled; catkins appearing before the leaves, sessile on the twigs, up to 4 em. long, and 1 em. thick or less; capsules about 5 mm. long, silvery-silky, on short pedicels up to 1 mm. long; bracts 1.5-2 mm. long, brown to blackish, thinly hairy; styles up to 1.5 mm, long; stamens 2; filaments glabrous. A Cordilleran species that comes into our region only in the upper Peace and Athabaska R. regions, the northern Rocky Mts., and in the upper Liard valley. The easternmost collection I have seen came from Athabaska Landing, just east of Lesser Slave L., Ball 2364 i it sheen of the lower surfaces of its leaves. It grows in wet meadows and on stream margins at low altitudes, and is a close relative of the eastern S. pellita (see note under S. pellita). Map 27. Ball (1951: 740-47) has changed the taxonomic status of S. swb- making it and S. bella Piper varieties of Salix Drummon- diana Barratt. In doing so he raises no question with to the typification and identity of S. Drummondiana itself, though this has been a debatable subject in American salicology for a long time. I vi i co. clusion that the species was of doubtful validity, and that its relation- ship was in the Sect. Chrysantheae with S. alazensis rather than in the Phylicifoliae with S. subcoerulea and S. pellita. At that time I had seen a photograph of what had been thought to be the type in the e' i of Kew Herbarium, two i the New York Botanical Garden, and a specimen in the tt Her- barium at Wesleyan University, Middle , Connecticut, labeled by Barratt himself as the type 0: col plant. They were collected Mountains,” probably in the region between the Saskatchewan and ‘Athabaska Rivers, and Barratt’s description was published in Hooker’s Flora Boreali-Americana 2: 144 (1838). The type material noted above shows certain characters that are of particular significance to the present discussion. The twigs are dis- tinetly pruinose. ider (Jour. Arn. Arb, 1: 87-9, 1919), studying limited material and with only a photograph of what he considered to 92 HUGH M. RAUP be the type (K), said that the species showed “almost an entire absence of the glaucous bloom of the branchlets.” The under surfaces of the leaves in the type specimens are thickly felted with a whitish, opaque ee and are not silvery in hcpeaatea es) ao th of these are imperfectly rene ath 3 appearing as though the shrub from which hey came were an F; hybrid of S. alaxensis var. longistylis and some other willow. Ball’s recent and rather detailed description of S. Drummondiana (Lc.) seems to have been made up from composite sources. In his citations of tn ea he mentions none of the type material, but he een influenced by Schneider’s statement that this species was aoe SHainose on the branchlets. On the other hand, he describes the andes surfaces of the leaves as “densely silvery-tomen- tose,” which suggests again that he had not seen any of the type material. Otherwise the SS. apparently was based largely upon specimens in the ee lea complex that answered to the mistaken requirements athse Bate a am inclined to look unon a mmondiona, as I did in 1934, as ormal form most closely related to the S. alaxensis group, but wiethiee the abnormality is due to local site conditions or to hybridiza- tion is too uncertain to allow any conclusions at ee If it could be proven that S. alarensis ani then the latter would have precedence, for it is the tee name. Salix arbusculoides Anders. in Vet. Akad. Handl. (Stockh.) 6: 147 (1867) Low shrubs or tall spindling trees up to 3 or 4 m. high with trunk diameters of 5 or 6 cm., the twigs reddish and shiny, glabrous or glabrate; leaves 2-6 em. long and commonly one-fourth to one-third hairs; catkins appearing with the leaves or a little earlier, sessile or with 2 or 3 small leaves on a short peduncle, the pistillate up to 5 cm. long; capsules 4-7 mm. long, pubescent, on pedicels 0.5-1.2 mm. long; bracts about 1 mm. long, blackish, sang sth up to 0.8 mm. long; staminate catkins up to 2.5 em. long; stamens 2; filaments glab- rous. A common species throughout most of the forested parts of our i las! common throughout the Mackenzie basin northward to the Mackenzie delta (Porsild, Le.; Raup, 1936: 239; 1947: 161), and along the ighway from western Alberta to cent tral Alaska. Hultén (1942: 549-50; 1949: 1733) gives it a wide range in the interior of Alaska and in the upper Yukon above 3S have seen several specimens collected tise hs lowe Colville R. in THE WILLOWS OF BOREAL WESTERN AMERICA 93 the vicinity of Umiat, Bliss & Cantlon 5058, 5112, 5131, 5262. Porsild (1951: 150-1) says that it is common in lowland spruce forest along the Canol Rd. northward to the vicinity of Mt. Sheldon, where it was found at altitudes up to 3500 feet. Although there is a record for it p 28. ‘Salir arbusculoides is a distinctive willow, growing mostly in low- end woods and thickets, but occasionally in rocky places above tim- loan spruce woods it often forms tall, slender, single-stemmed . Though extremely variable in growth habit, its leaf and catkin arses rs are remarkably uniform except for occasional glabrous- leaved forms that have been called var. Taber Anders. in Sv. Vet.- Akad. Handl. 6: 148 (1867 Salix sitchensis Eee in Bong., in Mém. Acad. Imp. Se. St. Pétersb. Ser. 6, Math. "2: 162 (1833) Shrubs up to 7 m. high, with dark brown to blackish twigs that are glabrous or occasionally pubescent; leaves commonly 3-7 em. long, 1.5-2.5 em. wide, oblong-obovate or strongly obovate, acute at the apex or occasionally obtuse, wedge-shaped at the base, entire or ob- scurely glandular-toothed on the margins, dark green above, pubes- mt above when young but soon glabrate, covered beneath with whitish silky pubescence consisting of short appressed hairs; catkins appearing with the leaves, 4-' 7 cm. long, slender and densely flowered, sessile or igen on short leafy peduncles; capsules 4-6 mm. long, silky-pubescent, sessile or nearly so; bracts brownish, about 1 mm. lone, thinly Baye styles 0.5-0.7 mm. long; stamen 1; filament glab- e Mostly restricted to the coastal areas of so! uthern and southeastern aska and southward (Hultén, 1942: 551-2; 1949: 1733). However, Hultén cites a record of it at Lake Lindemann, and it should be lool for a. the Teslin, hag ey Tacay +g lake regions. It is LITERATURE CITED ANpERSSON, N. J. 1868. Salicineae. In De Candolle, Prod. Syst. Nat. Regn. Veg. 16°: 191-331. BALL, CARLETON R. 19: 921a. Undescribed willows of the Section Cor- datae. Bot. Gaz. Pa: 426-37. __. 1921b, Notes on willows of Sections Pentandrae and Nigrae. Bot. Gaz. 72: 220-36. __, 1923. The willows of the Pacific States. Repr. from from Abrams, L. An illustrated flora of the Pacific re Washington, Oregon, ia. 1: iT. ——. 1926. Can: dian willows of the sections Pentandrae, Nigrae and bae. Can. Field-Nat. 40: 145-152, 171-175. ——. 1930. In Cooper, W. S. The seed-plants and ferns of the Glacier Bay National Monument, Alaska. Bull. Torr. Bot. Cl. 5 : 327-38. —. 1934, New or little known West American willows. tae Calif. ——. 1935. New varieties of western willows. Proc. Nat. Acad. Sci. 181-86. 94 HUGH M. RAUP ——. 1938. New varieties and combinations in Salix. Jour. Wash. sr Sci. 28: 443-52. ——. 1942. Far western novelties in Salix. Madrofio 6: 227-39. ——. 1948. Salix petiolaris J. E. Smith: American, not British. Bull. —. 1950a. Willow hybrids: Salix hebecarpa Fernald and S. simu- 52: 8-1 . 1950b. A review of Salix anglorum and Salix petrophila. Am. Midl. Nat. 43: 224-241. ——. 195la. New combinations in Salix (Sections Pellitae and Phyli- Reg eae Am. Midl. Nat. 45: a "8 Bee . 1951b. In Spetzman, L. A. geography and ecology of the eareae slope of Alaska. =n M.S. thesis, Univ. of Min ——. 1951e. In Jordal, L. H. A floristic and phytogeographic summary of the southern slopes of the Brooks Range, Alaska. Un- published Ph.D. thesis, Univ. of Michigan. 1952a. Salix. In Gleason, H. A. The new Britton and Brown illostrated fare of the northeastern United States and adjacent Canada. ——. 1952b. ie Davis, R. J. Flora of Idaho, 216-35. Dubuque, Iowa. BreEITUNG, A. J. 1947. Catalogue of the Ligeoos plants of central eastern Saskatchewan. Can. Field-Nat. Copy, W.J. 1956. New plant records for pea ‘Aitenin and south- ern Mackenzie District. Can, Field-Nat. 70: 101-30. CoviLLe, F. V. 1901. The willows of Alaska. Proc. Wash. Acad. Sci. 3: 297-362. Dutitty, A., E. Lepage, & M. Duman. 1953. Contribution a la flore du bassin’ de la Bute’ @Ungava. Contr. Arct. Inst. Cathol. Univ. Amer. 4: 1-104. ‘ALD, M. L. 1926. Two summers of botanizing in Newfoundland. Salix ¢ cordifolia Pursh. Rhod. 28: 181-88. ——. 1946. Technical studies on North American Plants. Rhod. 48: 27-38, 41-9. —— 1950. Gray’s manual of botany. Eighth ed.: 487-519. FRANKLIN, JOHN. 1823. Narrative of a journey to the shores of the Polar Sea in the oly 1819, 20, 21, and 22, with an Appendix on various subjects relating to science and natural history. London. —— 1828. put of a second expedition to the shores of the Plax Sea in the years 1825, ot and 1827, including an account of a —~— ofa detachment 0 the eastward by John Richardson. GrO@NTVED, JoHS. 1936. Vascular plants from arctic North America the Fifth Thule Expedition, 1921-24. Rept. 5th Thule Exped. 2: 32-35. Copenhagen. Hooker, W. J. 1829-1840. Flora Boreali-Americana. London. pees Eric. 1942. Flora of Alaska and Yukon. Lunds Universi- ift Avd. 2, Bd. 39. No. 1, Part 3: 4 — iy rhe Flora of Alaska and Yukon, Lunds Universitets Arsskrift Avd. 2, Bd. 39. No. 1, Part 10: 1732-3. JorpaL, L. H. 1951. A floristic and phytogeographic summary of the of the Brooks Range, Alaska. Unpublished Ph.D. THE WILLOWS OF BOREAL WESTERN AMERICA 95 Moss, E. H. 1953. For jes in rthwest Alberta. Can. Jour. Bot. 31: 212. =f 448-70. POLUNIN, N. ge Botany of the Canadian ingest i Arctic, Part 1. Ed Nai . Bull. 92: 1-408. POoRSILD, ALE. 1943. Materials for a flora of the cautiental North- west Territories of Canada. Sargentia 4: 1945. The alpine flora of the me elope of the Mackenzie Moun- tains, Northwest Territories. Nat. . Can. Bull, 101: 1-35. 1951. Botany of terial poh adjacent to the Canol Road. lee Mus. Can. Bull. 121: 1-400. —— 1955. The vascular plants of the bess Canadian Arctic Archipelago. Nat. Mus. Can. Bull. 135: 1-226. 1957. uate flora of the Canadian Arctic Archipelago. Nat. Mus. Can. Bull. : 1-209. Pursu, F. 1814. ra Americae Septentrionalis. I: 1-358, II: 359- 751. London. Ravp, HucH M. 1931. Salix glauca L. and its allies in the Athabasca —Great Slave Lake region. Rhod. 33: 241-44. 1934. Phytogeographic studies in the Peace and upper Liard River regions, Canada, with a catalogue of the vascular plants. Contr. Arn. Arb. 6: 1-230. 1935. Botanical investigations in Wood Buffalo Park. Nat. Mus. Can. Bull. 74: 936. Phytogeographic studies in the Athabaska — Great Slave Lake region, I. Catalogue of the vascular plants. Jour. Arn. Arb. 315. —— 1942. Additions to a catalogue of the vascular plants oe "ee eace and upper Liard River regions. Jour. Arn. Arb. 23: 1 1943. The willows of the Hudson Bay region and the pose Peninsula. Sargentia 4: 81-127. y Ei e botany of southwestern Mackenzie. Sargentia 6: 1- Sc NEIDER, Camitio. 1918-21, Notes on American willows. I. Bot. Gaz. 66: 117-142 (1918), Il. ibid., 66: 318-353 (1918), III. ibid., : 27-64 (1919), IV. ibid., 67: 309-346 (1919), V. Jour. Arn. Arb. 1: 1-32 (1919), Ne ibid., 1: 67-97 (1919), VII. ibid., 1: 147- 171 (1920), VIII. ibid., 211 “32 ( 1920), IX. ibid., 2: 1-25 (1 920), X. ibid., 2: 65-90 ( aa, XI. eh 2: 185-204 ( (1921), XIl. ibid., (1921). a H. J. 1957. The Flora of Manitoba. Nat. Mus. Can. Bull. 140: glee - the arctic An and J. B. Tarleton. Bull. N. Y. Bot. G: 149-87. : 1922. Flora of ‘the Rocky Mountains eal at plains. 2nd ~ ‘Bdition 1-1144. New York. ; —— 1932. Flora of the prairies and plains of central North America. 1-969. New Yor! SPORANGIA OF THE FERN GENERA ALLIED WITH POLYPODIUM AND VITTARIA? KENNETH A. WILSON The structure of the mature sporangium, particularly the nature and orientation of the annulus, has frequently been illustrated and has long been used as the principal taxono- mic character of the families of the ferns (e.g., Lindley, 1830; Mohl, 1845; Mettenius, 1856; Sadebeck, 1898). The status of most of the families has remained relatively un- changed since the middle of the nineteenth century, an there appears to be general agreement about their distinct- ness and composition. With the appearance of Bower’s series of morphological papers and, ultimately, the publication of the three volumes of “The Ferns” (1923, 1926, 1928), the “family” Polypo- diaceae was interpreted as being a polyphyletic taxon com- posed of heterogeneous groups. Bower’s phylogenetic ap- proach to the problem of fern classification and the evidence showing the diverse origins of the groups in the Polypodia- ceae served as an impetus for a complete re-evaluation of this “family.” This led to the appearance of four systems of classification of the ferns within a period of eleven years. They began with Christensen’s scheme in the “Manual of Pteridology” (1938), which was fairly conservative. The authors of the other, more radical, systems are R. C. Ching (1940), R. E. Holttum (1947, 1949), and E. B. Copeland (1947). (For an outline of the four systems of fern class- ification, see A. G. Stokey, 1951.) The confusion which now exists in the classification of the Filicales and more specifically in the large assembly of the Polypodiaceae may perhaps, in spite of cries of dis- couragement, prové to be valuable in itself. Comparatively little is known about the biology of the ferns. Our knowl- edge of their morphology is based to a large extent on Bower’s works. Little new morphological work has been published since the 1930’s, with the exception of Wagner’s comprehensive study of Diellia (1952a). The present status of fern taxonomy demands that new research in this group be directed towards a more thorough understanding of the ferns themselves, as has been pointed out by Holttum 1Part of a dissertation submitted in partial fulfillment of the requirements for the of Doctor of Philosophy in the University of Michi, 98 KENNETH A. WILSON (1954) and Wagner (1954a). Only with the accumulation of new evidence can a truly phylogenetic system of classifi- cation be approached. Only a few significant strides have been made in this di- rection. Among the contributions are the outstanding works of Manton (1950 et seq.) on the cytology of ferns, and of Stokey (1951 et al.) and Stokey and Atkinson (1952 et seq.) on the gametophyte generation. Morphological, anatomical, and cytological evidence has been used by Wagner (1954b) in unraveling the interrelationships of the Appalachian aspleniums. Haider (1954) has made a valuable contribu- tion to filling the gap in our knowledge of the morphology of the mature leptosporangium. The present study is limited to a consideration of the sporangium of genera of the Polypodiaceae and Vittariaceae as defined by Copeland in 1947. The systematic treatments of the plants that make up this assemblage vary consider- ably. Christensen (1938) placed these and a number of other groups in the Polypodiaceae, and treated these as the Beamline Vitteicoud Be a a Paliorioul ae, 14 » an 1 eae, (which he said may be better dealt with as families). Ching (1949) considered them as comprising eight families: Antrophyaceae, Vittariaceae, Loxogrammaceae, Cheiropleu- riaceae, Dipteridaceae, Platyceriaceae, Polypodiaceae, and Grammitaceae (sic). Holttum (1947) segregated them into the Polypodiaceae and Grammitidaceae and placed the Vit- tarioideae in the Adiantaceae. Copeland (1951), in his grapk G itis, apparently on the additional strength of unpublished evidence from the gametophytes, recognized the Grammitidaceae and stated that “there is no reasonable doubt that the group should have family status.” SPORANGIA OF FERN GENERA 99 istopteris, and Anetium are all like that of Vittaria. Cope- land (1947) listed these same sporangial characters in his description of the Vittariaceae. Goebel (1924) placed great emphasis on the structure of the sporangium in determining relationships in the vittari- oid complex. He reported that the sporangium of Pleuro- gramme (= Cochlidium) is so distinct that it can be immediately distinguished from those of the Vittariaceae. After an investigation of the sporangia of Xiphopteris and Ad horus (= Amphoradenium) he concluded that, on the strength of the sporangial morphology in addition to other evidence, Plewrogramme should be removed from the Vittariaceae and placed with Xiphopteris and Adenophorus. One of the few other ref to the sp i of a member of the Grammitidaceae was made by Copeland (1951), who described the sporangial stalk of Grammitis as being of “mostly a single row of cells.” Haider (1954), in his comparative survey of the mature sporangia of various fern families, illustrated the capsules of only nine members of the Polypodiaceae. He described the sporangium of the Polypodiaceae (sensu Copeland, 1947) as having two stomial cells with a several-celled epi- and hypostomium. Of most significance, an invagination occurs between the stomial cells; the epi-and hypostomial cells are swollen. (See, also, Goebel, 1924.) The sporangial stalk received only cursory mention in Haider’s paper. His results show that the sporangia of Dipteris and Cheiropleu- ria are distinctly different from those of the other so-called Polypodiaceae, whereas those of Platycerium agree with them in all respects. The structure of the mature sporangium has been of value in studies of ferns in families not included in the scope of this paper. For instance, Wagner ( 1952a), after compari- son of davallioid, lind id, and asplenioid sporangia, con- cluded that those of Diellia closely resemble those of the Aspleniaceae. Schnarf (1904) had earlier pointed out the differences of the sporangia of Asplenium, Phyllitis, and Ceterach from those of Athyrium and Diplazium. One of the primary bases for the establishment of the family Negrip- teridaceae by Pichi-Sermolli (1946) was the structure of the sporangium. : Haider (1954) pointed out several interesting results of his comparative study. He wrote that the sporangia of the Pteridaceae (excluding the Dicksoniaceae) fall into two groups corresponding to Bower’s Gymnogrammoid and 100 KENNETH A. WILSON Pteroid ferns. The sporangia of the genera of the Daval- liaceae agree closely with each other, while those of the gen- era of the Blechnaceae which he examined differ consider- ably one from the other. Considerably less work has been done on the nature and the systematic value of paraphyses in comparative studies than on the sporangia. A valuable contribution to our knowledge of paraphyses was made by Pirard in 1947. Fifty-one different species of ferns were examined by her, and she reported that most of the structures interpreted by various authors as “transformed sporangia” have nothing to do with the spore-producing organs. However, Vittaria scolopendrina was shown to possess secretory structures which are truly transformed sporangia. Of value also in Pirard’s paper is a systematic listing of the presence of paraphyses in the ferns as indicated in the literature by various authors. The value of paraphyses in systematic studies and as a guide to fern phylogeny is not yet known. The Hawaiian relatives of the genus Ctenopteris have been segregated by Copeland (1947) as the genus Amphoradenium on the basis of the presence of clavate, glandular paraphyses within their sori. A study has been made of the paraphyses of Polypodium virginianum by Martens (1943, 1949) and Martens and Pirard (1943). Simple-stalked or branching giands were found in P. virginianum, and their presence was said to furnish a clear-cut distinction between that species and P. vulgare. Further studies indicated that these glands were in reality aborted or transformed sporangia (Martens and Pirard, 1943). An unusual situation is presented by Copeland (1947), who, on the one hand, recognized the genus Amphora- denium on the single morphological basis of the paraphyses, yet, on the other hand, included with the genus Polypo- dium species with (1) no paraphyses, ( 2) filamentous para- physes, (3) clathrate peltate paraphyses, and (4) stellate paraphyses. ACKNOWLEDGEMENTS During sp course of this investigation I was greatly aided by the Rogers McVaugh have: read the manuscript and offered vatuable rs ; I am indebted to them for their efforts in my behalf. Dr. SPORANGIA OF FERN GENERA 101 A. Howard for making this expedition possible. I am indebted also to Mr. George R. Proctor and Mr. C. B. Lewis, of the Institute of manuscript. TERMINOLOGY Much inconsistency has prevailed throughout the years with respect to the terminology used to designate the vari- ous parts of the sporangium. Haider (1954) reviewed the of the loosely used and misused terms, and, in addition, in- troduced a few valuable new ones. (See also Renner, 1955.) The annulus may be complete (i.e., a complete ring) and have all of its cells thickened, as in Gleichenia, or it may be complete but only partially indurated so that a few of its cells are thin-walled, as in Loxsoma. (Cf. Bower, 1923.) In either case, the term “annulus” refers to all of the cells in the “ring,” whether or not they are indurated. The term “ring” has, in fact, frequently been used as a synonym for i di i has an incom- thicker walls than those above or below it, although never as thick as the cells of i Vittaria, only two the capsule by separating : 1 tinguished from the other stomial cells as the “separation cells” (Haider’s “Trenn-” or “Saumzellen”). The cells low the stomium are referred to as the “hypostomium,” while the thin-walled cells between the bow and the stomium may be called the “epistomium.” 102 KENNETH A. WILSON The two sides of the sporangium are distinctly different in their cellular construction, and a series of terms has been used to differentiate them. Haider equated the terms in this manner: “Stielseite=proximale (basale)—infraanu- lare=bisuturale, | Deckseite=distale (peripherische) = supraanulare—unisuturale Seite.” I question Haider’s choice of “inf J supraanu lare” in preference to others. He does not explain how one can distinguish the “upper” and “lower” sides of the sporan- zium in the Polypodiaceae. The same problem would arise were we to select “stalk side—cover side.” “Bisutural— unisutural” are also confusing terms and do not correctly portray the real condition in the sporangium. We are then left with “proximal” and “distal.’ These terms appear to be the most suitable. The proximal face of the sporangium includes the entire upper portion of the first capsular seg- ment, while the distal face is composed of the second and third segments (Cf. Wilson 1958a and 1958b). Even these terms may be criticized on the same basis as the others, but I hesitate to suggest any new terms at this time. A really satisfactory designation for the capsular faces cannot be proposed until our knowledge of the morphology of the spor- angia of the so-called “primitive leptosporangiate ferns” has been extended and correlated with facts concerning the sporangia of higher ferns. MATERIAL AND METHODS This comparative study of the mature sporangium is based on an examination of fifty-eight species of ferns from forty-nine different genera of the Polypodiaceae and Vittari- aceae as defined by Copeland. His (1947) treatment of the genera of ferns is the most comprehensive of the recent re- visions, and therefore Copeland is followed as a matter of convenience. The number of genera included in my study totals approximately 65 percent of all those included by him in the above-named families. Because of the intimate asso- ciation of ph ith sp gia, these also have been included. It is to be regretted that it was not possible to SPORANGIA OF FERN GENERA 103 The sporangia of all but two of the species examined are illustrated. A careful search was made for the presence of paraphyses in each species and, when present, these have also been illustrated. It may be significant to note that para- physes have been found in many species in which they had not been reported, or were previously reported as being absent. Except in a few cases where preserved material was available, all studies were made from herbarium specimens. Since, in drying, the sporangia shrink and become distorted it was not possible to study them directly from the her- barium sheets. However, it was found that by boiling the sori for about a minute in 5 percent sodium hydroxide and then carefully dissecting out the sporangia the cells of the stalk and capsule became clearly visible. In order that my identifications may be verified, speci- mens of all material used are available at the Herbarium of the University of Michigan, or, in rare circumstances, in other indicated herbaria. All illustrations are camera lucida drawings made from temporary water mounts of the sporangia. For each species both the proximal and distal face of the sporangium are illus- trated. The stalk, because of its extreme length, is seldom shown in its entirety. When it is important, however, the stalks are fully illustrated. In other cases, the following system has been followed: the upper portion of two-rowed stalks is generally illustrated to include the base of the third row beneath the capsule. One-rowed stalks are shown from base, et cetera. Occasionally it has been difficult to illus- trate the stalk correctly because of distortions in the mater- ial. However, when this was so, a correct illustration of the stalk may always be found in one of the two views of the sporangium. : With a knowledge of the ontogeny of the sporangium, each of the original sporangial segments may be distin- guished in the illustrations (cf. Wilson, 1958a and 1958b). In view of the potential significance of the sporangium in comparative morphology it is felt that without an accurate iliustration of porangial face no valid conclusions can be drawn. This, unfortunately, is a shortcoming of most 104 KENNETH A. WILSON illustrations of the sporangium. Haider, who criticized in- accuracies of earlier illustrations, himself fell short in fig- uring only one side and in incorrectly picturing the cell arrangement of many sporangial faces. MORPHOLOGICAL OBSERVATIONS There appears to be little constancy in the size of the sporangia. Differences in the dimensions of the capsule oc- cur between genus and genus, but there seems to be no significance in this variation. Haider (1954) has pointed out that sporangia may vary in their size and shape within a single sorus. Of the genera included in this study, Anar- thropteris (Pl. 5, figs. J, and J;) and Loxogramme (PI. 5, figs. E-I) possess the largest capsules, which measure up to 300 microns in diameter. However, within the genus Loxo- found. L l lata (Pl. 5, figs. I, and I.) and L. chinensis (Pl. 5, figs. G, and G,) have large capsules with a diameter of 275 to 300 microns, while those of L. brooksii (Pl. 5, figs. H, and H,) range from 200 to 250 microns. Grammitis (PI. 4, figs. A-C) , Cochlidium (PI. 4, figs. D, and D.), and Hecistopteris (Pl. 5, figs. C, and C,) are among those genera which have small capsules which measure only 100 to 150 microns in diameter. Most of the other genera studied have capsules with an average diameter of 250 mic- rons. In shape, the capsule of all genera except those of the Vittariaceae is ellipsoidal or biconvex. Those of the Vittari- a@ceae are spherical. In some genera such as Rip gate (Pl, 2, figs. C, and C.), Neocheiropteris (PI. 2, figs. F; and F,), and Prosaptia (Pl. 4, figs. I and J) the capsule is greatly elongated, while in the sporangia of other genera the length of the sporangial face is only slightly greater than the width. Haider described the length-width ratio of the sporangium of Pessopteris as being 3:2; this agrees with my measurements. THE SPORANGIAL FACES The proximal and distal faces of the sporangium may be easily recognized. The proximal face may be distinguished by the characteristic four cells in the upper portion of Seg- ment I (e.g, Pl. 1, fig. D,). In addition, only one of the rows of the stalk leads into face. On the other hand, two of the rows of the stalk lead into the distal face, and by follow- SPORANGIA OF FERN GENERA 105 ing the line between the two rows Segments II and III may be easily located. Except for Dipteris and Cheiropleuria the cell arrange- ment in the sporangial faces of the genera studied is essen- tially similar. Some genera have a greater number of cells contributing to the sporangial faces, but their arrangement reflects the ontogeny of the sp i (eg., A th us: PI. 5, figs. D; and D,). There appears to be no reason to doubt that the development of these capsules follows the same pattern of those of Phlebodium (Wilson, 1958a), Xi- phopteris, and Pyrrosia (Wilson, 1958b). The great simil- arity of the cell arrang t in the sp ial faces in these genera indicates that this is so. Dipteris (Pl. 1, figs. A: and A,) and Cheiropleuria (PCS figs. B, and B;) have a cell arrang t in the sp ial faces that distinguishes them not only from each other but also from all other sporangia studied. It is not possible for me to | logize the sp ial faces of these two genera with those of the other genera investigated. There is no doubt that their ontogeny is different. Bower (1915) sug- gested that they develop from the initial by an opposite, two-rowed segmentation, but he did not describe the devel- opment of the capsule. e diff of the sporangial faces of the two genera are striking, but, for a clearer understanding of their structure, the ontogeny of their spor- angia should be studied in detail. A few genera have sporangia which bear hairs arising from the cells of the sp gial faces. Polypodium plu- mula (Pl. 1, figs. E, and E.) produces short hairs which arise from the cells of the distal face. A few sporangia of this species are naked and others may have up to five hairs, but more quently each sporangi bears only two or The hairs on the sporangia of Pessopteris have been fre- quently described. The two extremes have been fi here: one sporangium with thirteen hairs (PI. 2, fig. C:) The sporangium of Ctenopteris exornans bears an ex- tremely long seta (PI. 4, fig. H.). Occasional sporangia may be found to be naked; less frequently they bear two setae. THE ANNULUS Dipteris (Pl. 1, figs. A, and A,) and Cheiropleuria (Pl. 1, figs. B, and B;) are the only two genera which have an 106 KENNETH A. WILSON oblique and complete annulus. All of the other genera studied have a vertical ring which is interrupted by the stalk. Furthermore, the sporangia of Dipteris do not have a well-defined stomial region: almost all cells of the ring are indurated, although the cells opposite the stalk are usually thinner-walled. Copeland (1947) described the stomium of Dipteris as “ill developed.” Cheiropleuria, on the other hand, has a clearly differentiated stomial region. The Bow Although the number of bow cells varies within a single species, there is a general tendency for this variation to fall within a certain range, and usually one particular count oc- curs more frequently than any other. Wagner (1952a) re- ported the range and the average of the number of bow cells in the species he examined. Haider (1954), however, indicated the range and the most frequently occurring count of the cells rather than the average. Haider’s method of recording the counts seems to be the better one ; even though, except in unusual circumstances, the most frequently occur- ring count would probably not vary greatly from the mean. Table I presents the range and most frequent count (italics) of bow cells in ten sporangia of most of the species exam- ined in this study. Haider counted the bow cells of seven species of the Poly- podiaceae, including Platycerium bifureatum and Cyclo- phorus [Pyrrosia] winkleri, and concluded that “die Zahl der BZ [Bogenzellen] bei den Polypodium-Arten nahezu konstant 14 betrigt.” As may be seen from Table I, there is a tendency for most species of the Polypodiaceae to have about fourteen bow cells, but there is no suggestion of con- stancy, even in the genus Polypodium. The greatest number of bow cells was found in the spor- angia of Cheiropleuria, in which the count of nineteen was found to occur as frequently as twenty-four. The sporangia with the smallest number of bow cells were those of Gram- mitis graminea; in this species the number ranged from seven to ten, with ten being the most common. _ Colysis elliptica shows an unusual condition in the indura- tion of the cells of the annulus. Frequently one of the cells of the hypostomium may be found to have its walls thick- ened in the same manner as the bow cells (PL. 3, fig. E,). : cells become indur: ( SPORANGIA OF FERN GENERA 107 TABLE I COMPARISON OF NUMBER OF BOW CELLS* Dipteris conjugata Cheiropleuria bicuspis Platycerium andinum ium plumula Sey RR Wes ifoli Thylacopteris jis Dictymia atte! Pleopeltis ehunbergiana aes M. piloselloid Solenopteris bifrons Marginariopsis wiesbaurii Campyloneurum phyllitidis Pessopteris crassifolia i reum Eschatogramme furcata Paltonium lanceolatum Pp bull * vik I Weatherbya accedens — edie spicata la Sona piloselloides Microsorium pun tum M. scolo} ia Leptochilus axillaris Colysis elliptica Dendroglossa —— ans Ctenopteris exornans Prosaptia contigua brooksii figures represent sporangium for each species, while the figure or 18 — 19 — 24 — 27 17 — 18 — — 20 4— 15 12 — 15 — 17 — 19 12 12 14 — 16 — 21 13 — 1h 12 — 13 — 14 — 16 14 — 15 — 16 18 — 44 14 — 15 14 — 15 18 — 14 — 17 13 12 — 13 — 15 13 — 14 14 — 15 14 —17 14 14 — 15 17 — 18 — 23 17 — 18 — 19 14 — 15 14 i —4—15 14 12 — 14 11 — 13 14 — 15 7— 8— 9 9 — 10 — 13 13 — 14 9 — 10 12 — 18 8 — 9— 10 12 — 13 13 —1 1h — 15 — 16 — 17 11 — 1 13 — 14 the range in the number of bow cells per or figures in italies indicate the most 108 KENNETH A. WILSON A 1 is di pteri 14 — 15 Antrophyum reticulatum 13 — 144 — 15 Anetium citrifolium 10 — 11 Polytaenium lineatum 12 — 14 — 15 Scoliosorus ensiformis 13 — 15 — 16 Hecistopteris pumila 17 — 20 — 23 Vittaria remota 17 — 20 A i ifoli 13 — 14 Vaginularia paradoxa 17 — 18 — 19 The Stomial Region The greatest degree of variation in the structure of the capsule occurs in the stomial region. In Dipteris (Pl. 1, figs. A, and A,) the stomial region is not differentiated from e bow, while Cheiropleuria (Pl. 1, figs. B, and B,;) pos- sesses a well-defined stomial region with three stomial cells and usually three epistomial and three hypostomial cells. e sporangia of all species of the Polypodiaceae and Grammitidaceae examined are remarkably similar in the structure of the stomial region. All have two stomial cells clearly separated from the bow by thin-walled epistomial cells and with a well-defined hypostomium. The epistomium is generally three-celled, although in some cases it was found to be two-celled (e.g., Polypodium plumula: Pl. 1, fig. E:), or four-celled (e.g., Polypodium vulgare: Pl. 1, fig. D,). The epistomium of Platycerium (Pl. 1, figs. C, and C2) typically consists of four cells, while that of the species of Loxogramme (P1. 5, figs. F-I) ranged from three cells (Pl. 5, fig. H,) to as many as six cells (PI. 5, figs. E, and I,). The stomium of all sporangia examined in these two famil- ies was seen to have the characteristic invagination be- tween the stomial cells described by Haider. Unfortunately, not all illustrations show this condition since many sporan- gia were drawn after the separation of the stomial cells. Nevertheless, examination of other sporangia showed this invagination clearly, and it is evident in the sporangia well before the cells of the bow become thickened. This condi- tion may be clearly seen in the illustration of the proximal face of Belvisia (PI. 2, fig. I,). Other figures show the stom- ium from slightly different angles which frequently obscures this feature. In sharp contrast to the stomial region of the above fam- ilies, that of the members of the Vittariaceae has a four- celled stomium (e.g., Scoliosorus: PI. 4, figs. N, and N.), an epistomium of typically two cells, and a one- to three-celled hypostomium. Only in rare cases are there only three stom- SPORANGIA OF FERN GENERA 109 ial cells present (e.g., Vaginularia: PI. 5, fig. B.). There is no invagination to be found between any of the stomial cells of this group. THE SPORANGIAL STALK The sporangial stalk was found to be composed of one, two, three, or four rows of cells. The stalks of Dipteris (Pl. 1, figs. A, and A,) and Cheiropleuria (Pl. 1, fig. B, and B,) are four-rowed, but they are not identical to each other. That of Dipteris is elongated and is formed by four regular rows of cells. (Only two of the rows may be seen in each view of the sporangium.) Cheiropleuria, however, has a more massive stalk which becomes constricted at the base of the capsule. The arrangement of cells does not always appear to be as regular as it is in Dipteris. The sporangial ontogeny of these two genera is in need of investigation, since earlier reports are questionable. Three-rowed stalks (not figured) may be found in Platy- cerium, Marginariopsis, and Pyrrosia (Wilson 1958b), but they are rare. Generally their sporangial stalks are two- rowed except at the base of the capsule where they become three-rowed rowed. Two-rowed stalks are frequent and have been observed in all members of the Polypodiaceae sensu stricto. An un- usual condition is seen in the stalk of Dictymia (PI. 1, figs. H, and H.), which has a single small cell at the base of the stalk of Prosaptia linearis (Pl. 4, figs. J, and J.) is one- rowed for most of its length and does not differ from the others in this family. ly the one-rowed portion consists of two cells (PI. 5, fig. Fy): The stalks of L. brooksii (Pl. 5, figs. H, and H,) and par- ticularly of L. lanceolata (Pl. 5, figs. 1, and I,) are very i . That of L. lanceolata may be two-rowed at the base, but the rows of cells do not conform to the sporangial 110 KENNETH A, WILSON gramme lanceolata and Anarthropteris dictyopteris are the The sporangia of all species of Vittariaceae agree in having a sporangial stalk that is one-rowed at the base. This conforms with Bower’s (1928) and Copeland’s (1947) observations. PARAPHYSES Many of the species of ferns examined bore paraphyses. e most ly t 1 type was a simple hair composed of two to six cells. This type is found in Eschato- gramme (PI. 2, fig. D.), Paltonium (PI. 2, fig. E.), Lepto- chilus (Pl. 3, fig. D.), and many other genera (see especial- ly Plate 3). The stalked clathrate scale was also frequently found (e.g., Pleopeltis: Pl. 1, fig. Iy; Marginariopsis: P|. 2, fig. A.). These clathrate paraphyses are either peltate (PI. 1, fig. I,) or laterally affixed (Pl. 2, fig. H.), and frequently both types are found in the same sorus (Pl. 2, figs. I, and I,). lat ted with the sporangia of Stellate h are Platycerium (PI. 1, fig. C2), Pteropsis (Pl. 3, fig. A;), and Pyrrosia yf iow hy while M. piloselloides has clathrate, laterally affixed ones (not figured). Except for Anetiwm and Polytaenium, which bear no paraphyses, each genus of the Vittariaceae has a type en- tirely different from the others. two or three rows of cells (Pl. 5, fig. J,) ; Copeland (1947) suggests that they may be metamorphosed sporangia. Still other types are found in Dipteris (PI. 1, fig. A.), Cheiropleuria (Pl. 1, fig. B.), Goniophlebium (Pl. 1, fig. SPORANGIA OF FERN GENERA 111 F,), Solenopteris (Pl. 1, fig. J,), and Ctenopteris (PI. 4, fig. 4). Aborted, tannin-filled sporangia in various stages of development were observed in Polypodium plumula, Escha- togramme, Pseudodrynaria, Crypsinus, Selliguea, and Pros- aptia contigua. DISCUSSION OF THE SIGNIFICANCE OF SPORANGIAL MorPHOLOGY TO FERN TAXONOMY During the last several decades it has been the general practice to describe the sporangia of the so-called “advanced leptosporangiate ferns” as being simply of the “polypodia- ceous type.” With this, all sporangia which are stalked and possess an erect annulus were considered to be fundament- ally similar. From the facts presented in the previous sec- tion, it is shown that the parts of the sporangia of different fern groups are by no means identical in their structure. The morphology of the sporangium can be used as a valu- able guide to indicate generic interrelationships, and may serve in clarifying the status of problematic genera and species. The sporangia of the vittarioid genera stand out as the most distinct and easily recognizable group among those included in this study. These sporangia are characterized by having four stomial cells, a thin-walled epi- and hypos- tomium, and a one-rowed sporangial stalk. There has been very little disagreement with respect to the composition of the Vittariaceae. Ching (1940) was the only one who divided this assemblage into two families. This he did on the basis of the structure of the spores, but both types of spores he used as the basis of separation are of the number of bow cells in classification, since their num- ber appears to be so variable. In all other respects the sporangia of the genera of the Vittariaceae, sens. lat., are very similar to each other. guishing characteristic of the Vittariaceae, as pointed out by Stokey (1951), is in the gametophyte, which she said is “sharply distinct from that of any other 112 KENNETH A. WILSON known higher fern.” The morphological distinctness of the sporophyte was discussed by Goebel (1924). Manton and Sledge (1954) reported that the Vittariaceae might be placed with Adiantum, as was suggested by Holt- tum (1947), “without doing violence to the cytology.” However, very little is known about the sporangium of Adiantum. Haider (1954) illustrated the capsule of A. peruvianum and showed three stomial cells. (Unfortunately the stalk was not figured.) But three stomial cells are rare in vittarioid sporangia, and a single incomplete drawing is insufficient for adequate comparison. Surprisingly little is known about the gametophyte of Adiantum (Stokey, 1951). Much more work is needed in the pteroid ferns before we can arrive at any conclusion in this problem. Suffice it to say that Copeland’s treatment of the Vittariaceae is well supported by the evidence from the sporangium. The sporangia of Dipteris and Cheiropleuria do not show any indication that these genera are closely related or even remotely allied to the polypodioid ferns. A four-rowed con- dition of the stalks is not in itself evidence of relationship, particularly when these stalks differ from each other. The structure of the capsule of the one genus is different from that of the other. Haider (1954) has already stressed these differences. The removal of the two genera from the Poly- podiaceae, and their placement in two separate families as done by Ching, is now supported by the structure of the sporangium as demonstrated in the present investigation. Evidence from the gametophyte as reported by Stokey (1945) and Stokey and Atkinson (1945a) also justifies this treatment. Wagner (1952b) found that the combination of h: ters in the sporophyte of Cheiropleuria supports the establishment of the family Cheiropleuriaceae. Manton (1954) reported that the “n’”’ number of chromosomes is “ce, 33.” This is significantly different from that of the typical Polypodiaceae which ranges around thirty-seven. On the basis of this accumulated evidence, the recognition of the families Dipteridaceae and Cheiropleuriaceae is now strongly justified. Ching placed Platycerium in the isolated family Platy- ceriaceae. The sporangial structure does not, however, jus- tify this action. The sporangia of Platycerium do not differ in any fundamental respects from those of the other mem- bers of the Polypodiaceae. Haider has reported that there is “kein Zweifel, dass die Gattung mit Polypodium verwandt ist.” I can only emphasize his conclusion. Stokey and At- SPORANGIA OF FERN GENERA 113 kinson (1954b), in a study of the gametophytes of five spe- cies of Platycerium, also found no evidence to justify the family Platyceriaceae. Manton and Sledge (1954) have shown that cytologically this genus agrees with other mem- bers of the Polypodiaceae. Wagner (1952b) reported on other similarities of Platycerium to the polypodiaceous genera Pleopeltis and Pyrrosia. The evidence is unusually clear and detailed that Platycerium is rightly placed in the Polypodiaceae. Most recent authors agree that the grammitid ferns are distinct from the Polypodiaceae. Whereas the sporangia of the members of the two groups are remarkably similar in the structure of the capsule, the one-rowed stalk clearly distinguishes the Grammitidaceae from the Polypodiaceae. The sporangia of both families have a capsule with an erect annulus which is interrupted by the stalk. In both, the stom- ial region is clearly differentiated and possesses two stomial cells and a thin-walled epi- and hypostomium. Both have a deep invagination between the two stomial cells. However, the stalk of the Polypodiaceae is two-rowed at the base, while that of the Grammitidaceae is one-rowed. The problematic species Polypodium plumula, which shows some characters in common with the grammitid genus Ctenopteris, has a two-rowed stalk, and on this basis its inclusion in the Polypodiaceae is supported. The significance of the single cell at the base of the stalk of Dictymia is uncertain. This may be the result of the orientation of the first oblique wall in the sporangial initial which may not quite reach the level of the receptacular cells. Additional investigations may indicate that the typical two-rowed polypodiaceous stalk may also sometimes occur in this genus. Nevertheless, the presence of a single small cell at the base of the stalk is at present not considered to be significant in this genus. An investigation of the nature of the first division of the sporangial initial would help sot leaf texture.” The significance of this statement is not clear to me, and such an association has not been suggested by others. Moreover, Ching’s inclusion of Anarthropteris dic- 114 KENNETH A, WILSON tyopteris in this genus is not supported by evidence from the sporangium. The three-rowed stalk is rare in the Polypodiaceae and has been found only in a very few individual sporangia out of a two-rowed majority in three different species. The sporangia of Prosaptia contigua are peculiar in the structure of their stalk. They do not agree with the typical- ly grammitid stalk of P. linearis. This may be a result of abnormal develop t. The abund. of aborted spor- angia is the sori of P. contigua indicates this may be the case. The genus Lorogramme has turned out to be a problem- atic one from the standpoint of p gial structure. It has been placed in the Polypodiaceae by Holttum, in the Grammitidaceae by Copeland, and in its own family, Lozo- grammaceae, by Ching, who considered it closely related to the vittarioid ferns. Christensen (1938) treated it as “a distinct genus of very doubtful relationship, perhaps related to Grammitis, but in vegetative characters not unlike Hymenolepis [Belvisia] and Paltonium.” A widely diver- gent view has recently been expressed by Nayar (1955) who gested that Loxog be placed in the Pterid- aceae and idered a derivative of the “Syngrammoid” ferns. The structure of the p gia of Loxog ne does not agree entirely with either that of the Polypodiaceae or the Grammitidaceae. They differ from these families in having the basal one-rowed portion of the stalk composed of ene or two cells and having generally five instead of three epistomial cells. Their sporangial stalk is difficult to inter- pret, and a study of the sporangial ontogeny is needed to clarify its structure. The stalks of Loxogramme brooksii and L. lanceolata only add to the confusion. I would tenta- tively suggest that Loxogramme may best be treated as a distinct family, but a more satisfactory treatment must await a comprehensive study of the genus. e association of L with Syng is du- bious, at least from sporangial evidence. Nayar (1955) makes the following comparison: “The characteristic three- rowed stalk of sporangium, the average number of cells con- stituting the annulus [bow], and the stomium with pro- truding lip cells and separated from the stalk and annulus ick also i one-rowed, and the three-rowed portion subtending the cap- sule cannot validly serve as a basis for comparison since SPORANGIA OF FERN GENERA 115 this is characteristic of sporangia of the Polypodiaceae, Grammitidaceae, Vittariaceae, Aspleniaceae etc., and re- flects the ontogeny of the capsule (Wilson, 1958b). As has been shown, the number of bow cells is of questionable sig- nificance. Moreover, the sporangium of Syngramma has not been studied, nor have those of the other genera of the Pteridaceae.? The gametophyte of Loxogramme is imperfectly known. Manton (1953) reported the chromosome number of the genus, but this still Mhid not solve the problem of its relation- ship: cytological evidence indicates that it is not associated with the Vittariaceae, but places it in the polypodioid-gram- mitid complex; the chromosome numbers of the members of the Polypodiaceae and Grammitidaceae are not basically unlike. The sporangia of Copeland’s monotypic genus Anarthrop- teris present an equally perplexing problem. Copeland (1947) stated that “Anarthropteris seems most nearly re- lated to Loxogramme,” and placed it with the grammitid ferns. The sporangium is certainly not grammitid in its structure. The evidence of vertical intercalary walls during stalk development is unusual, although its significance is obscure. The primordial sesh indicates that the first division is most likely t teris (Wilson, 1958b). But again this is not conctiaiée evidence of re- lationship. The only resemblance of its sporangium to that of Loxogramme is in its stalk which, by a slight stretch of the imagination, might be compared to that of L. lanceolata. From the structure of the sporangium it appears that Anar- thropteris does not belong either in the Polypodiaceae or the Grammitidaceae. Its association with Loxogramme is questionable. The actual affinities of this little known fern cannot be determined without further investigation. On the basis of the structure of the paraphyses, there ap- pear to be three major groups of genera in the Polypodia- ceae. The first of these, pawns Sah and may bea hel and Pteropsis, has stellate para may be a close related group of genera. Wagner (1952b) discussed t similarity of Platycerium to Pyrrosia, and Copeland foc ?Since this paper was prepared, a . pebiention, by A. enema a chegeieggerd Kenntnis der einiger 2 359- 426. 1931) has come to my attention. Landmann deseribed the sporangia ee Syn- gramma as short-stalked possessing elo arrow, brown stomial more doubtful that Lozogramme and Syngramma are 116 KENNETH A. WILSON suggested that the affinity of Pteropsis is “manifestly to rosa.” Bower (1923) suggested that “it is but a step from... stellate hairs to a peltate scale with central attachment.” It seems more likely in my opinion, however, that the pel- tate paraphyses are not derived from the stellate ones. The development of the stellate paraphyses (Wilson, 1958b) seems to be quite different from that of the peltate scale as described by Bower. Pleopeltis, Marginariopsis, Lemma- phyllum, Neocheiropteris, Weatherbya, and Belvisia all have peltate, clathrate paraphyses. The last three genera also have clathrate paraphyses which are laterally affixed; these are also found in Microgramma. piloselloides. On this basis, these genera may represent, therefore, the second closely related group in the Polypodiaceae. Copeland (1947) also suggested this association. The genus Microgramma, however, has species with different types of paraphyses. Microgramma lycopodioides has branched filamentous paraphyses. This suggests either that the genus is in need of revision or, as is more likely, the paraphyses are not always adequate indicators of re- lationship. The third group might be composed of those genera having simple filamentous paraphyses. This would include Eschatogramme, Paltonium, Microsorium, Leptochilus, Colysis, Dendroglossa, Pseudodrynaria, 2 pigeon Sel- d P liguea, is, and Phlebodit semblage of genera is not considered to be a closely eters group by any author. Unbranched filamentous paraphyses may arise in many different ways, and without a precise knowledge of their origin they cannot point to interrelationships of spe- cies or genera. The value of paraphyses as indicators of fern phylogeny is not known. What do the widely different types of para- physes described in the species of Polypodium or Micro- gramma signify? Much more study is needed to solve this problem. Alston (1956) suggested that Stenochlaena belongs in the Polypodiaceae. Copeland ( 1947) placed it in the Blechna- ceae, while Holttum put it in the subfamily Pteridoideae of the Dennstaedtiaceae. Stokey and Atkinson (1952) found that its gametophyte has much in common with that of Blechnum. Manton and Sledge (1954) reported that the aoe = Stenochlaena does not conform very well — either Copeland’s or Holttum’s view. Cytological eviden SPORANGIA OF FERN GENERA 117 gives support to Alston’s placement of the genus in the Poly- podiaceae. However, Haider (1954) examined the sporan- gia and said that in the structure of the stomial region they are very similar to those of the Aspidiaceae. The problem of the affinity of Stenochlaena thus also remains unsolved. The sporangia of the rare Bornean fern Holttumiella flabellifolia are not known. Copeland reported that they are “apparently sessile.” On the basis of this description and of the findings of the present investigation, there is some doubt about its pl t in the Polypodii Our | ledge of the sp ium of the genera of other fern families is meager. It is evident that the structure of the sporangium can contribute a great deal to our under- standing of fern phylogeny. There is no doubt that the study of sporangial morphology can serve as important new evidence in fern taxonomy in a manner like that which has been demonstrated for the cytology and the gameto- phyte generation. SUMMARY The structure of the sporangia of fifty-eight species of ferns placed by Copeland in the Polypodiaceae and Vittari- aceae is discussed and compared. It is shown that the struc- ture of the sporangium can be used as a valuable guide in systematic studies of ferns. On the basis of sporangial morphology, the Vittariaceae has been shown to be a distinct family. The families Dip- teridaceae and Cheiropleuriaceae are justified by the spor- angial structure. There is no evidence to support the recog- nition of a family, Platyceriaceae. The two-rowed stalk of in a separate family. Both Loxogramme and Anarthrop- teris are in need of intensive investigation. Some probable t ic implicati h are discussed but V. of parap their signifi in phylogenetic studies remains obscure. LITERATURE CITED Aston, A. H. G. 1956. The subdivision of the Polypodiaceae. Taxon 5: 28-25. rey jowER, F. O. 1915. Studies in the phylogeny of the Filicales. V-. Cheiropleuria bicuspis (Bl.), Presl, and certain other related ferns. Ann. Bot. 29: 495-529. ss a 917. Studies in the phylogeny of the Filicales. VI. Ferns showing the “Acrostichoid” condition. Ann. Bot. 31: 1-29. COMPARISON OF SPORANGIAL STRUCTURE WITH PROPOSED TAXONOMIC SYSTEMS* SPORANGIA GENERA” CHRISTENSEN® CHING (1940) COPELAND HOLTTUM ASSEMBLAGE OF OTHER (1938) (1947) (1947, 1949) DISTINCTIVE CHARACTERS Stalk 4-rowed; an- — Dipteris Dip loid pterid Polypodi Polypodi Sol stem; spores el- nulus complete; sto- liptie; gaunetopyte with mium — undifferenti- thick midrib; n = Stalk 4-rowed; an- Cheiropleuria Polypodioideae Ch i P 1 Polypod Protostelic or solenostelic nulus complete; sto- em; spores tetrahedral; mium well defined. stipes nonarticulate; gameto- phyte narrowly cordate. Stalk 2-rowed; an- Platycerium Polypodioid Plat Polypodi Polypodi Dictyostelic stem; spores el- nulus interrupted; ee Polypodioideae Polypodi Polypodi Polypod liptie; tipes articulate; stomium 2-celled esate nel Polypodioideae Polypodiaceae — Polypodi. Polypodi sonlen mostly reinente! copte ge = Polypodiaceae _ ametophyte date; Piero Pol lioid Polypodi Pol Polypodiaceae ws, 37; leaf mainly Pedenatee eined, Microg na Polypodioid Poly padi Balspodi is _ Solenopteris Polypodiaceae — ae is s v- r I _ Campyl Polypodioid Polypod' Polypodi = Pessopteris _ . ly podi s lypodi — Phlebodium ose families which are in bold face type are those ise ewe which are feat favereal by the evidence at present. Genera not specifically mentioned “Those fi by an author are indicated by a dash, are Lars baalnead in this study and follow the treatment of Copeland (1947); synonyms and differing generic taxonomic treatments of the *The g other authors are not indica’ *Christensen treats site ‘tern groups as subfamilies of the Polypodiaceae in the very broad sense. 811 NOSTIM ‘V HIGNNAY TABLE II (continued) ASSEMBLAGE OF OTHER DISTINCTIVE CHARACTERS Dietyostelic stem; spores el- liptic; stipes _ articulate; scales clathrate; leaf reticu- Solenostelic Bosal spores ripes nonarti- SPORANGIA GENERA CHRISTENSEN CHING (1940) COPELAND (1988) (1947) (1947, 1949) Dias Palsnediota Polypodi Pol . ae eed Polypodioid Polypodi Pol _ Polypodioid Polypodi Pol Polypodi ihn Palypodiold Polsnodi E ; Polypodi Wesketoe _ _ P ee Belvisia Polypodioideae = Polyp Polypodi yrrosia —- Pol ioid Polypodi P Polypodi Pteropsis —_— Pol ioid Polypodi Pp Polypodi i Polys 7 I +: Pp Bolvondl Leptochilus Poly id Polypodi Polyp Polypodi 0 Pol. 4 Polypod ri Balenant Dendroglossa _ Pp = Pseudod ia Polypodioid Polypodi F = Drynaria _ Polypodioid Polypodi Polyp Polypodi I, ype PY P 1 At 33 P 1 Ai Pe p Pp 1 Al Crypsi _ —_ Polyp ae Selliguea —_ Polypodioid P i Polypodi Polypodi Auth ete | Rolppanioid Polypod Polypodi Polypodi Stalk 2-rowed; 1- Dictymia — Polypod Polypod _ celled at base; an- nulus interrupted; stomium 2-celled, late-veined. Stalk 1-rowed; an- Grammitis Polypodioid i Polypod 'G nulus interrupted; Cochlidium Pol dioid i Pol 1 'G iti stomium 2-celled, — Sel. l Polypodioid Polypod " G i “The family Grammitidaceae was recognized by Copeland in 1951 and these genera were placed in it. culate; pecien nonclathrate; VUENID Nudd dO VIONVUOdS 611 Taste IT (eontinued) GENERA CHRISTENSEN CHING (1940) COPELAND HOLTTUM ASSEMBLAGE OF OTHER SPORANGIA (1938) (1947) (1947, 1949) DISTINCTIVE CHARACTERS Xiphopteris _ — Polypodi r itid elongate; n Caly i Polypodioid i Polypodi : itid 36, ‘leat mainly free- Stenop Polypodioid Polypodi (wel fa ee os } ° is ae |p) a Hs Dilcnaal 1¢ va Pp * OVP + . sive Prosaptia —_ Polypodioid G Polypod Grammitidaceae Stalk 1 row of 2 Lowog Polypodioid 3 Polypodi Poll Di stem; spores cells; annulus inter- elliptic or peo: stipes rupted; stomium 2- ticulate scales clath- celled, fe n = 86. Stalk irregular, 2- or Anarthropteris _ Polypodi Polypodi re ae Dictyostelic stem es 8-rowed; annulus in- elliptic; stipes nenareculates terrupted; stomium leaf reticulate-veined, 2-celled, Stalk 1-rowed; an- Antrophyum Vittarioideae Antrophyaceae Vittariaceae Adiantaceae — Protostelic or siphonostelic nulus — interrupted; Polytaenium Vittarioideae Antrophyaceae _Vittariaceae Vittarioideae stem; spores _ elliy stomium 4-celled. Ananthocorus Vittarioideae A F Vittari _ tetrahedral; stipes nonarti- Anetium Vi ioid pl Vittariaceae culate; scales clathrate; Hecistopteris Vittarioideae Vittariaceae Vittariaceae Vittarfoidess eaineinahite elongate; n = Vaginularia — Vittarioideae Vittariaceae Vittariaceae 30; laminar spicular cells; Vittaria Vittarioideae Vittariaceae Vittariaceae Vittarioideae 4 mainly reticulate vein- Scoliosorus - — Vittariaceae OI NOSTIM *V HLANNGY SPORANGIA OF FERN GENERA 121 —. The Ferns. Vol. I. Analytical Examination of the Cri- teria of Preity ache 359pp. iver ay Press, Cambridge, England. —. 192 The Ferns. Vol. Il. The Eusporangiatae and other slaively primitive ferns. San University Press, Cambridge, ngial ——. 1928. e Ferns. Vol. III. The se ig ee Ferns. 306pp. University Press, Cambridge, Englan CuinG, R. C. 1940. On natural classification om the family “Polypo- diaceae.”
Gallus 51: 155-184.
Renner, G. 1955. Bau und Funcktion des Ringes am Celey eae
gium. Sitzs. Bayer. Akad. Wiss. math.-naturwiss. 1954
SADEBECK, R. _, 1898. shoei In Engler and Pun This
tiirlicl ili (4): 1-91.
ScunarF, K. 1904. mueaee: zur Kenntnis des Sporangienwand-
baues der Polypodiaceae und der Cyatheaceae und seiner system-
atischen Bedeut utung. Sitzs. Akad. Wiss. Wien, math.-naturwiss.
So A. G. 1945. The Gametophyte of Dipteris conjugata. Bot.
. "1951. The ibuti n by the gi phyte to classificati
of the h ferns, hol : 39-58.
Stoxey, A. G., pL. R. ATKrNSON. 1952. The pene of
Stenoehlaena abuts (Brum. - Bedd. ee
bicuspis (Bl. ) Preak
~ Piytomorthology 4: 192-201.
The seme pay of five species of Platycerium. Phyto-
Ea acre 4: 165-172.
Wacner, W. H. Jr. 1952a. The fern genius pega its structure,
affinities and taxonomy. Univ. Cal. Publ. Bot. 12.
——. 1952b. pores of foliar dichotomy in eine awe Amer. Jour.
Bot. 39: 578-5!
——. 1954a. The evidence used in recent yeas of the ferns.
Huititme Congr. Intern. Bot. Rapports & Communications.
2,4,5,6: 9-15.
. 1954b. re evolution in the Appalachian aspleniums.
a ypreeionss = 103-11;
A. 1055a. Ontogeny of the sporangium of Phlebodium
Poigpedtien) aureum. Amer. Jour. Bot. 45: 483-491.
——. 1958b. Ontogeny of the sporangia in Xiphopteris serrulata
and Pyrrosia nuda. Jour. Arnold Arb. 39: 478-493.
SPORANGIA OF FERN GENERA 123
ve I. Mature sporangia and paraphyses — Fig. A. Dipteris conjugate
(Philippines; Gates 7663). — Fig. B. Cheiropleuria bicuspis (Ryukyu Is.: Walker
& Tawada 6891). — Fig. C. Platycerium andinum (Peru: Klug 3872). — Fig. D.
olypodium vulgat (“in paeninsula Hel m icum.” Trela). Fig. E.
Polypodium plumula (Bermuda: Taylor 49-1163). — Fig. F. iophlebium persici-
foliu (Si : Bartlett 8470). — Fig. Th teris papil tra:
artlett 6753b). — Fig. H. Dictymia attenuata (Australia: Simonds 1895). — Fig. L.
Pleopeltis thunbergiana (Ha Topping 3712). —Fig. J. Solenopteris bi-
rons (Colombia: Klug 1838). — Fig. K. Microgramma lycopodioides (Cuba:
Jervis 1441B). In all eases the two faces of the sporangia are shown: paraphyses,
when present, are also shown.
124 KENNETH A. WILSON
—Fig. A s wiesbaurii
2637). — Fig. B. shana tod phyllitidis (U1 Mich. Bot.
“hs pteris crassifolia (Costa Rica: Scamman, March 1951).
eyig I Rbchitayrasins | younhs (Haiti: Ekman 4714). — Fig. E. Paltonium
lanceolatum (Dominica: Cooper II 92). —Fig. F. Neoc teris phyllomanes
ina: a Fig. G. Lemmaphyllum eieoshelion (China:
Lau 835). — Fig. . Weatherb ya accedens (Sumatra: Boeea 6968). — Fig. I. Bel-
visia spicata (Sumatra: Bartlett 8004).
PLATE Ir. Mature sporangia and parap . Marginaric
Skutch 2
SPORANGIA OF FERN GENERA 125
Piate Ill. Mature sporangia = paraphyses.
(Indo-China: Clemens 3333). B.
Copeland 1535). — Fig. c.
re Fig. E.
Dendroglossa cantoniensis ( ina:
(Univ. Mich. Bot. Gard. 19978; Koelz
‘ak c istian Col
lege 15108). ewe J. Lecanopteris carnosa (Sumatra
Fig. L. Arthromeris juglandifolia (India:
Selliguea feei (Sumatra: Bartlett 7819).
Duthie 6295).
126 KENNETH A. WILSON
Pate IV. Mature sporangia and paraphyses. — Fig. A. Grammitis graminea
(Jamaica: Wilson & Murray 558). — Fig. (Jamaicaé:
(Mexico: Matuda 2907).
SPORANGIA OF FERN GENERA 127
PLA’ Mature sporangia and paraphyses. — Fig. A. nama lineatum
(Jamaica: "Wilson & Webster 468). — Fig. B. pa: 5
Yuneker 9070). — Fig. C. Hecistopteris pumila (
D. Ananthacorus angustifolius (Briti: S gas
icifolic ‘sang 72). — Fig.
H. Loxogramme brooksii (euumicn: Bartlett — Fig. L
ceolata (Philippines: Topping 429). — Fig. J. Anartiropteria Paesears (New
Inverarity 1868).
THE SOMATIC CHROMOSOMES OF RUDBECKIA AND
RELATED GENERA OF THE COMPOSITAE’
ROBERT E. PERDUE, JR.*
INTRODUCTION
The research reported in this paper is the outgrowth of a
study that was directed primarily toward an understanding
of the taxonomy and relationships of species of the genus
Rudbecki ata on cl number and somatic chro-
mosome morphology were obtained which reflect upon the
relationships of species within the genus as well as upon re-
lationships between F kia and related genera. While
these data did not provide the primary basis for taxonomic
judgment, they aided considerably in guiding the taxonomic
approach and provided evidence which fully supports deci-
sions based upon conventional morphological study. Study
of the chromosomes of this genus indicated where the line
should be drawn between 2 subgeneric groups and where
the taxonomist should expect to find distinct morphological
differences.
The fulfillment of the objective of the investigation called
ideration of 3 additional genera, Dracopis, Ech-
bida as separate genera, in the most recent edition of Gray’s
Manual, Fernald (1950) regarded Dracopis as a section of
Rudbeckia. In the latest edition of Britton and Brown’s
Flora (Gleason, 1952), Cronquist treated Dracopis as a
distinct genus. Battaglia (1952) expressed the opinion,
based primarily on embryological data, that of these genera,
tRevised from a portion of a thesis submitted to the Department of Biology,
Harvard University, in partial fulfillment of the requirements for the degree of
of Philosophy. I am especially indebted to Dr. Reed C. Rollins under whose
mducted. A was: leted
direction this study col large portion of the work
during tenure as Associate Botanist, Texas Research Foundation, was supported
by a grant from the h Foundation, During the early stages of the research,
nald
used.
2U, S. Department of Agriculture, Agricultural Research Service, Crops Research
Division, Beltsville, Maryland.
130 ROBERT E. PERDUE, JR.
only Echinacea should be segregated from Rudbecki
Morphologically these 4 genera can readily be distinguished
rom one another and these distinctions are supported by
the cytological data presented in the present paper.
Rudbeckia is a genus of North America Compositae,
Tribe Heliantheae. The 15 species form 2 very natural sub-
generic groups, subg. Rudbeckia and subg. Macrocline,
whose limits are morphologically very well defined, though
a number of the characters which distinguish them are
comparatively obscure. The morphological basis for the dis-
tinction of the 2 subgenera is discussed in detail in another
paper (Perdue, 1958).
RPiudh Bia. RP.
77,
: includes 9 species, consisting
of a total of 22 taxa (Perdue, 1957 ). The species are very
distinct (morphological inter lation between species does
not occur), so distinct that interspecific hybridization would
not appear to be possible in most combinations even though
the species were to be brought into contact. Most of the
species of this subgenus occupy natural habitats in con-
junction with 1 or more other species. Extensive field and
herbarium studies have not revealed even the most remote
ggestion of interspecific hybridization. Essentially, subg.
Rudbeckia is a group of eastern North America species.
Only a single taxon, R. hirta var. pulcherrima, is distributed
generally beyond the eastern borders of the Great Plains.
This common weed is found from southern Canada, through
all the United States, into northern Mexico.
Rudbeckia subg. Mi line includes 6 species, encom-
passing a total of 16 taxa (Perdue, 1958). In contrast to
those of subg. Rudbeckia, the species of this group are very
closely related. Morphological intergradation is not common,
but this is apparently due to the restriction of the species to
geographic areas well separated from the ranges of their
close relatives. The occurrence of 2 species in the same
habitat or even in the same county is rare. Several of the
species of subg. Macrocline appear to be so closely related
that hybridization might be expected were they to be
brought togeth
er.
Although many species of Rudbeckia are well known,
judging from the arrangement of species in floristic manu-
als the existence of 2 well-defined subgeneric groups
species with very narrow, grass-like leaves that are unlike
those of any other species of the genus. The narrow-leaved
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 131
R. mohrii (subg. Macrocline) is distributed through a com-
paratively restricted area in western Florida and southern
Georgia. The narrow-leaved R. graminifolia (subg. Rud-
beckia) is a very narrow endemic restricted to a small area
of western Florida within the geographic area of R. mohrii.
Gray (1884) recognized the true relationships of most of
the species of Rudbeckia but placed R. mohrii and R. gram-
inifolia together and apart from their true relatives because
of their unique, narrow leaves. The understandable failure
to recognize the existence of the 2 subgeneric groups has
persisted into our most recent floristic manuals.
MATERIALS AND PROCEDURE
A majority of the plants examined cytologically were
grown in the greenhouse from seed. Other plants were col-
lected in nature, potted and intained in h
Each lot of seed or of living plants was assigned an acces-
sion number. The culture numbers used in Table I reflect
the accession numbers and in those cases in which 2 or
more plants were grown from a single accession, they are
distinguished by secondary numbers.
Considerable difficulty was experienced in germinating
seed of most species of the 4 genera. Seed of the annuals
(Rudbeckia hirta, R. mollis, R. triloba and Dracopis amp-
lexicaulis) germinated without treatment in 5 to 7 days
when placed on moist filter paper in a petri dish and main-
tained at room temperature. Seed of the other species fail-
ed to germinate after 2 weeks in petri dishes even after an
initial over-night soaking in water. Difficulty in germinating
seed of the perennial species appeared to be due to im-
permeability of the achene wall to moisture. When the
eg
During the early stages of the work, several attempts
were made to germinate seed on circles cut from ordinary
paper towels rather than on filter paper. In each instance
the cotyledons increased in size and turned green; the radi-
cle of the embryo began to develop but soon turned brown
and died. Apparently, a substance which acts as an inhibi-
tor of root growth was present in the paper toweling.
132 ROBERT E. PERDUE, JR.
About 5 to 7 days after the seed germinated, the young
plants were of sufficient size to be handled. Twenty-five or
more plants were transplanted from each petri dish to a
large pot. Several weeks later, selected plants were trans-
planted to small individual pots.
Although seed could be germinated at any time of the
year and at any stage after they matured, growth of young
or mature plants was negligible during the winter. Murneek
(1940) found that seedlings of a horticultural form of
Rudbeckia hirta (as R. bicolor var. superba) grow very
slowly during the fall and winter until about the middle of
March, when the photoperiod increases to about 12 hours.
At this time the leaves of the young rosette, extremely flat
during the winter, assume a more upright position and vig-
orous growth begins. These observations seem to apply to
all species of Rudbeckia that I have grown in the green-
house. Although this factor was not studied closely, it ap-
peared that the plants were essentially dormant, or at least
very sluggish in growth, until the photoperiod reached a
satisfactory length. Within a period of a few days after
factory for growth, leaves that had
been flat or even drooping somewhat, raised slightly, and
shortly new leaves began to develop. Root tips collected
after this change were very vigorous and provided excellent
material for study. Except on plants which became pot
nd, vigorous root tips could be collected through the
spring and summer up to the time when development of the
flowering stalk was initiated. Root tips collected after this
stage were few and most provided poor material for study.
The most satisfactory time of day for collection of root
tips for cytological study was between 9 and 11 a.m.,
although vigorous material could be collected at any hour
on comparatively cool days. Apparently, a daily cycle of sig-
popes increased root growth does not occur in species of
Excised root tips were pretreated in colchicine and fixed
in 1:3 acetic-alcohol. — preparations were made by a
Feulgen. jue.
Slides were pines and suitable chromosome comple-
ments were photographed at a magnification of 1350x or
1425x. From each complement photographed, a diagram
was made with a camera lucida, primarily for accurate
identification of each negative. In these di each
chromosome was represented by a line conforming to the
angle formed by the chromosome and in length proportional
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 133
to it. In instances where it was considered that a censtric-
tion, observable to the eye, might not be clearly seen in the
photograph, a short line showing the position of the con-
striction was placed across the line representing the chro-
mosome. The diagrams were annotated as necessary to in-
sure accurate interpretation of the photographs. These dia-
grams proved invaluable in properly interpreting certain
photographs in which 2 or more chromosomes were close
together or lying across one another.
Considerable difficulty was encountered in growing plants
of subg. Macrocline to maturity ; consequently voucher spec-
imens were not obtained from many of the plants examined.
Fortunately, all species and varieties of this subgenus are
readily identified in the rosette stage and there is no doubt
but that the determinations given are correct. Vouchers
were obtained for most of the plants of subg. Rudbeckia
examined. Available specimens will be deposited at the
Gray Herbarium and in the herbarium of the Texas Re-
search Foundation.
CHROMOSOME NUMBERS
During this study, chromosome numbers, obtained from
mitotic metaphases in root tips, were determined for 88
plants representing all 6 species of Rudbeckia subg. Macro-
cline, 8 of the 9 species of FR’ su and
4 species of the related genera Dracopis, Echinacea, and
Ratibida. The eed Priors of Rudbeckia not represented is
R.h (sul dbeckia), a very rare species of the
southeastern Mews States. For subg. Macrocline counts
were made for a single variety of each species; for subg.
Rudbeckia, — were made for 14 of the 22 specific and
subspecific tax:
Chromosome s onbets of Rudbeckia, Dracopis, and Ech-
inacea previously reported in the literature are as follows:
Rudbeckia laciniata n = 382-87 Fagerlind, 1946.
0 ee Ne £5 ttaglia, 1946c.
. laciniata
var. flore pleno Hort. bia Geass Battaglia, 1946¢.
R. bicolor (= R. hirta) a> 19 Battaglia, 1946a.
R. ioc (= R. hirta) 2n= 38 Battaglia, 1947.
R. hirta n= Battaglia, 1946a.
R. hirta
var. meine F: Hi n= 19 Battaglia, 1946a.
R. speciosa (= R. fulgida var.) 2n= 76 Battaglia, 6b.
ample. ulis n= 16 Battaglia, 1946a.
n= 11 Battaglia, 1946a.
134 ROBERT E. PERDUE, JR.
Chromosome numbers determined during the present
study, most of which are reported herein for the first time,
are given in Table 1, along with culture numbers, and col-
lector and place of collection.
TaBLe I. CHROMOSOME NUMBERS IN RUDBECKIA AND RELATED GENERA
SPECIES Cutture No. 2N Source
Rudbec!
subg. Macrocline
R. cali ica
var. californica 201-2 36 Above Rancheria Creek,
ca. 1 mile east of Hunt-
gton Lake, Fresno,
California.
was
202-2 36 Same source as 201-2.
202-4 36 Same source as 201-2.
R. laciniata
var. laciniata 42 72 Near Rock Hill Station,
Bucks Co., Pennsylvania.
M. G. Henry.
ca. 72 Same source as 4-2.
4-5 ea. 72 Same source as 4-2.
4-8 102+ Same source as 4-2.
28-1 54 Ca. 15 miles west of in-
tersection of Routes 31
and 92, Mahnomen Co.,
Minnesota.
B. Ownbey
28-2 36 Same source as a.
R. maxima 225-1 36 Along Texas Route 64,
R. mohrii 205 36 0.9 mile west of junc-
tion of Routes 71 and
22, Wewahitchka, Gulf
Co., Flori
R. E. Perdue, Jr. 1399.
207-2 36 205.
207-4 36 Same source as 205.
208 36 Same source as 205
R. nitida
var. texana 224-1 36 1 mile southwest of
Laporte, Harris Co.,
Texas,
R. E. Perdue, Jr.
224-2 36 Same source as 224-1.
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 135
Es Braye talis
occidentalis
. fulgida
var. palustris
R. fulgida
var. speciosa
224-3
57-1
253-2
55-2
251-1
36
36
ca. 76
ca. 76
Same source as 224-1.
Near Moran, Jackson
Hole Wildlife Park,
Teton Co., Wyoming.
C. L. Porter.
Same source as 57-1.
Near Naceville, Bucks
Co., Pennsylvania.
Heed
Same source as 2-1.
Near Finland, Bucks
Co., Pennsylvania.
. G. Henry.
Salem Valley, 3 miles
northeast of Ringold,
About 2 miles north-
west st Pe Marks along
Hyde Park, 1.6
miles phe junction with
Route 30, Wakulla Co.,
Florida.
R. E. pons Jr. 1658¢.
Same as
R. E. Ponies, Jr. 1658d.
3 miles northwest of
Greeley, Reynolds Co.,
Mi 2
ton Co., Arkan:
H. Iltis.
Along Turtle Creek
near per Turtle
Creek ad
miles north of junction
_ Route 16, Kare Co.,
aE E. Perdue, Jr. 1861.
Same source as 251-1.
from
era theses original-
ly oe Mitchell Co.,
Illinoi:
136 ROBERT E. PERDUE, JR.
Vee .
87 ca. 76 E. Battaglia
R. fulgida
var. umbrosa 3-1 76 Seed from a_ garden
ea
43-2 ca. 76 Same pie as 43-1.
R. graminifolia 209 38 0.9 miles north of Route
6.6 miles west of
Wewahitchka, Gulf Co.,
Florida.
R. E. Perdue, Jr. 1408.
210 38 Same source as 2
R. E. Perdue, Jr. 14084.
211 38 Same source as 209.
R. E. Perdue, Jr. 1409.
R. grandiflora
var. alismaefolia 252-1 38 Near Route 146, 2 miles
ea of Seabrook, Har-
is Co., Texas.
R. E. Perdue, chee 1971.
252-4 38 Same source as 252-1.
Ri a
var. grandiflora 227-1 38 Along railroad Pico
way, near Route 70, 12
miles iat ae Hugo,
Choctaw Co., Oklahoma.
R.
K
oO
#
Same source as 227-1.
hirta
var. angustifolia 52-3 38 Navco, Mobile Co.
abama.
G. N. Perdue.
R. hirta
var. hirta 18-1 38 Seed a herbarit
about 3 ies
west of Genesis, Ten-
E. 4 C. 14138.
22-4 38 Seed from herbarium
R. E. S., F. W. W., and
E. H. C. 14822.
R. hirta
ar. pulcherrima T-3 38 Harvard Forest, Peter-
sham, Worcester Co.
ee
R, E. Perdue, Jr.
12-1 38 Avenel, Silver Spring,
Maryland.
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 137
R. missouriensis
R. mollis
R. subtomentosa
12-2
2
13-3
15-10
37-1
239
246
45-1
46-1
46-2
ca. 38
38
38
38
BS
BS BS
D. S. Correll.
Same source as 12-1.
Seed from herbarium
s en: Be Co.
Tennessee.
T. Walker 16054.
10 miles south of Grand
Ownbey.
Hamilton Co, Ohio.
E. Bra
Tilinois
H. E. Ahles.
Massa Creek, south of
Jonesburg, Warren Co.,
White (junction
Routes 27 and 47) along
ug 27, Suwanee Co.,
Flori
R. E. as: Jr, 1663.
Near Route 20, 2.4 miles
east of Hawt
(junction of Routes 20
— Ph Putnam Co.,
E. or. Perdue Jr. 1817.
138 ROBERT E. PERDUE, JR.
R. triloba
var. triloba 1-4 57 A. F. Blakeslee.
19-1 ca. 38 Seed from herbarium
specimen: northeast of
Morristown, Hamblen
‘o., Tenne:
R. E. S., F. W. W., and
E. H. C. 8830.
19-3 38 Same source as 19-1.
20-1 38 Seed from herbarium
specimen: bottoms of
Turnbull Creek, near
Craggie Hope, Cheath-
am Co., Tennes:
R. E. S., E. H. C., and
Ws ~ 14788.
26-1 ca. 57 Near Sturbridge, Wor-
cester Co., Massachu-
setts.
R. E. Perdue, Jr.
26-2 57 Same source as 26-1
32-1 ea. 57 Hamilton Co., Ohio.
E. L. Braun.
32-2 ca. 57 Same source as 32-1.
34-1 57 ina.
E. Ahles
34-2 57 ie source as 34-1.
Dracopis
amplexicaulis 126-1 32 Near __ McLennan
Co., as.
M. P. Mauldin,
126-2 32 Same source as 126-1.
26-6 32 Same source as 126-1.
Echinacea
pallida 38 22 Iilinois.
. BE. Ahles.
233-1 44 + junction of Coit and
Road, ca. 2 miles
west of Rich 5
las Co., Texas.
R. E. Perdue, Jr.
233-7 44 Same source as 233-1.
Ratibida
columnaris 63-1 27 10 miles south of Scott
City, Scott Co., sas.
L. & O. S. Fearing.
63-2 26 ie source
R. pinnata 31-2 28 Adams Co., Ohio.
E. L,
31-10 28 Same source as 31-2.
All species of subg. are apparently diploid
: i Macrocline di
with a ThagtareegsSavesgtke Ohne A laciniata
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 139
ploid plants were found. In reporting a tetraploid plant of
R. laciniata as having 2n=76 chromosomes, Battaglia
(1946c) was apparently misled by the fact that other Rud-
beckia species that he studied do have a diploid number of
38
All plants of R. laciniata var. laciniata from which chro-
mosome counts were made were grown from seed. How well
these plants would survive in nature is not known. Of the
6 plants examined, only the specimen with 102+ chromo-
i ly in the I There
&
sembles ry d
collected only from several high mountains in a small area
of western North Carolina.
The four triploid collections listed in Table I were grown
from seed collected in the northern states and are fairly
typical of the northern form. The two diploid collections
form. One can only speculate as to the origin of the cyto-
140 ROBERT E. PERDUE, JR.
that hybridization between var. rupestris and the typical
southern form of var. triloba is responsible for the present
existence of triploidy and the present morphological varia-
tion among the northern representatives of var. triloba.
R. fulgida is a highly polymorphic species with 7 vari-
eties. Of the 4 varieties from which counts were obtained,
tetraploid plants were found in all but one. It is perhaps
more than coincidence that, for the species as a whole, the
polyploid collections originated from localities more or less
within the heart of the over-all area of distribution. In
contrast, the diploid collections are all from areas that are
at the periphery of the over-all distribution of the species.
Although no locality data were available for the collection
of var. umbrosa, this variety has so far been found only in
an area that lies almost at the exact center of distribution of
the species. R. fulgida is a very distinct species and it
seems unlikely that its polyploid nature originated as a
result of interspecific hybridization, but it may well have
resulted from hybridization between varieties.
y counts agree with those reported by Battaglia
(1946a) for Dracopis amplexicaulis (2n=32). A plant of
Echinacea pallida from one source proved to be diploid
(2n=22) ; plants from another source proved to be tetra-
ploid (2n=44). The diploid number of this species agrees
with the chromosome number reported by Battaglia for E.
purpurea. In Ratibida apparent diploid numbers of 26 and
28 occur; one plant was aneuploid with 27 chromosomes.
CHROMOSOME MORPHOLOGY
: Somatic chromosomes of the species of the 4 genera stud-
ied are comparatively numerous and though very variable
in size they are quite large. The position of the primary con-
striction (centromere) is usually very clear and in well-
spread figures can be observed in every chromosome. Sec-
ondary constrictions can occasionally be observed in species
of Rudbeckia. Easily distinguished satellites appear only
in pl ts of Rudbeckia hirta and Rati-
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 141
beckia subg. Macrocline, 8 species of Rudbeckia subg.
Rudbeckia, 2 species of Ratibida, 1 species of Dracopis and
1 species of Echinacea.
With the exception of the cl of Echii
pallida and Rudbeckia triloba, all of the chromosome com-
plements examined in the present study were diploid. The
use of tetraploid complements of Rudbeckia species was im-
practical as there were usually many chromosomes that
could not be accurately measured. As the data within each
of the genera and subgenera are comparatively uniform, it
is unlikely that analysis of polyploid complements would
contribute anything new. For most of the species or vari-
eties studied, 2 photographs were examined but in most
cases these were from the same plant. For Rudbeckia gram-
inifolia and R. triloba, single photographs were examined.
Three photographs representing Echinacea pallida were
studied.
The photographic records were made of cells in which
the chromosomes were darkly stained and, with a few ex-
ceptions, sufficiently dispersed so that the length of each
chromosome and the position of its primary constriction
could be clearly seen. It was impossible to get perfect
records in every instance, but there were never more than
a few chromosomes in any photograph for which the length
and constriction position could not be measured. From the
negatives exposed at a magnification of 1350x or 1425x,
photographic enlargements were prepared at a total mag-
nification of approximately 4000 diameters. From these en-
largements lengths of the chromosome segments were
measured in milli As the pari to be made
are relative, the measurements are hereafter referred to as
units.
Secondary constrictions were disregarded. In most cases
they could not be seen. When they were apparent, they
could usually be distinguished from the primary constric-
tion. In a few cases, the 2 constrictions were equally prom-
inent and it was necessary to compare the troublesome chro-
mosome with the data from other phot hs to determi:
which of the constrictions should be disregarded.
For each chromosome the total length was measured and
from this a relative length was calculated by dividing the
length of each chromosome by the average length of all of
the chromosomes in its complement. This was desi;
primarily to eliminate variation in chromosome length be-
tween different materials resulting from slightly different
142 ROBERT E. PERDUE, JR.
treatments with colchicine, or differing response to the
same colchicine treatment by different chromosome com-
plements of the same root tip or collection of root tips from
a single plant. Additional records made for each chromo-
some included measurements of the length of each of the 2
arms, and a value representing the position of the constric-
tion obtained by dividing the length of the long arm by the
length of the short arm. This method of obtaining a value
for the constriction position was selected as it always yields
a value of 1.0 or higher. Thus a value of 1.0 represents a
chromosome with an exactly median constriction, whereas a
value of 5.0 represents a chromosome which may be classi-
fied as subterminally constricted, the intermediate values
representing chromosomes in which the constriction occurs
at positions between these two extremes.
One of the major limitations in this study was the dif-
ficulty in obtaining sufficient data. When working with
plants with such high chromosome numbers, it is difficult
to obtain a large number of cells in which the chromosomes
are sufficiently spread so that the length of each and the con-
striction position show up clearly. In studies of plants with
lower chromosome numbers, where this difficulty is either
not encountered or is not so great, this method of chromo-
some analysis could be carried much further.
A number of errors were unavoidable even though good
photographs were used and extreme care was exercised in
making measurements. It is very difficult to get all the chro-
mosomes in a cell in perfect focus in one photograph. Any
chromosomes below or above the level of focus appears short-
er than it does when in perfect focus. This affects the meas-
urement of length and can affect the relative position of the
constriction, especially in measurements of short chromo-
somes and in measurements of long chromosomes when only
one end is out of focus. The error due to this source is believ-
ed to commonly be 0.5 unit and, as the error can involve both
ends of a chromosome, the total error for a single chromo-
some can readily be 1.0 unit. In certain instances it was
profitable to expose 2 or more negatives at different levels
of focus in order to get more accurate measurements for
the chromosomes lying above or below the level common to
most members of the complement. Although the chromo-
some measurements were carefully made, there was some
discrepancy between 2 independent sets of measurements
from identical photographs. This type of error is much
smaller than the type mentioned above and is not great
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 143
enough to cause any appreciable concern. A third source
of error is the differential shrinking of the chromosomes
by colchicine. While this does not affect the relative position
of the constriction it makes any comparisons of chromo-
some lengths difficult. In this study comparisons of chro-
mosome lengths were for the most part unnecessary.
During the course of the study, it was necessary to use two
different microscopes, the optical system of one having a
magnification of 1350x and that of the other a magnification
of 1425x. This difference is so slight as to be relatively
insignificant, especially since measurements of length play-
ed only an indirect part in interpretations of the data.
The data derived from the measurements of the chromo-
somes were organized in tabular form to show the total
length of each chromosome, the relative length, the length
of the long segment (L), the length of the short segment
(S), and the value representing the position of the con-
striction (L/S).
The large number of chromosomes in each complement
made a direct comparison of the tables very difficult. In
this study, scatter diagrams proved most satisfactory for
the demonstration of similarities or differences in karyo-
types. This graphic method of presentation has several
advantages. The data can be vividly presented without the
need for setting up arbitrary classes for chromosome
length or constriction position. The picture presented by
each graph is as accurate as could possibly be obtained;
facts, not possibly arbitrary interpretations, are presented.
Furthermore, it is much easier to compare the chromosomes
of one plant with those of another by study of the graphs
than it is by study of the tables. Finally, patterns of varia-
tion in chromosome length or constriction position that can
be observed in the tables or that are suggested by the tables
are very clearly confirmed or denied by graphs.
The data Le toate in the tables were used to prepare the
accompanying graphs. Graphs 1-12 pertain to species of
Rudbeckia subg. Macrocline and Graphs 13-28 to species of
subg. Dracopis is represented by
graphs 29 and 30, Ratibida by Graphs 31-34, and Echinacea
by Graphs 35-37. Relative chromosome length is plotted
along the vertical axis and L/S, the value representing the
position of the constriction, is plotted along the horizontal
axis. Two or more chromosomes of a complement of equal
length and with constrictions in the same position are rep-
resented on the graphs by the appropriate number of dots
144 ROBERT E. PERDUE, JR.
grouped closely around the correct point. This representa-
tion is judged to present a truer impression of the comple-
ment than would be obtained if only a single dot was used.
For example, ar single dot is used to represent duplicate
20 cannot be adequately por-
trayed on Graph 27 and 16 chromosomes cannot be adequate-
ly portrayed on Graph 28. The following diagram indicates
the numerical measurements used along the ordinate and
abscissa in Graphs 1 to 37.
Graphs 1-12 illustrate details of chromosome morphology
for all 6 of the species of Rudbeckia subg. Macrocline. The
species are similar in gross chromosome morphology and a
very definite pattern is established for this subgenus. The
chromosomes can be readily divided into 2 natural groups of
18 chromosomes each. One group includes comparatively
small chromosomes more or less medianly constricted (L/S
values mostly 1.8 or less) ; the other includes comparatively
=
—
2 1501
o
oe
o 125)
§
or oe
&
Ss ars
S
© 050,
2
BS o2s
oD
ec
T T T T T T T
10 20 30 40 5.0 6.0 7.0
Constriction Position (b/s)
large chromosomes the constrictions of which range from
submedian to a condition approaching subterminal (L/S
values mostly 1.7 or greater). Differentiation of the chro-
mosomes into 2 groups is not so distinct in the graphs rep-
resenting R. californica (Graphs 1 and 2) but is very dis-
tinct in the graphs representing the other 5 species.
From the standpoint of general morphology and geo-
graphic distribution it is very clear that R. californica and
R. occidentalis are more closely related to one another than
either is to another pair of closely related species, R. maz-
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 145
ima and R. nitida. The first 2 species are even more distantly
related to R. mohrii, the morphological extreme of the sub-
genus. If significant differences do occur between individual
species or species-groups within subg. Macrocline such dif-
ferences would be expected to show up in a comparison of
4 riots 4
7 ee i -
agit”
ee
] 28 7
T T T r T T T
Graph |. Rudbeckia californica Graph 2. Rudbeckia californica
var. fornica var. californica
4 Pane 4
s s . .
5 App US ge 7 ? .
° .
= e 4
e 2
4 Pes é 4 °
Graph 3. Rudbeckia occidentalis Graph 4. Rudbeckia occidentalis
var. occidentalis var. occidentalis
Graph 5. Rudbeckia laciniata
var. lacimata
146 ROBERT E. PERDUE, JR.
4 os 4
4 Bolt Se ie 4 Cneore
af, a.
3 7 2
° ona,
| = eee
oS ae T T T T
gig 7 Rudbeckia maxima Graph 8. Rudbeckia maxima
s
5 a 4 ° s 7
T T oo ee T T
Graph 9 Rudbeckia nitda Graph 10. Rudbeckia nitida
var texana var texana
Ag
%
| = | #
T
Graph IL Rudbeckia mohru Graph 12. Rudbeckia mohru
the chromosomes of R. 29 hate and R. occidentalis on the
one hand with those of R. maxima and R. nitida on the
other. Either species pair might also be expected to show
differences setting them apart from R. mohrii. From the
graphs it is clear that such distinctions are not indicated
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 147
T T r T
Graph 13. Rudbeckia hirta Graph I4 Rudbeckia hirta
var. pulcherrima var. pulcherrima
F T T T
Graph 15. Rudbeckia missouriensis Graph I6. Rudbeckia missouriensis
eR es EAT
Graph IZ Rudbeckia fulgida Graph 18. Rudbeckia ae
var. fulgida var. fulgida
within subg. Macrocline and it is likely that additional data
would show the same uniform pattern for all the species.
Details of chromosome morphology of 8 of the 9 species
of db su dbeckia are portrayed in Graphs
13-28. More or less ‘clumping of the dots is characteristic of
148 ROBERT E. PERDUE, JR.
| aoe
:
.
J ‘i J heen
ae
4 a :
.
T T T T ots T
Graph I9. Rudbeckia fulgida rat 20. ae fulgida
var. palustris ar. palustris
4 = a
: one 4 %
.
eee | ~ S
.
4 ze" J 2.
T r T T T T T T
Graph 21. Rudbeckia mollis Graph 22. Rudbeckia mollis
ss . 4
“ a 2
er ?
1 &R 1 oe
= 1 <
oe .
T T T T J T T T
Graph 23. Rudbeckia grandiflora Graph 24. Rudbeckia grandiflora
vor. grandiflora var grandiflora
all species of this subgenus; that is, there is no separation
of the chromosomes into groups on the basis of either length
or constriction position. The constriction position varies
from median to more or less submedian.
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 149
Graph 25. Rudbeckia subtomentosa
T T
Graph 27 Rudbeckia triloba
‘ar, triloba
4 ES
4 Cabiee
os i
yy * .
a .
Graph 29. Dracopis amplexicaulis
T T T
Graph 28. Rudbeckia graminifolia
1 A
+ ace .
fe
aed
Graph 30. Dracopis amplexicaulis:
:) ee
In contrast to the morpl
subg.
Macrocline, in which all of the species are fairly closely re-
lated, subg. Rudbeckia includes species and species groups
that are much more diverse in their relationships. Ex-
amples of species that are highly distinct morphologically
150 ROBERT E. PERDUE, JR.
Graph 31 Ratibide columnoris Graph 32. Retibida columnaris:
Groph 33. Ratibida pinnata Graph 34. Ratibida pinnata
are R. hirta (Graphs 13 and 14), R. triloba (Graph 27), and
R. graminifolia (Graph 28). R. missouriensis (Graphs 15
and 16) and R. fulgida (Graphs 17-20) are much more
closely related to one another than to any other species.
This is also true of R. grandiflora (Graphs 23 and 24) and
B. subtomentosa (Graphs 25 and 26). The graphs do not
indicate any differences of greater magnitude between the
most distantly related species than between closely related
species or between 2 varieties of the same species. Compare
the 2 varieties of R. fulgida in Graphs 17-20. Even though
the graphs indicate that the morphological patterns of the
chromosomes of species of this subgenus are comparatively
uniform, it is possible that significant differences could be
established by study of additional material.
Graphs 29 and 30 represent the chromosomes of Dracopis
amplexicaulis, the only species of the genus. Although the
data for this species are so limited as to be very inconclusive,
Graph 29 was derived from an excellent photograph
(Fig. 19) and probably represents fairly accurately the
pattern of chromosome morphology in this species. This
graph suggests that the chromosomes can be separated into
2 groups. In the first group, consisting apparently of 1 pair,
the chromosomes are more or less subterminally constricted
(L/S values of 3.25 and 4.00). In the second group the
position of the constriction varies from median to sub-
median (L/S values of 2.0 or less). There is no separation
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 151
&
4 me s Selec . zs =
T T T T T T
Graph 35. Echinacea pallida
4 res
bce arte’ ilies
2 Seng eee
Graph 36. Echinacea pallr
°
£* ‘
a $ Pid te ao ° per
r T T r
Graph 37 Echinacea pallida
of the members into groups on the basis of size. As evi-
denced by the illustrations (compare Fig. 19) the chromo-
somes are much smaller, on the average, than are those
any other of the species studied. This smaller size was ap-
parent in all of the material examined.
152 ROBERT E. PERDUE, JR.
Graphs 31-34 illustrate details of chromosome morph-
ology of 2 species of the genus Ratibida. Dispersion of the
dots is characteristic of this genus, the result of consider-
able variation in chromosome size and especially of varia-
tion in position of the constrictions. The chromosomes can
be separated into 3 groups. The first group, consisting ap-
parently of but a single pair, is relatively long and more or
less medianly constricted (L/S values of 1.0-1.6). Another
group of chromosomes is comparatively short, but also more
or less medianly constricted (L/S values of 1.0-2.0). The
third group includes chromosomes of median length that
are constricted from the submedian position to the sub-
terminal position (L/S values mostly greater than 2.0).
Graphs 33 and 34 represent Ratibida pinnata, which has a
chromosome number of 2n=28. Graph 32 represents R.
1: is (R. col: ifera of manuals), which has a chro-
mosome number of 2n=26. The constriction of one long
chromosome could not be observed and this chromosome is
not represented on the graph. An aneuploid plant of R.
l is with 27 ek is rep ted by Graph 31.
The graphs clearly show that the aneuploid plant of R.
columnaris differs by the presence of an extra, medianly
constricted, small chromosome. The graphs further indicate
that R. pinnata differs from R. columnaris in having 2 addi-
tional, small, medianly constricted chromosomes.
Graphs 35-37 all represent Echinacea pallida. A diploid
vlant with 22 chromosomes is portrayed in the first graph;
a tetraploid plant with 44 chromosomes is represented by
the last two. The chromosomes are more or less uniform in
size as evidenced by the distribution of the dots in a more
or less straight, horizontal line. There is great variation,
however, in the position of the constrictions and on this
pasis the chromosomes can be easily separated into three dis-
tinct groups. One group is more or less medianly constricted
(L/S values of 1.0-1.8), a second is constricted from sub-
medianly to more or less subterminally (L/S values of 2.0-
4.0), and the third is constricted in a position that is dis-
tinctly subterminal (L/S values of 4.0 or greater).
CONCLUSIONS
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 153
presented for the other genera are inconclusive and must
be regarded as tentative although they probably present a
fairly accurate picture, especially for Ratibida. If minor
exceptions are disregarded, definite patterns of chromosome
morphology are clearly established in the major taxa con-
sidered, and uniformity is the rule within each of these
groups. The most important conclusion to be drawn from
the data presented is the fact that morphological differen-
tiation of Rudbeckia into 2 distinct subgenera has been ac-
companied by visible di tiation in ck struc-
ture as well as by a change of chromosome number. These
data support the recognition of subg. Macrocline and subg.
Rudbecicia as 2 distinct very natural taxa, and they support
th 2x. of DR. Ih ya .
e Ratibida, Dracopis and Echi-
nacea as distinct genera. The genera and subgenera dis-
cussed can be distinguished from one another by the mor-
phology of their somatic chromosomes as well as by their
chromosome number.
The patterns of chromosome morphology exhibited by
Rudbeckia, Ratibida, Dracopis, and Echinacea, based upon
the material examined (limited as it may be for the last
two genera), are compared in Table 2. For each genus or
subgenus considered in this paper, the table lists the basic
morphology. For the genera studied the information de-
rived from the chromosomes is as much within the realm of
morphology as are the characteristics of the achene. These
are additional morphological characters, however, in im-
portance, characters of the greatest magnitude.
The chromosome numbers of R ibeckia subg. J
(2n=36), Ratibida pinnata (2n=28), and Dracopis
(2n=82) raise a question as to the possibility that the last
genus was derived from a hybrid between the other two.
Tf an egg cell of a species of Ratibida with n—14 was fertil-
ized by pollen from a species of Rudbeckia subg. Macrocline
with n=18, the resulting plant would have a chromosome
number of 2n=32. Apomictic development would be neces-
rr,
Rudbeckia subg. Rudbeckia
Qn = 38
No, species: 9
No, species examined: 8
Average chromosome length:
16,87 units. eee 8s
divided into natural
as to length or ee
position,
womes medianly to sub-
medianly constricted, L/S 1,0-
2
OMT
TABLE 2.
Rudbeckia subg. Macrocline
2n = 36
No, species: 6
No, species examined: 6
chromosome length:
units, Chr eae os
Average
14,25
vided fate two naturs
a th a and Dae
Group 1, Short ehromosomes
nee to sub-medianly con-
icted, L/S mostly 1,0-1.8
2, Long chromosomes
more or les!
constricted,
L/S mostly 1,7-3.3.
CYTOLOGICAL DIFFERENTIATION OF RUDBECKIA AND
Dracopis
2n = 82
No, species: 1
No, species examined: 1
Average chromosome lengt!
8.56 units, Chromosomes ai
vided into two Liles ‘ie:
8 to constriction
Group 1, Chromosomes medi-
anly to sub-medianly constric-
ted. L/S 2.0 or less,
Group 2 Chromo:
or less he teeny constric-
ted. L/S
(Data oa Bactanietee)
RELATED GENERA
Ratibida
2n = 26, 28
No, species: 5
examined: 2
No. species
°
Average chromosome length:
16.32 units, Chromosomes di-
vid into three r
groups as to length and con-
striction position
Group 1, Long chromosomes
medianly to —sub-medianly
constricted, L/S 1.0-1.6,
Group 2. Short chromosomes
medianly to sub-medianly
stricted, L/S 1,0-2.0,
Group 8. Chromosomes — of
medium length, sub-medianly
to sub-terminally Ses e:
L/S mostly greater than 2.0,
Echinacea
2n = 22
No. species: 5
No. species examined: 1
and dn races)
Average chromosome length:
15.94 units, Chromosomes di
vided i
into three natural
groups as to constriction posi-
tion,
sub-
1.0-
Group 1. Medianly to
medianly constricted, L/S
1.8.
Group 2. Sub-medianly to sub-
terminally constricted. L/S
2.0-4.0,
8. Strongly sub- ee
ra P aaestoea, L/S
VST
‘ar ‘andudd “a Luadou
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 155
sary to the reproduction of such a plant and its establish-
ment in nature. An examination of the chromosomes of the
three genera, while not completely voiding the possibility,
does not support a theory of hybrid origin for Dracopis.
First, the chromosomes of Dracopis are conspicuously small-
er than those of species of Rudbeckia and Ratibida. While
the morphological patterns of the chromosomes of Rud-
beckia subg. Macroline and Ratibida are somewhat similar,
those of Dracopis are distinctly different from each of the
other two. An intergeneric hybrid between species of
Rudbeckia and Ratibida would also display a distribution
of 3 more or less distinct groups of chromosomes and
would be expected to have more chromosomes with subterm-
inal constrictions. From these data a theory of the hybrid or-
igin of Dracopis cannot be completely negated but it seems
very unlikely unless the chromosomes of this species have
changed considerably since the initial hybridization.
Battaglia (1946a) reported that Dracopis is fully sexual,
reproducing according to the F’ ritillaria-type of embryo-sac
formation. This fact also argues against the possibility of
hybrid origin of the genus.
BIBLIOGRAPHY
BATTAGLIA, E. 1946a. Ricerche iologi ed 1 he sul genere
Rudbeckia (Asteraceae). I-V: Il gametofito femminile e maschile di
R. bicolor Nutt., R. hirta L., R. hirta L. var. meine freude Hort., R.
amplericaulis Vahl, e R. purpurea L. (= Echinacea purpurea
Moench.). Nuovo Gior. Bot. Ital., n.s. 53:1-26.
——. 1946b. Ricerche iologiche ed briologiche sul genere
Rudbeckia (Asteraceae). VI: Apomissia_in Rudbeckia speciosa
Wender. Nuovo Gior. Bot. Ital., ah 53:27-69.
. 1946c. Ricerche iologiche sul genere
Rudbeckia (Asteraceae). VII: Apomissia in Rudbeckia laciniata
L. issia nella sua varieta a fiori doppi. Nuovo Gior. Bot.
482.
Met Pec pe Me
S|
. 1947. Ricerche iologiche ed genere .
beckia (Asteraceze). XII: Il gametofito femminile e maschile di
Rudbeckia flava Greene, con particolare riguardo al suo comporta-
mento di ibrido strutturale. Nuovo Gior. Bot. Ital., n.s. 54:560-567.
1952. ili: i di i briol ici in funzione di una
migliore conoscenza della sistematica del genere Rudbeckia (Com-
positae). Atti Soc. Toscana Sci. Nat. (Pisa). Ser. B. 59:217-221.
FAGERLIND, F. 1946. Sp y i
d bild bei Rudbeckia laciniata L. Acta Horti
Bergiani 14:39-99. ;
FERNALD, M. L. 1950. Gray’s Manual of Botany. 8th ed. American
Book Co., New York.
GuEason, H. The New Britton and Brown Illustrated
Flora of the Northeastern United States and Adjacent Canada.
New York Botanical Garden, New York.
156 ROBERT E. PERDUE, JR.
ee Asa. 1884. Synoptical Flora of North America. Vol. 1. Pt. 2.
URNEEK, A. E. 1940. of day and temperature coe in
gree Bot. Gaz. 102:269-279.
E., JR. 1957. Synopsis of Rudbeckia subgenus Rud-
bekia Rhodora 59: 293-299.
. Monograph of Rudbeckia subgenus Macrocline. Contrib.
Tex. eaaee Found. (In press).
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 157
oe
’ am Gi)
o O
—- |
~ os | “~G =
oe kh # or ary
3 oe Ot od
=~"
s Sed
S
sua = =
ated ta‘? es
<— ha » f Sn ,
= -
Fig. 1 Rudbeckia species, mitotic metaphases
from root tips.
R. ar
ROBERT E. PERDUE, JR.
~
ce
> ee
5 a aw
= fT ; sy
,ore ¢ é
Fg ad ve 4!
we ‘x
= of”
:
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 159
160 ROBERT E. PERDUE, JR.
CHROMOSOMES OF RUDBECKIA AND RELATED GENERA 161
18 a
q z y — py
-<
~ ~ a.
of Rudbeckia, Dracopis, and Ratibida species, mitotic
18. Rudbeckia triloba var. trileba, 2n= ST .
Fig. 20. Ratibida pinnate
Fig. 18-20. Chromosomes
metaphases from root tips. Fig.
Fig. 19. Dracopis amplexicaulis, 2n=82 (1350x).
(1350x).
162 ROBERT E. PERDUE, JR.
Fig. 21-28. Chromosomes of Ratibida and Echinacea species, mitotie metaphases
from root tips. Fig. 21. Ratibida columnaris, 2n=27 (1425x). Fig. 22. Echinacea
Wlida, 2n=22 (1425x). Fig. 23. E. pallida, 2n=44, (1350x).
eS FROM THE GRAY HERBARIUM
F HARVARD UNIVERSITY
Edited by
Reed C. Rollins and Robert C. Foster
NO. CLXXXVI
A MONOGRAPHIC STUDY OF THE CYRILLACEAE
os :
Joss L. THomas
Published by
THE GRAY nee OF HARVARD UNIVERSITY
RIDGE, MASS., U. S. A.
1960
CONTRIBUTIONS FROM THE GRAY HERBARIUM
OF HARVARD UNIVERSITY
Edited by
Reed C. Rollins and Robert C. Foster
NO. CLXXXVI
A MONOGRAPHIC STUDY OF THE CYRILLACEAE
By
JoaB L. THOMAS
Published by
THE GRAY HERBARIUM OF HARVARD inarerncsis
CAMBRIDGE, MASS., U. S. A.
1960
Issued May 27, 1960
A MONOGRAPHIC STUDY OF THE CYRILLACEAE
BY
JoaB L. THOMAS
INTRODUCTION
The Cyrillaceae is a family of shrubs and small trees dis-
tributed on the Coastal Plain of southeastern United States,
in middle America, including the West Indies, and in north-
ern South America. The family, as recognized in the present
treatment, is composed of three genera and thirteen species.
Although the three genera are very distinct morphologically,
the family appears to be a natural unit, and is easily distin-
guishable from related groups. On the basis of a study of
fossil pollen it is evident that the Cyrillaceae is also an old
family, with definite Cyrillaceous pollen occurring in de-
posits of Upper Cretaceous age (Steeves, 1959). The genera,
also, have apparently been distinct at least since Upper
Oligocene (Traverse, 1955).
In addition to a taxonomic revision of the three genera
the present treatment includes studies of the morphology,
geographic distribution, paleobotanical history, evolution,
species relationships, and infraspecific variation. The study
of herbarium material was supplemented by two summers
of field work in southeastern United States, and a short
pronounced. ‘A ;
Although considerable information has been obtained in
the present study, there are several interesting problems
in the Cyrillaceae which need further work. Additional field
4 JOAB L. THOMAS
studies are needed on the ecology and infraspecific variation
of several species of Purdiaea in eastern Cuba. Further work
is also needed on the origin and development of root sprouts
in Cyrilla, particularly morphogenetic studies of the early
stages of bud formation. The cytology of the entire group
is virtually unknown ons ere en study.
KNOWLEDGEM
I am deeply indebted to eons Gat C. Rollins, Director of the
Gray Herbarium, under whose guidance the research was carried out.
His helpful suggestions and criticisms during the preparation of this
work, and his assistance in obtaining funds for research and field
work are greatly appreciated.
Dr. Robert C. Foster has answered many questions on nomenclature,
and Mrs. L. Schwarten has given valuable assistance in the library.
The Latin diagnosis was prepared by Mr. Leslie A. Garay. Dr. C. E.
Kobuski has been very helpful in obtaining loans of herbarium speci-
mens and in providing storage and work space in the Harvard Uni-
versity Herbarium. Helpful suggestions and criticisms have been
offered by several other persons associated with the Herbarium, in-
eluding Dr. C. E. Wood, Dr. K. A. Wilson, Dr. Arthur S. Barclay,
Mr. George Argus, and Dr. W. P. Adams.
Professor Ralph H. Wetmore has shown a great deal of interest
in the morphological studies and has offered many valuable sugges-
tions. Professor Elso S. Barghoorn has given much assistance in the
paleobotani tanical studies and has generously provided equipment for the
of p
Dr. R. K. Godfrey of Florida State University has kindly provided
cytological material and several special mass collections of dried
material. A summer of field study was made possible by a grant from
the Fernald Fund For Study in Systematic Botany, provided through
the generosity of Mr. F. W. Hunnewell of Wellesley, Massachusetts.
Research the summer of 1958 was supported by a National In-
stitute of Health grant-in-aid through the Department of Biology,
Harvard University.
I wish to extend special thanks to my wife, Marly Dukes Thomas,
cient whose assistance and understanding this study would not have
sible.
I am indebted to the a of the following herbaria for the loan
of specimens used in this work:
Arnold Arboretum (A); pds Natural History Museum (F);
University of Florida (FLAS); Gray Herbarium (GH); Herbario de la
Salle, Habana (Ls); University of Michigan (MicH); Missouri Bot-
anical Garden (Mo) ; Université de Montréal Qt) 5 New York Botan-
ical Garden (Ny); Estacién Habana (sv);
University of Texas (TEX); University of California (uc); United
States National Museum (us).
HISTORY OF THE GENERA
CYRILLA
Cyrilla racemiflora was described by Linnaeus in 1767
MONOGRAPH OF CYRILLACEAE 5
from material sent to him by Dr. Alexander Garden, a resi-
dent of South Carolina. The name Cyrilla was given in
honor of Dr. Domenico Cirillo, an Italian physician and
professor at Naples. Michaux (1803) described a second
species, Cyrilla antillana, apparently from material collected
in the West Indies, but no specimen or locality was cited.
A few years later, however, Poiret (1812) noted that the
characteristics used to distinguish the two species inter-
graded to such an extent that he could not separate them,
and concluded that they must be varieties of the same spe-
cies. This seems to have established a precedent, for in the
years to follow ten more species were described in this
genus, and in most cases later workers found the species
difficult to segregate due to numerous intermediate forms
which seemed to link one species to another. Botanists who
were familiar with this genus in the field were particularly
aware of these intermediate forms.
Rafinesque (1840) noted that the various collections of
Cyrilla from the United States showed considerable varia-
tion in leaf size and shape, but stated that he had found
in the fruits “... characters for three distinct, blended spe-
cies.” Subsequent authors agreed that these species were
blended, and two of the three species were not recognized
in any later works, the names seldom appearing even as
synonyms. The third species, Cyrilla parvifolia Rafinesque,
has been recognized by several workers, although the author-
ship is usually attributed to Shuttl th (1896) who des-
cribed a similar plant under the same name. Sargent (1921)
reduced Cyrilla parvifolia to a variety of C. racemiflora,
stating that the plants of the variety “... afford no mor-
phological characters by which it can be distinguished spe-
cifically from that species.” One additional species in the
United States, Cyrilla arida, was described by Small (1924)
from material collected in the southern lake region of Flo-
rida.
N. E. Brown (1901) described a new South American
species of Cyrilla, C. brevifolia, from material collected in
British Guiana. However, Gleason (1931), after studying
additional collections from this area, stated that “...the
differences between this species and the following (C. race-
miflora) are slight and scarcely entitle it to specific rank.”
Three additional species of Cyrilla have been described
from eastern Cuba: one by P. Wilson (1902) — Cyrilla
cubensis, and two by Urban (1926) —C. nitidissima and
C. nipensis. These species have been recognized in most
subsequent treatments of the group, but at least one worker,
6 JOAB L. THOMAS
Carabia (unpublished)', concluded that all of the repre-
sentatives of Cyrilla growing in Cuba belonged to the ori-
ginal Linnaean species, C. racemiflora.
In spite of widespread disagreement as to the validity of
the various species of Cyrilla, Mattick (1935) recognized
nine of the above species, excluding the two Rafinesque
species only. This treatment was also followed by Uphof
in Engler and Prantl (1942). Yet there was still con-
siderable disagreement in the literature, and some authors
(e.g. Lawrence, 1951) continued to refer to Cyrilla as a
monotypic genus. The situation was accurately summarized
by Gleason (1952) as follows: “The genus is by some
considered to be monotypic, by others as many as ten spe-
cies are recognized.”
CLIFTONIA
In contrast to Cyrilla, Cliftonia has been treated as a
monotypic genus by every author who has dealt with the
group. However, it is such a striking and unusual looking
plant that it has been described as a new genus on several
different occasions. It was originally described by Lamarck
(1791) as a species of Ptelea, P. monophylla. The original
description was based on fruiting material only, and ap-
parently the winged fruit of Cliftonia was the characteristic
which caused Lamarck to include it in the genus Ptelea.
Fourteen years later Gaertner f. (1805) described a new
genus and species, Cliftonia nitida, unaware that it was
the same as Lamarck’s species of Ptelea. Britten (1905)
pointed out that the name Cliftonia was given in honor of
D.G. Clifton, Chief Justice of West Florida, and not Francis
Clifton, the English physician, as was stated by Sargent
(1892). Four years after Gaertner’s description of Clif-
tonia, this taxon was again described as a new genus and
species, Mylocaryum ligustrinum, by Willdenow (1809).
A fourth generic name for this taxon, Walteriana, was
published in stoereate by Endlicher (1841) from the manu-
script of Frase
There was citeahdeeabe confusion in the nomenclature
of Cliftonia for ee years as various combinations of
re used by different authors. Finally
Britton (1889) ities out by implication that the correct
combination was Cliftonia monophylla. He did not make
the combination, however, and the first author to use the
correct combination was Sargent (1892), who attributed
? This appears on annotation labels which read, “Revision of Cuban Plants — det.
31F. utaheas er apparatus torscnutioas bis published.
MONOGRAPH OF CYRILLACEAE 7
the combination to Britton. In recent years there has been
little confusion in either the nomenclature or the taxonomy
of Cliftonia.
PURDIAEA
The genus Purdiaea and a single species, P. nutans, were
described by Planchon (1846) from material in the Hooker
Herbarium. It was named in honor of Purdie who col-
lected it in New Granada (Colombia). The second species
of Purdiaea was described by A. Richard (1853) as a new
genus and species, Costaea cubensis, from material col-
lected in Cuba by Valenzuela. For unexplainable reasons,
most subsequent authors adopted the later name Costaea,
treating the older name Purdiaea as a synonym. The correct
combination Purdiaea cubensis was first made by Urban
(1926).
third species, Purdiaea stenopetala, was described by
Grisebach (1860) from the collections of Charles Wright
in Oriente Province, Cuba. Three additional species from
eastern Cuba (P. velutina, P. microphylla, and P. shaferi)
were described by Britton and Wilson (1915) from the
extensive collections of Shafer in Oriente Province.
From collections of Purdiaea in Peru made by Weber-
ved Gilg (1931) described a new genus and species,
Uoiosepalum weberbaueri. Mattick (1935) noted that Gilg
ie later become aware that his new genus was actually a
species of Purdiaea but was prevented by his death from
making the transfer. Mattick made the transfer to the
Sei Purdiaea, recognizing it as a distinct species, P. weber-
“The — treatment of the group is a revision of the
Cuban species of Purdiaea by Marie-Victorin which was
edited pee published posthumously by J. Brunel in 1948.
In that treatment, four species (P. ekmanii, P. moaensis,
P. nipensis, P. ophiticola) and three varieties (P. cubensis
var. albosepala, P. nipensis var. alaini, P. ophiticola var.
parvifolia) were described as new.
CYRILLOPSIS
The genus Cyrillopsis Kuhlmann (1925) was included in
the Cyrillaceae by Mattick (1935) and Uphof in Engler and
Prantl (1942). This genus is excluded from the Cyrillaceae
in the present treatment on the basis of wood anatomy,
pollen morphology, and several aspects of the fiower struc-
ture (discussed below). It is a that this genus
possibly belongs to the Ce
8 JOAB L, THOMAS
ECONOMIC IMPORTANCE
Members of the Cyrillaceae are of only minor economic
importance. Both Cyrilla and Cliftonia are occasionally cul-
tivated as ornamentals in southeastern United States, but
are seldom cultivated outside of this area except in botanic
gardens. These plants can be grown considerably north of
their natural range. Cliftonia has been grown as far north
as Delaware and southern New Jersey ; Cyrilla as far north
as Massachusetts (Rehder, 1940). A specimen of Cyrilla has
been in cultivation at the Arnold Arboretum in Jamaica
Plain, Massachusetts since 1926. Although the plant is some-
what dwarfed and the reproductive structures do not de-
velop normally, it is a very attractive shrub and flowers
abundantly each year.
The bark of Cyrilla is rich in phenolic compounds, and
there are a few reports in the literature of the use of this
bark as a styptic or astringent (Sargent, 1892). According
to Elliott (1821) the bark of Cyrilla can also be used as
“| a serviceable application to wounds or ulcers, where
the indication is to cicatrize them.”
In some areas of southeastern United States Cyrilla is
valued in the production of honey. The glandular, secretory
, the long flowering period, and the large number of
flowers produced each year combine to make this plant an
unusually good source of nectar. The honey is known locally
as “titi” honey, in reference to the colloquial name for
Cyrilla.
MATERIALS AND METHODS
The plant materials used in the morphological studies
were primarily from my own collections, although herb-
arium material was used in a few instances. A list of all
collections used, giving the area from which they were col-
and the herbarium in which the collections were de-
posited, is included in the appendix. —
Floral buds were fixed in either Carnoy’s solution with
chloroform, or Formalin-Acetic acid-Alcohol. Those fixed
in Carnoy’s solution were transferred after two to five hours
into 70% ethyl alcohol; those fixed in FAA were stored in
the same solution. The buds were dehydrated in a normal-
butyl alcohol series, embedded in paraffin and sectioned with
a rotary microtome. The sections were stained with either
safranin and fast green or crystal violet, and mounted in
Stems and older wood were softened in hot water and
sectioned, without embedding, with a sliding microtome.
MONOGRAPH OF CYRILLACEAE 9
Wood sections were stained with Heidenhain’s Haematox-
ylin and safranin, and mounted in diaphane.
The leaf preparations were made by hand sectioning with
a razor blade. In a few cases leaves fixed and stored in FAA
were used, but it was found that satisfactory preparations
could be made from dried material. The dried leaves were
softened in warm water, with a little detergent added, for
approximately two hours before sectioning. The sections
were mounted, unstained, in glycerin.
VEGETATIVE MORPHOLOGY
STEM. — In Cyrilla and Cliftonia the young stems are gla-
brous and are usually angled or ridged. The ridges tend to
form a spiral pattern around the stem with each ridge evi-
dent for a distance of approximately two nodes, ending at
the base of a leaf. The ridges become less prominent as the
stem increases in diameter, and by the end of the second
growing season the stems are more or less terete. In Pur-
diaea the external morphology of the stem varies consider-
ably from one species to another, ranging from the small,
delicate, somewhat vine-like stems in P. parvifolia, to the
robust, rigidly erect stems of P. velutina. The robust stems
are usually more sharply ridged than the smaller stems, but
in all members of the Cyrillaceae the young stems are angled
to some extent. The stems of Purdiaea velutina and P. moa-
ensis are always densely pubescent near the end. In other
species, such as Purdiaea cubensis and P. nutans, the stems
are always glabrous, whereas in P. nipensis the stems may
be either glabrous or puberulent.
Cork formation begins early in the stems of all three
genera. Although the periderm layer is relatively thin in
young stems, it often becomes quite thick and spongy in
older stems, especially in Cyrilla. It is apparently this fea-
ture which accounts for the common name “Leatherwood”
which is often applied to Cyrilla and occasionally to Cliftonia
in the southeastern United States. A single layer of phello-
derm is developed in the young stems of Cyrilla, Cliftonia
and probably of Purdiaea, although it has not been verified
in the latter genus. The primary cortex contains numerous
thick-walled parenchyma cells with wide lumina (Metcalfe
and Chalk, 1950). The cortex, however, is rapidly destroyed
by the periderm which develops rather early in the inner-
most layers of the cortex. The primary vascular system
of the stem is a eustele with numerous primary vascular
bundles surrounding the pith. The primary xylem is com-
10 JOAB L. THOMAS
t ¥ owth rin I
ction N ly thickened cell w
radial
monophylla, m © t wood showing an un
ial section thre a
MONOGRAPH OF CYRILLACEAE 11
posed of spirally-thickened tracheids in the protoxylem with
a rapid transition in the metaxylem to tracheids with bor-
dered pits, fiber-tracheids, and vessels (Fig. 5). The pith
is 5-angled in transverse section. It is composed of relatively
thick-walled, parenchymatous cells, many of which are filled
with densely-staining phenolic compounds.
MATURE WooD. — Growth rings, present in all three gen-
era, consist of from two to five rows of fiber-tracheids which
are thicker-walled and usually somewhat smaller in radial
dimensions than the fiber-tracheids in the “spring wood”
(Figs. 1 and 2). The vessels are usually abruptly smaller
and fewer in number near the end of a growing season than
earlier in the season. In the tropical representatives the
growth rings are obscure, and in some preparations it is
difficult indeed to designate a definite growth ring. This, of
course, is expected in a tropical or sub-tropical tree growing
along stream margins where there is little seasonal fluctua-
tion in either temperature or moisture.
The vessel elements are small with a tangential diameter
of 25-50 micra for Cyrilla and Cliftonia (Metcalfe and
Chalk, 1950). No old wood of Purdiaea has been seen, but
the vessel size in the young stem is comparable to that in
Cyrilla and Cliftonia at a similar stage of development
(Fig. 6). In Cyrilla the vessels are solitary; in Cliftonia
pore multiples occur, particularly in the late wood ; and in
Purdiaea pore multiples occur h t the
secondary xylem of the young stems. The vessels are numer-
ous in all three genera, more so in Cliftonia and Purdiaea
than in Cyrilla. The vessel elements are thin-walled and
somewhat angular in cross-section. The end walls are ob-
lique, and there is considerable overlap of consecutive ele-
ments in the vessel (Fig. 7). This phenomenon accounts for
most of the apparent tangential pairs of vessels seen in
transverse section of Cyrilla. The perforation plates are
scalariform, consisting of numerous small bars (Fig. 8).
There is an average of 30-40 bars per end wall, but as many
as 65-70 have been counted in some preparations of both
Cyrilla and Cliftonia. The intervascular pitting is rather
variable, even in a single plant. Scalariform, transitional,
and opposite intervascular pitting have been seen in a single
preparation, although transitional to opposite pitting is by
far the most common type. The mean length of the vessel
elements in Cyrilla and Cliftonia is approximately 0.8 mm.,
with a range of 0.6 to 1.0 mm. (Bailey and Tupper, 1918).
Dark-staining deposits occur frequently in older vessels, but
12 JOAB L. THOMAS
-
of the Cyrillaceae. Fig. 7. Cy emiflora, tangential
ing the upright cells on the ys. Fig. 8. Clif-
f mature wood, showing a typical vessel end wall
section of mature
ng of tracheids
eids.
e the
beyond the pit border.
sf and
‘e extends only slightl:
MONOGRAPH OF CYRILLACEAE 13
no tyloses have been seen in any members of the Cyrillaceae.
e imperforate tracheary elements are primarily fiber-
tracheids in the older wood, but some tracheids occur in
both Cyrilla and Cliftonia (Fig. 9). The fiber-tracheids pro-
duced early in a growing season have relatively thin walls;
those produced late in the season have moderately thick
walls. Small, circular-bordered pits are abundant on both
the radial and the tangential walls (Figs. 4 and 9). The
mean length of the fiber-tracheids is approximately 1.0 mm.
with a range of 0.6 to 1.3 mm. (Bailey and Tupper, 1918).
The rays are heterogeneous and may be either uniseriate
or multiseriate (Fig. 4). The uniseriate rays are usually
only 3-5 cells high, but may be up to 12 cells high. Those
with 3-5 cells are usually composed of only upright or square
cells, whereas those containing 8 or more cells are com-
posed primarily of procumbent cells. The multiseriate rays
contain mostly procumbent cells, but there are usually up-
right cells on each end, often forming wings (Fig. 7). Most
of the multiseriate rays are 4-6 cells wide and 20-30 cells
high, although exceptionally large rays up to 8 cells wide
and 50-60 cells high have been observed. The cell walls in
the rays are usually very irregular, with conspicuous thick
and thin areas. This is particularly striking in the upright
cells on the ends of multiseriate rays. Occasionally some of
the multiseriate rays in Cyrilla broaden on the periphery
external to the cambium. This characteristic is particularly
noticeable in some of the fossil preparations of Cyrilla from
the Brandon Lignite in Vermont (Spackman, 1949).
The xylem parenchyma is relatively abundant and diffuse
in Cyrilla, usually occuring as apotracheal strands. It is
less abundant in Cliftonia, where it is more often found ad-
jacent to vessel elements.
The nodes are unilacunar in all members of the Cyrilla-
ceae. There is some evidence from the external ridges on
the stem that the vascular traces leave the stele approxi-
mately two nodes below the level at which they enter a leaf,
but this has not been verified by anatomical studies.
LEAVES. — The leaves are simple, entire, estipulate, and
moderately to heavily coriaceous. They are highly variable
in size and shape in Cyrilla and Purdiaea, but rather uni-
form in Cliftonia. In a few species of Purdiaea the leaves
are approximately the same size and shape in all of the
specimens seen; but these are poorly collected species, and
future collections will undoubtedly show a much wider range
of variation. The high degree of variation in the leaves of
Cyrilla is discussed in detail below in the section on variation
14 JOAB L. THOMAS
in C. racemiflora. In general, the leaves of Purdiaea range
in length from 1-1.5 em. in P. microphylla, to 11-12 cm. in
P. moaensis; those of Cyrilla range from 2-15 cm. long and
those of Cliftonia range from 3-6 cm. long.
The venation is pinnate in Cyrilla, Cliftonia, and the
South American species of Pwrdiaea. In all other species of
Purdiaea, however, the leaves have one to five pairs of
strongly arcuate, lateral veins which run longitudinally,
more or less parallel to the margin of the leaf. The midvein
is prominent beneath in all members of the Cyrillaceae. In
most species of — i lateral veins are prominent
beneath and p t above; whereas in
Cliftonia the lateral ae ae are > scarcely if at all prominent on
either surface. In Cyrilla there is considerable variation in
the degree of prominence of the lateral veins.
There is a fairly heavy layer of cuticle on both surfaces
of the leaves, particularly in the more coriaceous leaves of
Purdiaea and the leaves ofthe
of Cyrilla. The cuticle on the lower surface i is often irregu-
larly ridged in Cliftonia. Stomata are fairly abundant and
confined to the lower surface in all three genera. The guard
cells are not flanked by any specialized accessory cells, but
by normal epidermal cells. Below the upper epidermis there
are two layers of rather compact, columnar, palisade cells
in Cyrilla and Purdiaea, and one to two layers of more
loosely arranged palisade cells in Cliftonia. According to
Solereder (1899) the mesophyll in Cyrilla and Cliftonia con-
sists solely of palisade tissue, but in all of the preparations
that I have seen (which include samples from the United
States, Cuba, and South America) there is a well-developed,
spongy mesophyll of loosely arranged, more or less isodia-
ene cells, with numerous large air spaces. These air
are particularly large and numerous in leaves of
Cliftonia w which have only a single layer of palisade cells. In
these leaves the mesophyll region occupies approximately
2/3 of the cross-sectional diameter of the leaf. In some
leaves of Cliftonia there is a gradual transition between the
columnar cells of the palisade layer and the more isodia-
metric cells of the mesophyll.
Druses are abundant in the leaves of all three genera.
Solitary crystals have been seen in Purdiaea and Cyrilla and
were reported in Cliftonia by Beauvisage (1920). The vas-
cular bundles in the veins are surrounded by a sheath of
thick-walled sclerenchymatous cells. In the midvein of Cy-
rilla and Cliftonia, and in the midvein and large, lateral
veins of Purdiaea, the metaxylem is composed of cells with
MONOGRAPH OF CYRILLACEAE 15
relatively thick, secondary walls. The tracheary elements
in the smaller veins have spirally-thickened, secondary walls.
There are no stipules in any members of the Cyrillaceae,
but extending over the axillary buds and laterally to each
side there are several bright red, ligulate, glandular struc-
tures. A pair of similar structures is also found at the base
of the bracts, occasionally at the base of the bracteoles, and
rarely even at the base of the sepals. The nature of these
structures is unknown. The similarity between the struc-
tures at the base of the bracts and those in the axils of
leaves may be merely superficial. The position of these
peculiar structures is suggestive of bud scales in the leaves,
but they are more suggestive of vestigial stipules at the
base of the bracts where they always occur in pairs, one on
either side of the bract.
ROOTS. — The roots of Cyrilla and Cliftonia are woody,
much branched, and quite shallow. They are composed of a
diffuse network of branches with no primary or tap root.
The major branches bend abruptly a few inches under the
ground and run horizontally, more or less parallel to the
surface of the ground. They tend to remain parallel to the
surface even when the plants are growing on small mounds
or in depressions. Numerous smaller branches which are
fibrous in texture run both horizontally and vertically into
the soil. Along the length of the horizontal roots numerous
adventitious shoots arise, eee large vegetative clones.
(The nature and d shoots
are discussed in detail below. —
The primary vascular system in the roots of Cyrilla, the
only genus in which the roots have been examined anatomi-
cally, is a triarch protostele (Fig. 19). Cambial activity
begins early, and growth rings are present in all but ex-
angular in cross section, with relatively thin, secondary
walls. The perforation plates are scalariform, consisting of
25-50 bars per end wall. Vessel pitting is transitional to
opposite, the pits being rather small. The imperforate trach-
eary elements consist of both tracheids and fiber-tracheids,
with secondary walls varying from quite thin in the trach-
eids to relatively thick in some of the fiber-tracheids. The
rays are heterogeneous, and may be either multiseriate or
uniseriate. The multiseriate rays are usually only 2-3 cells
wide in younger roots, occasionally becoming 4-5 cells wide
in older roots. The uniseriate rays are 3-10 cells high,
consisting of both procumbent and upright cells. Xylem
16 JOAB L, THOMAS
+]. ‘. } ]
parenchyma is rather scanty and app i
usually occurring in vertical strands which are adjacent to
a ray. The outer layer of the bark of young roots consists
of a layer of spongy, relatively thin-walled cork cells. Just
inside the cork cells there is a layer, several cells thick, of
parenchymatous cells, many of which contain densely-stain-
ing phenolic compounds. Small lenticels occur infrequently
in the roots.
PUBESCENCE. — Although both Cyrilla and Cliftonia are
glabrous, most species of Purdiaea are to some degree pu-
bescent, particularly on the reproductive structures. The
degree of pubescence is highly variable in some species of
Purdiaea, occasionally varying from glabrous to pubescent
in a single species. In Purdiaea cubensis and P. nipensis
this variation occurs within a small geographic area.
Two different types of trichomes have been found in Pur-
diaea. In Purdiaea velutina, P. moaensis, and P. shaferi the
racemes and, in the latter two species, the young stems are
clothed with long, silky, usually appressed trichomes ; where-
as in most of the other species the trichomes are short and
more rigidly erect. The long trichomes have a small, bulbous
base usually composed of 6-8 cells, and a long, unicellular,
non-septate “blade”. The blade is terete, pointed, and rela-
tively thick-walled. The small trichomes do not have a multi-
cellular base, the entire structure being a single, elongate
epidermal cell. They are pointed, thick-walled, and covered
externally with an irregular layer of cutin.
FLORAL MoRPHOLOGY
RACEMES. — The flowers are borne in either terminal
or axillary racemes which vary in length from 2-3 cm. in
diaea microphylla to 25 cm. in some populations of
Cyrilla racemiftora. Flowering proceeds progressively from
the base to the apex of the racemes. The rachis is ridged,
each ridge extending for approximately two nodes and end-
ing at the base of a pedicel. The pedicel is also ridged,
strongly so in Cyrilla, where there are fine, sharp ridges
that extend the full length of the pedicel. Each ridge termi-
BRACTS. — The bracts which subtend the flowers of the
Cyrillaceae are highly variable in size, shape, and texture.
In Cyrilla they are lanceolate, medially thickened, and usu-
MONOGRAPH OF CYRILLACEAE 17
ally claw-shaped, varying in length from 5-30 mm. In Clif-
tonia the bracts are spatulate and slightly concave with
relatively little variation in size. In Purdiaea the bracts, at
maturity, vary in size and shape progressively from the base
to the apex of a raceme. Characteristically, the bracts near
the base of the raceme are long and lanceolate ; those further
up the racemes become increasingly shorter and more del-
toid. The long, basal bracts often attain a maximum length
of more than twice that of the short, deltoid, apical bracts.
There is but one vein in the bracts and bracteoles of
Cyrilla and Cliftonia, whereas those of Purdiaea have sev-
eral prominent, longitudinal veins, and usually a distinctly
visible, reticulate pattern of smaller veins. In Purdiaea and
Cliftonia the bracts are articulate at the point of attachment
to the peduncle; and in the latter genus they are deciduous,
usually shedding well before anthesis. A pair of bracteoles
is borne in varying positions on the pedicel in Cyrilla and
Cliftonia. The bracteoles are of the same texture as the
bracts, and in Cyrilla they are approximately the same shape.
In Cliftonia the bracteoles are usually lanceolate and quite
different from the spatulate bracts. There are no bracteoles
in Purdiaea.
SEPALS.— All members of the Cyrillaceae have, as a rule,
five sepals, although six sepals are rarely encountered in
Cyrilla, and as many as eight have been reported for Clif-
tonia (Sargent, 1892). In Cyrilla and Cliftonia the sepals
are deltoid, usually thickened medially, and are equal in
size. Rarely the sepals are laterally united at the extreme
base in Cyrilla. This phenomenon occurs with greater fre-
quency in some populations than in others, but apparently
it is always sporadic in occurrence, and is without geograph-
ical or ecological significance.
In Purdiaea the sepals are highly specialized and quite
different from those found in Cyrilla and Cliftonia. Sepals
a quincuncial manner. t is, two are exterior, two are
interior, and one has one edge exterior and one interior.
A similar type of sepal insertion is found in the Theaceae,
Clethraceae and Hyperi The qui ial type of sepal
insertion is greatly exaggerated in Purdiaea, with the two
exterior sepals completely enclosing the two interior sepals,
and all but one edge of the middle sepal. The size and shape
18 JOAB L. THOMAS
are often quite different in three of the five sepals, and it is
therefore necessary to describe them individually. They are
numbered for convenience, according to their position in the
spiral. The lowermost of the two exterior sepals is number
1; the other exterior sepal is number 2; the middle sepal
which has one exterior and one interior edge is number 3.
The two interior sepals are usually the same size and shape,
and thus are not numbered, being referred to simply as the
“«nterior sepals”. In a similar way, the first two sepals which
enclose the other three are referred to collectively as the
“exterior sepals’’.
The exterior sepals are approximately equal in length
in Purdiaea stereosepala, P. stenopetala, and P. moaensis.
However, in all of the other species, the first sepal is clearly
longer and usually broader than the second sepal. In fact,
in P. nutans, P. cubensis, and P. nipensis the first sepal is
as much as twice as long as the second sepal at maturity.
The third sepal is usually somewhat asymmetrical. The
edge which extends laterally between the two exterior sepals
is rounded, whereas the other edge which is completely en-
closed by the exterior sepals is flattened. The flattened mar-
gin is usually ciliate, even in species in which the other
margin is glabrous. Also, the surface of the enclosed portion
of the third sepal is often more densely pubescent than the
surface which is exposed. The interior sepals are approxi-
mately equal in size in all species, and are considerably
smaller than the outer three. They are very narrow, lan-
ceolate, and somewhat thickened basally. The ventral sur-
face is adjacent to the ovary, and the sepals are usually
concave, following the shape of the ovary. The interior sepals
are characteristically more densely pubescent than the other
three sepals. Moreover, the interior sepals are always pu-
bescent, even in species in which all other parts are glabrous.
The sepals of all the Cyrillaceae are persistent in fruit.
Those of Purdiaea increase considerably in overall dimen-
sions during the maturation of the fruit, whereas those of
Cyrilla, and Cliftonia attain near maximum size soon after
anthesis. In the latter two genera there is a single, very
small, and obscure vein which extends longitudinally for the
entire length of the sepal. In Purdiaea, there are several,
more or less p inent, longitudinal veins and a well-de-
veloped, reticulate system of smaller veins.
In Cyrilla the sepals are composed of more or less isodia-
metric yma cells. They are 6-7 cells thick near the
middle, gradually tapering laterally to only two cells thick
MONOGRAPH OF CYRILLACEAE 19
along the edge. The dorsal and ventral surfaces are covered
with a thin, irregular layer of cutin. The cutin fills the
spaces between the epidermal cells, and stands out from the
surface in numerous microscopic ridges.
PETALS. — The corolla consists of 5 (rarely 6) petals
which are delicate, and membranaceous. In Cyrilla and
Cliftonia they are white or pinkish-white ; in Purdiaea they
range from deep pink to violet. The venation of the petals
is quite different in the three genera. In Purdiaea there are
numerous small, longitudinal veins which branch repeatedly,
covering the entire petal. In Cliftonia there are one large
central vein and two smaller side veins which extend the
full length of the petal, branching only a few times. In
Cyrilla there is only one vein which runs the full length of
the petal, and usually there are 1-2 pairs of very small lat-
eral veins which arise near the middle of the petal.
The petals of Cyrilla are morphologically quite distinct
from those of the other two genera due to a characteristic
median and basal thickening. This thickened area is com-
posed of fairly thick-walled, glandular cells, filled with
densely-staining compounds. These cells are nectariferous,
secreting a sweet, sticky “honey” which is very attractive
to bees. In this thickened region the petals are 8-10 cells
thick, but lateral to this on either side they taper abruptly
to 3 cells in thickness. There is a thin layer of cutin on
both surfaces of the petals. As in the sepals, the cutin pro-
trudes in numerous microscopic ridges. The outer walls of
the epidermal cells are also cutinized and thickened, more
so on the dorsal than on the ventral surface.
ANDROECIUM. — The stamens of Cliftonia and Purdiaea
are in two whorls with five stamens in each whorl. The
stamens in the outer whorl are opposite the sepals; those
in the inner whorl, opposite the petals. In Cyrilla, only the
outer whorl is present. The stamens of the two whorls are
approximately equal in Purdiaea, but in Cliftonia those of
the outer whorl are considerably longer than those of the
inner whorl. The anthers are versatile in all members of
the Cyrillaceae. The filaments are attached near the middle,
on the dorsal side in Cyrilla and Cliftonia, but on the ventral
side in Purdiaea. The filaments of Cyrilla and Purdiaea are
narrow, terete, and subulate; those of Cliftonia are laterally
expanded and petaloid below the middle, narrowing abruptly
above the middle, becoming terete and subulate. The fila-
ments of all members of the family are vascularized by a
single, longitudinal trace.
20 JOAB L. THOMAS
In Cyrilla and Cliftonia the anthers are ovoid and shed
their pollen by means of longitudinal slits. In Purdiaea, on
the other hand, the anthers are quite different. They are
oblong with a caudate base, and shed their pollen by means
of terminal pores. The filament is folded back on itself in
bud, and the anthers develop in an inverted position. When
the flowers open the filament straightens, rotating the an-
thers through 180 degrees, placing the pores in an apical
position and the cauda in a basal position. It is interesting
to note that in Cyrilla and Cliftonia where the filaments are
attached on the dorsal side of the anthers, the two lobes of
the anthers are free below the point of attachment; whereas
in Purdiaea, where the filaments are attached on the ventral
side, the anther lobes are free above the point of attachment
(after the filaments have straightened at anthesis).
The developmental morphology of the anthers was studied
in detail in Cyrilla only. They are composed of two pairs of
elongate microsporangia. The tissue of the partition between
the two sporangia of each pair remains intact and the spor-
angia do not become confluent. The outer layer of the
anther wall becomes flattened and stretched during the
growth and development of the anther, but the cells usually
remain intact and uniformly adjacent in mature anthers.
ae Sas fibrous bands develop in the cells of the anther
wall.
The endothecium is well-developed, becoming elongate at
ee with conspicuous fibrous bands extending later-
mic in both of these tissues. The tapetum persists more or
= intact through meiosis but rapidly disappears during
devel t of the mi p
2
e sO =
In early stages the archesporial cells are seen as a plate
of tissue, usually 2-3 cells wide and approximately 25 cells
MONOGRAPH OF CYRILLACEAE 21
long in longitudinal section, and 3-4 cells across in trans-
verse section. The primary sporogenous cells undergo occa-
sional divisions, giving rise to a slightly larger number of
microspore mother cells. The divisions of the microspore
mother cells are of the so-called simultaneous type (Mahesh-
wari, 1950). That is, the first and second meiotic divisions
are pleted before cytokinesis begins. With cytokinesis
a tetrad of microspores is formed in the shape of a tetrahe-
dron. The tetrads break up soon after cytokinesis, with
the beginning of exine formation.
POLLEN. — All pollen preparations used in this study were
prepared by acetolysis, and are in the pollen reference col-
lection at Harvard University. A detailed description of the
acetolysis procedure used is given by Traverse (1955).
The pollen of Cyrilla has received considerable attention
since this genus was found to be one of the most abundant
elements in the Brandon Lignite. A brief description of the
pollen of the Cyrillaceae was given by Erdtman (1952), and
a more detailed description of Cyrilla pollen by Traverse
(1955).
His description of the pollen of Cyrilla racemiflora is as
follows:
Tricolporate and tetracolporate pollen grains with very
wide longitudinal furrows, which are seen in polar view to
narrow toward the poles, producing a Y-shape. As seen at
polar view there is a characteristic overhang of exine at the
edge of the furrow . . . Transverse furrows elliptical, con-
stricted at the contact with the longitudinal furrow. Margos
and bow-shaped longitudinal costae very prominent. Psilate
sculpture. Size ca. 25 micra. Thickness of exine: ca. 2.7 mic-
ra.
The pollen grains of Purdiaea and particularly those of
22 JOAB L. THOMAS
Fic. 10-16. Pollen grains of the Cyrilla Fig. 10. Cliftonia monophylla, polar
view, mid-central focu: 1, Cliftonia cnaekeae high focus of the same grain
shown in Fi rilla racemifiora, polar view, mid-central foeus. Fig.
ig. 10. Fig. C
13. Cli i monophylla, equatorial view, cane foeus. Fig. 14. Cyrilla racemi-
flora, equatorial view, mid-central oe Fi, Purdiaea nutans, polar view, id-
cent a Fig. 16. Purdiaea nutans, passed view, mid-central foeus. Fig.
10-16, ia iibaiinn oar 1000 x.
MONOGRAPH OF CYRILLACEAE 23
era, being quite fine and reticulate, visible only under very
high magnification. The separation of Cyrilla and Cliftonia
on the basis of pollen is much more difficult. The pollen
grains of Cyrilla are generally more rounded and have a
thinner ektexine though a thicker total exine. They have
a slightly more prominent exine sculpturing. Also the exine
usually overhangs the edge of the furrow to a greater
extent in Cyrilla than in Cliftonia, These distinguishing
characteristics are all variable, and the two genera occasi-
onally overlap. However, on the basis of a combination of
these characters more than 90% of the pollen grains of these
two genera can be distinguished with accuracy.
The close similarity of the pollen grains of all members
of the Cyrillaceae is of considerable significance, particu-
larly in view of the fact that the genera are so distinct, and,
indeed, quite dissimilar in several characteristics. There is
some evidence from the fossil record that the morphology of
the pollen of this group has remained rather constant. In
the Magothy Formation of the Upper Cretaceous (dis-
cussed below) several grains of cyrillaceous pollen have
been found. These grains are very similar to those of
Cyrilla and Cliftonia, strikingly so considering the age of
the deposit.
Pollen studies have also added further evidence to a close
relationship between the Cyrillaceae and the Ericales. The
only group which has pollen that closely resembles that of
the Cyrillaceae is the genus Clethra. The pollen of Clethra
is larger than that of the Cyrillaceae, but the grains are
similar in several characteristics. Possibly the pollen of the
Ericaceae and the Pyrolaceae would show a closer resem-
blance to that of the Cyrillaceae were it not for the fact that
in these families the pollen remains in terads.
In the Aquifoliaceae and the Celastraceae, on the other
hand, the pollen is quite unlike that of the Cyrillaceae as
was pointed out by Erdtman (1952). An examination of
the pollen of Cyrillopsis has added further evidence for ex-
cluding this genus from the Cyrillaceae. The pollen of
Cyrillopsis is similar to that found in some members of the
Celastraceae, but quite unlike that of the Cyrillaceae.
GYNOECIUM. — The ovary is 2-5-locular with pendulous
ovules, one in each locule of Cliftonia and Purdiaea, three in
each locule of Cyrilla. The ovules are anatropous in Cyrilla
and Cliftonia, but in Purdiaea they are attached to the funi-
culus in an orthotropous manner, and could thus be des-
cribed as pendul thotrop In all of the Cyrillaceae
24 JOAB L, THOMAS
there is a central column of vascular tissue which extends
longitudinally from the receptacle to near the apex of the
ovary. Copeland (1953) has shown in his illustration of
Cyrilla and Cliftonia that this vascular bundle divides near
the apex of the ovary with a single trace extending into each
locule. A similar type of development is also found in Pur-
diaea. In Cliftonia and Purdiaea the funiculus is attached
directly to a small column of tissue surrounding this trace.
In Cyrilla the trace passes into a short placental arm which
branches twice to give rise to three ovules per locule.
There is a column of elongate parenchyma cells extending
from the ovule to the apex of the style. According to Cope-
land (1e.) this column of cells broadens at the apex in
Cyrilla and Cliftonia forming the stigmatic surface. In
Purdiaea there is a single, elongate style which is not lobed
at the apex. The style is composed of a thin layer of epider-
mal cells enclosing 4-5 columns of elongate, parenchyma
cells, similar to those leading from the ovule to the stigma
in Cyrilla. These columns do not fuse; and if the epidermal
layer is removed they readily separate into discrete intact
strands, each leading from the apex of the style to an ovule.
In the ovules of Cyrilla and Cliftonia there is a single
integument, and a thin, short-lived nucellus. The nucellus
usually disappears shortly after tetrad formation during
megasporogenesis. Copeland predicted the existence of a
tenuinucellate condition in this group although his material
was too old to show any nucellus. In later stages of megas-
porogenesis, there is a fairly well-developed integumentary
tapetum.
The embryo sac begins development relatively early. An
expanding tetrad of megaspores is usually present before
the microspore mother cells of the same bud have begun to
enlarge prior to meiosis. Subsequent development in the
embryo sac is slower, however, and the male and female
gametophytes mature at approximately the same time in a
given flower. The mature embryo sac is 8-nucleate, os
one of the antipodals is occasionally either difficult to discern
. cc This phenomenon was also noted by Copeland
Intermediate stages of seed development have not been
seen, but the mature seed is apparently devoid of a seed coat.
Planchon (1846) practically described this situation i in Pur-
— of the seed he says: integumento tenuissimo
albicante, albumini carnoso adhaerente .
MONOGRAPH OF CYRILLACEAE 25
Similarly Gilg (1896) describes the seed of the Cyrillaceae
as having seed coats not clearly differentiated (“ . . .un-
deutlich abgegliederte S hale. . .”). Copeland inter-
preted the fruit of Cyrilla as a 2-seeded caryopsis, thus
intimating that the seed coat is fused to the ovary wall.
However, in fruits in which no seeds develop (partheno-
carpy is of common occurrence in the Cyrillaceae) the struc-
ture of the ovary wall is the same as it is in fruits which
do contain seeds. There is no evidence in the latter of any
extra layer which could be interpreted as a seed coat. The
cells of the outer layer of the endosperm are somewhat
differentiated, but this layer is composed entirely of thin-
walled cells and is clearly a part of the cellular endosperm
and not a seed coat. This unusual characteristic of seeds
without a seed coat is found in all members of the Cyrillaceae
and serves as a reliable character bv which this family can
be distinguished from related families.
CYTOLOGY
metaphase II configurations were found in which the chrom-
osomes could be counted with accuracy. The number that
was found in most cells was n = 20. Unequal distributions
were apparently quite frequent in this collection, however,
for numbers ranging from 18 to 22 were obtained in several
cells. In one pair of daughter cells, one cell contained 19
chromosomes and the other, 21 chromosomes. Several bridge
formations were observed and there was considerable vari-
ation in the size of the chromosomes. Possibly the discrep-
ancy in chromosome number was due partially to chromo-
some fragments rather than to unequal divisions.
More cytological work is badly needed in this group, not
only on Cyrilla but also on Cliftonia and Purdiaea which
have never been studied cytologically. On the basis of my
experience with preserved material, it seems advisable that
cytological work in this group be attempted only if living
material is readily available.
ADVENTITIOUS SHOOTS
Reproduction in both Cyrilla and Cliftonia is primarily
26 JOAB L. THOMAS
by means of retofios*, that is, vegetative sprouts from hori-
zontally positioned, underground roots. As discussed above,
the root systems of Cyrilla and Cliftonia are shallow, and
the major branches grow in a horizontal direction, parallel
to the surface of the ground. Along the length of these
horizontal roots numerous retofios arise. These develop un-
derground and, as they emerge from the soil, resemble nor-
mal seedlings.
Propagation by means of retofios accounts for the vast
majority of individuals or apparent individuals of Cyrilla
and Cliftonia throughout the range of these genera. Hun-
dreds of shrubs and small trees of Cyrilla have been exam-
ined in populations along the coastal plain of southeastern
United States from Virginia to eastern Texas, and in the
Provinces of Pinar del Rio and Oriente in Cuba. In every
case, exposure of the root systems revealed that the plants
were interconnected with several and often numerous other
“individuals” near by. Thus, what appears superficially to
be a population composed of separate individuals is usually
a large clone.
The total area covered by one of these clones has been
found, in many instances, to be quite extensive. For example,
in southeastern Virginia, near the North Carolina border
(Thomas 458), I excavated an area approximately 15 feet
wide and 30 feet long, containing over 50 bushes of Cyrilla,
and found that all of these bushes were part of a single clone.
In numerous other areas, by tracing the root systems from
one bush to the next, I have found clones which extended
for even greater distances.
three or more inches below the first branch. The entire sys-
tem was still int ted, thus indicating that the plants
in a clone remain interconnected for several years.
It is difficult, on the other hand, to determine the parent
plant of a clone. In a situation such as the one described
above, in which the clone is large and contains several large
used by the Mexicans to denote sprouts from the roots as
by
Due to the awkwardness of the English equivalent, the term retofio will be used
here to refer to this phenomenon ; that is, the situation in which vegetative shoots orig-
inate from underground roots.
MONOGRAPH OF CYRILLACEAE 27
shrubs of approximately the same size, it is virtually impos-
sible to select the parent plant. In certain areas one can
find small clones in which there is a single, large bush and
a number of smaller ones more or less, radiating from it.
Clones such as this are exceptional, however, and even in
these one cannot be certain that the supposed parent plant
did not originate from the roots of other bushes that once
grew in the area. Numerous clones were found which seemed
to be composed entirely of young retofios, but these could
usually be traced to old stumps in the vicinity. This is
common along highway right-of-ways that are clear-cut
periodically.
In some plant groups the formation of retofios is related
to disturbance or injury. In Guayule (Parthenium argenta-
tum), for example, Lloyd (1911) has shown that there is a
direct correlation between disturbance and the formation of
retofios. The ratio of retofios to seedlings was found to vary
greatly with the habitat, although both seedlings and ret-
ofios can be found in practically any habitat in which the
species grows. On stony slopes, where there is evidence of
disturbance, the retofios were found to be more abundant
than the seedlings, whereas on more level and less disturbed
soil just the reverse was true. Apparently, in Guayule the
so-called normal retofios are formed only when the end or
a portion of the root is exposed. “Induced retofios”, on the
other hand, can be stimulated to develop by injuring the
root system or by severing it from the plant. Also, in
Guayule the retofios become separate and independent soon
after they emerge from the root of the parent plant. As the
retofio develops, the proximal part of the root connecting
it with the parent plant undergoes little or no further
growth and soon dies. The distal portion of the root con-
tinues to grow and forms the equivalent of a tap root for
the developing retofio. The retofio, however, is distinguish-
able from a seedling by the shape of the taproot and its
position in the soil (Muller, 1946).
In Cyrilla there is no apparent correlation between dis-
turbance or injury and the formation of retofios. They form
regularly on un roots 3 to 8 inches underground in
unusually abundant in cut-over areas, but this is more likely
to be a result of the ability to colonize rapidly (by means
28 JOAB L. THOMAS
of vegetative sprouts) than a result of disturbance caused
by the cutting.
One of the most striking examples of rapid colonization
by Cyrilla of a cut-over area was seen in the Sierra de Moa
in Oriente Province, Cuba. A large part of the vegetation
of this area was second growth following extensive lumber-
ing operations which were terminated approximately fifteen
years ago. One of the dominant elements of this second
growth, particularly on the moist sites, was Cyrilla. One
area near the town of Moa was of particular interest in
this connection. Ona rather gentle slope, 350-400 feet above
sea level, an area of roughly 500 acres showed signs of
having once supported a sizeable pine forest, as evidenced
by numerous tree stumps. However, in the summer of 1957,
the area was covered by a fairly dense growth of Cyrilla.
An examination of several of these plants in different parts
of the region indicated that the colonization by Cyrilla had
been the result of vegetative propagation. Moreover, the
presence of hundreds of young retofios indicated that the
colony was still expanding rapidly. There was little evidence
of recent disturbance in the area and the young retofios
were developing from unexposed roots. There was some
evidence of erosion, but the area appeared to have been more
or less stable for several years, and colonization by vegeta-
tive means was continuing.
ANATOMY OF DEVELOPING RETONOS.— The development
of vegetative shoots from roots is a fairly common phenom-
enon, especially among the dicotyledons, and has been rec-
ognized for several centuries. It was first pointed out by
Hieronymus Tragus in 1546 in ovat ae pe arvensis. Since
then the number of species d by
this method has continued to i increase. she “(1857) listed
42 species, Warming ( 1877) 87 species. and Wittrock (1884)
132 species which The most compre-
hensive treatment of this subiect was “made by Beijerinck
(1887) who referred to 246 different genera in which one
or more species are known to propagate by means of root
sprouts. In many of these, however, the anatomy and devel-
opmental morphology have not received detailed study.
The present study of the origin and early development
of adventitious buds within the roots was made on material
of Cyrilia racemiflora collected in Virginia and North Caro-
lina. Several roots bearing young retofios were collected
in late May and early June, and fixed and stored in Formalin-
Acetic Acid-Alcohol. The material was softened for four to
MONOGRAPH OF CYRILLACEAE 29
Anatomy of
ze in the develor
of the bud.
30 JOAB L. THOMAS
five weeks in concentrated hydrofluoric acid, embedded in
paraffin, and sectioned with a rotary microtome. The sec-
tions were cut at a thickness of ten micra, and stained in
safranin and fast green.
In sectioning roots from which very young retofios were
emerging, I was fortunate enough to pass through several
buds in various stages of development. An examination of
these buds added a great deal of valuable evidence, particu-
larly in interpreting early stages in the ontogeny of the
adventitious buds. In early stages the bud consists of a
rounded cluster of small, densely cytoplasmic, meristematic
cells. A recognizable pith of darkly-staining, tanniniferous
cells develops during the early stages of bud formation. At
the earliest stages seen the bud is located in the region of
the phellogen or cork cambium and apparently originated
either directly from the phellogen or from recent derivatives
of the phellogen. Thus, the origin of the adventitious bud
yaust be exogenous. (A median section of the bud in an
early stages of development is shown in Fig. 17, and an
enlargement of the same bud is shown in Fig. 18).
The evidence that adventitious buds continue to originate
in roots that have undergone two or cee seasons aa cam-
bial activity is a further indicati
of the buds. The pericycle is procera soon after the. initi.
ation of cambial activity, thus eliminating the possibility
of any subsequent buds originating endogenously from the
pericycle.
Adventitious buds have not been seen in roots lacking
secondary tissues, and therefore it is not known whether
buds ever develop in the primary body of the root. Since all
the young buds that have been seen apparently originated in
the vicinity of the phellogen, it is conceivable that adventi-
tious buds are pein only after the development of a sec-
ondary dermal s
During the me bsp of bud development a noticeable
swelling appears on the root, external to the bud (Fig. 17).
This swelling becomes more extreme in later stages, as cells
derived from the bud begin to elongate at right angles to
the long axis of the secondary xylem in the root (Fig. 21).
In still later stages radial rows of parenchymatous cells can
be seen extending laterally on each side of the developing
shoot (Fig. 20). These rows of parenchyma cells are not
derivatives of the bud, but develop from the cambium of the
root, leaving a gap in the stele.
An unusual aspect of the development of these retofios
MONOGRAPH OF CYRILLACEAE 31
is the long period of time between the initiation of the bud
and its emergence from the root. In every case that has
been examined the bud has remained within the root for
three or more years before emerging. During this three-or
four-year period the developing shoot increases considerably
in size. The rate of growth in diameter of the bud increases
as it grows outward, and as a result the developing shoot
becomes somewhat obconical in shape (Figs. 19 and 20).
During the first three or four years of development the
rate of growth in the shoot is just sufficient to keep pace
with the growth of the secondary body of the parent root.
As the bud increases in cross-sectional diameter, procambial
cells begin to differentiate basipetally, eventually establish-
ing vascular connections with the stele of the root. (An
intermediate stage in this process is shown in Fig. 22). The
establishment of vascular connection and the emergence of
the adventitious shoot from the root occur at approximately
the same time. It seems likely that the shoot must establish
these connections before it can emerge from the root.
By the time the young shoot has emerged it already has
a rather large cross-sectional area of procambial cells that
are blocked out, and a relatively large area of differentiated
primary xylem. The differentiation of the primary xylem
and the development of a cambium is a rapid process in the
young shoot once it has emerged. There is also a deflection
of some secondary xylem from the root into the shoot. Fig.
20 shows a radial section through the base of a young shoot
collected during the middle of the growing season of the
year that it emerged from the root. This photomicrograph
shows the deflection of secondary xylem from the root into
the shoot and also the well-developed primary xylem in the
shoot. It also shows that the tissues of the young retonos
can be traced well into the stele of the root. A transverse
section of a root bearing a shoot at approximately the same
stage of development as the preceding one is shown in Fig.
1
Again, the tissues of the shoot can be traced well into the
stele of the root, and the age of the retofio is indicated by
the number of growth rings in the root. The tissues of the
retofio can be traced through three or possibly four growth
rings in the root. This bud must have originated soon after
the initiation of cambial activity. There are, however, defi-
nite radial rows of secondary xylem in the root, internal to
the tissues of the developing shoot.
Occasionally two adventitious buds are found in close
32 JOAB L. THOMAS
proximity. Fig. 22 shows a transverse section through a
root containing two buds developing quite close together.
(Although one of the buds cannot be seen at this level it
became visible in later sections, approximately 70 micra
deeper at the position indicated by an arrow). These two
buds apparently originated at approximately the same time
and developed at the same rate. This simultaneous develop-
ment of two buds in close proximity may be of significance
in later studies, particularly in connection with the possib-
ility of hormonal control of the development of these adven-
titious buds.
In summary, the retofios of Cyrilla originate exogenously
as adventitious buds in the region of the phellogen, develop
at a rate that is just sufficient to keep pace with the sec-
ondary growth of the parent root during a period of three
to four years, and finally emerge as a vegetative shoot. The
factor, or factors, which controls the origin and development
of these buds is not known. On the basis of this limited
anatomical study it seems that the buds must establish
vascular connections with the stele of the parent root before
they emerge from the root. Further studies are needed on
the origin and development of retofios, including compara-
tive anatomical studies on other groups, and particularly
morphogenetic studies to determine the nature and source
of the stimulus that initiates the buds, and the possibility
of hormonal control of their sub t devel t
1S
PHYLOGENY
In the history of phylogenetic classifications involving
the Cyrillaceae most authors have considered the relation-
ships of the family to be with either the Ericales or the
Celastrales, and it has been shifted back and forth between
these two groups many times. Jussieu (1789), the first to
attempt a natural or phylogenetic classification of Cyrilla,
included it in his Ericae. Lindley (1836) placed Cyrilla and
a second genus, Cliftonia, which had been described after
Jussieu’s treatment, in the Celastraceae. Torrey and Gray
) also included these two genera in the Celastraceae,
but added a note saying that the relationships seemed closer
to the Ericaceae and that these genera probably should be
considered as a sub-group of that family. Endlicher (1840)
included the Cyrilleae in his supplement, indicating that
they were related to the Ericaceae. The following year,
however, he changed their position and included the Cyrilleae
asa up within his Ilicineae (Endlicher, 1841). Five
years later Planchon (1846) described the genus Purdiaea,
MONOGRAPH OF CYRILLACEAE 33
stating that it was closely related to Cliftonia and Cyrilla,
but also pointed to relationships with Clethra, Pyrola, and,
oddly enough, Sarracenia. In his classification, however,
Planchon placed Purdiaea and the ericaceous genus Eltiottia
along with Cyrilla and Cliftonia in his Cyrilleae, which he
included in the Ericaceae. This treatment of the Cyrilleae
as a sub-group of the Ericaceae was followed by Walpers
(1847) and Agardh (1858). Even Lindley (1847), who was
the first to include Cyrilla and Cliftonia in the Celastraceae,
revised his treatment and recognized the Cyrillaceae as a
separate family related to the Olacaceae and the Ericaceae.
Chapman (1860) also gave family status to the Cyrillaceae
and placed it near the Aquifoliaceae and the Styracaceae, but
in his treatment these familes were also placed near the
Ericaceae. Bentham and Hooker (1876) placed the family
Cyrillaceae near the Ilicineae, but also emphasized a rela-
tionship with the Olacaceae, indicating further that the
apparent relationship between Purdiaea and Clethra was
merely a superficial resemblance. This opinion was base
primarily on the work of Decaisne.
With very few exceptions this treatment of Bentham and
Hooker has been followed in the major botanical works up
to the present day. The following are among those whose
treatment of this group essentially agrees with that of
Bentham and Hooker: Durand (1888) ; Gilg, in Engler and
Prantl (1892); Mohr (1901); Dalla Torre and Harms
(1900-1907) ; Brown (1901); Small (1903, 1913, 1933) ;
Bessey (1915) ; Wettstein (1935) ; Mattick (1935) ; Uphof
in Engler and Prantl (1942) ; Fernald (1950) and Gleason
(1952). One interesting exception to the above is the work
of Baillon (1892), in which he separated the Cyrillaceae
from the Ericaceae, but left the genus Purdiaea in the latter
family. This is one of the few times in which one of the
three genera of the Cyrillaceae has been treated in a family
separate from the other two.
The more recent workers concerned with the Cyrillaceae,
however, have pointed to further evidence of the relationship
between the Cyrillaceae and the Ericaceae and related fam-
ilies. Marie-Victorin (1948) was of the opinion that the
present family is clearly related to the Ericales, and that
the problem at hand is primarily that of the position of the
Cyrillaceae within the Ericales. Copeland (1953) presented
further evidence from comparative morphology, particularly
of the reproductive parts, in favor of a close relationship
between the Cyrillaceae and Ericales, although he stated that
34 JOAB L, THOMAS
this conclusion did not necessarily negate the former asso-
ciation of the Cyrillaceae with the Aquifoliaceae. He also
cited the work of Heimsch (1942) on the comparative anat-
omy of the Terebinthales and Gruinales of Wettstein (1935).
Heimsch concluded, on the basis of wood anatomy, that
the Cyrillaceae did not belong in this group. The wood of
the Cyrillaceae is more primitive than that of the other
groups in Wettstein’s Terebinthales and Gruinales, but is
in many respects similar to the wood of the Ericales. As
further aspects of the total biology of the Cyrillaceae have
been studied, the evidence for a close relationship with the
Ericales has become increasingly strong, and in my opinion
the Cyrillaceae should be included in that order. The follow-
ing is a summary of the anatomical and morphological fea-
tures of the Cyrillaceae which are also found in the Ericales:
primitive wood structure with small vessels having oblique
end walls and scalariform perforation plates; rays uniseri-
ate and multiseriate, heterogeneous ; imperforate tracheary
elements primarily fiber apotra-
cheal; crystals and tannin deposits common; unilacunar
nodes ; 5-angled pith; leaves alternate, entire, simple, coria-
ceous, and estipulate; flowers in axillary or terminal
racemes, flower parts in 5’s; sepals inserted in a quincuncial
manner, persistent; anthers inverted in bud, dehiscing by
terminal pores; glandular nectariferous disk at the base of
the ovary; ovary 2-5-loculed; ovules with a single integu-
ment ; pollen grains tricolporate, with wide longitudinal fur-
rows, longitudinal costae, psilate sculpturing, and with the
exine overhanging the furrow margins.
PALEOBOTANICAL HISTORY
The Cyrillaceae has a rather lengthy paleobotanical his-
tory. Cyrillaceous pollen grains have been found in the
Upper Cretaceous Magothy formation from a well drilling
near Brookhaven, Long Island (Steeves, 1959). These pollen
grains are not assignable to any of the living representatives
of the Cyrillaceae, but in view of the age of the deposit this
is not surprising. However, the grains are very similar to
those of Cyrilla and Cliftonia, and definitely belong to the
Cyrillaceae. In some characteristics these ancient pollen
grains appear intermediate between those of Cyrilla and
Clifton‘a and possibly represent a stage in the history of
family before these two gene. Se RAEN distinct.
The most th 5 ae ted paleontological record
of this family is the occurrence of Cyrilla in the flora of the
MONOGRAPH OF CYRILLACEAE 35
Brandon Lignite of Vermont. This rich, fossil flora consists
of numerous fruits, seeds, fragments of leaves, an abundance
of pollen and spores, and a large amount of well-preserved
wood including both stems and roots. The Brandon Lignite
is one of very few deposits of Tertiary age in northeastern
North America (Barghoorn and Spackman, 1949). For this
reason our knowledge of the flora of the Northeast during
this period is rather meagre, and the flora of the Brandon
Lignite is highly significant.
The composition of this flora long remained a mystery,
and until 1949 only three of the different woods had been
tentatively identified. These were — as Pityoxylon
wliton, Laurinoxylon
Grendonaai Jeffrey and Chrysler, pee a third very doubt-
ful member which was tentatively placed in the genus
“Betuloxylon”. A most important clue to a real understand-
ing of the nature of this flora was the discovery by Barg-
hoorn and Spackman (1949) that the most abundant wood
in the deposit, the puzzling “Betuloxylon”, was actually the
wood of Cyrilla. With this discovery, other elements began
to fall into place. The Pityorylon of Knowlton was found to
a member of the genus Gordonia in the Theaceae, and
the Laurinoxylon brandonianum of Jeffrey and Chrysler was
tentatively assigned to the genus Persea in the Lauraceae.
These three elements can be found growing together at the
present time along the margins of acid swamps on the
Coastal Plain of southeastern United States.
Further evidence of the occurrence of these plants in the
Brandon Lignite was presented by Traverse (1955) in a
study of the fossil pollen, spores, and other micro-fossils of
the deposit. Pollen of both Cyrilla and Gordonia was found
to be very abundant, that of Cyrilla being second only to
ercus in order of abundance in the deposit. The pollen of
Persea, like that of other members of the Lauraceae, does
not preserve, and thus does not occur in the deposit. How-
ever, fossil cupules were found which apparently belong to
that genus. :
The identification of the Cyrilla wood in the deposit was
rather difficult, due primarily to the unspecialized nature
of the wood anatomy of that genus. A reliable identification
was possible, however, due in part to the great abundance
of the wood of es in the deposit (49.8%). Because of
ras aban Leahl well-preserved wood, ia
tively good asa iris were obtainable. These showe
several as of the anatomy including radial, tangential,
36 JOAB L. THOMAS
and transverse, nodal sections of the stem. Detailed studies
of these sections, and comparisons of them with equivalent
sections of living material of Cyrilla were made by Barg-
hoorn and Spackman (1949), and were illustrated by Spack-
man (1949). Though not identical in every detail, the living
and fossil material were found to be remarkably similar.
And in view of the age of the deposit, estimated by the above
authors to be Upper Oligocene, the close similarity between
the two is indeed remarkable.
One extremely puzzling feature of the Brandon deposit is
the absence of fruiting material of Cyrilla. Throughout its
present range, Cyrilla is found to produce abundant fruits
each year. Even a cultivated bush in the Arnold Arboretum
in Jamaica Plain, Massachusetts, approximately 600 miles
north of the present range of the genus, produces abundant
fruits each year. Yet no fruits of Cyrilla have been found
in the Brandon Lignite.
One explanation for this may be related to the structure
of the fruit and the seed. In Cyrilla the ovary wall of the
mature fruit is the functional seed coat. The seed itself is
composed of an elongate embryo, surrounded by a mass of
cellular endosperm, but with no seed coat. Moreover, germ-
ination of the seed involves a breakdown of the ovary wail
which occurs readily under moist conditions. This would
tend to lessen the chances of fruits being preserved. Never-
srg if kiaghoer fruits were produced at or near the site
tly as they are produced throughout
oh range of the genus today, one would expect some fruits
to have been preserved in the deposit.
The determination of the age of this deposit has also
been a rather difficult matter. The stratigraphy of the
entire vicinity of the Brandon formation is very confus-
ing, and gives little insight into the age of the deposit.
The conclusions as to the age of the Brandon Lignite were
eee primarily on the degree of “modernization” of the
technique consists of comparing the fossil flora
ai ‘the living flora of the same area, and plotting the per-
centages of exotic or extinct taxa in the fossil flora against
geologic time. Reid (1920) employed this technique in
studying the Pliocene floras of Europe. His plot yielded a
curve which showed a range from over 90% exotic species
at the beginning of the Pliocene to less than 10% at the end
of this period. Similarly, _ ootageip (1951) fee developed
; curve for the Tertiary floras of North America, and a
certain amount of control was added by including several
MONOGRAPH OF CYRILLACEAE 37
floras for which fairly reliable dates were available from
other evidence. On the basis of this curve the Brandon Lig-
nite was placed in the latter part of the Upper Oligocene.
The Barghoorn curve also shows a sharp rise during
the Oligocene which correlates with the climatic deter-
ioration during the Tertiary. Thus at the time of depos-
ition of the Brandon Lignite a rapid “modernization” of
the flora of northeastern North America had already begun.
On the basis of present-day distributions of the var-
ious elements found in the Brandon Lignite, it would seem
that this flora was decidedly warm temperate or sub-tropi-
cal. The above authors have designated the general region
of the Coastal Plain of southern Georgia and Florida as
the nearest present-day equivalent of the climatic and eco-
logical conditions that must have characterized the Bran-
don area at the time of the deposit. The extreme abund-
ance of Cyrilla wood in the formation indicates that there
must have been fairly extensive, shallow swamps bordered
by Cyrilla thickets, comparable to the Cyrilla thickets com-
monly found on the Coastal Plain of southern Georgia and
northern Florida today. The abundance of roots as well
as stems in the deposit further indicates that these thick-
ets probably consisted of large, vegetative clones of Cyril-
la, as is true of the Cyrilla thickets at the present time.
Thus, there is some evidence that the capacity of vegeta-
tive Bi giaeccnes may have developed quite early in this
genu:
The pollen of Cyrilla in the deposit was found by Trav-
erse (1955) to be rather variable, particularly in size. He
states that the range of variation is conceivable within a
single species of Cyrilla, particularly when one takes into
consideration that anthers containing immature pollen are
commonly found in the deposit. It is possible, neverthe-
less, that more than one species of Cyrilla was present.
In fact, on the basis of this variation, Traverse described
a new species, Cyrilla barghoorniana. Although the genus,
as it is represented in the present-day flora, is composed
of a single species, it is quite conceivable that there were
two or more species of Cyrilla present during the Upper
Oligocene. It seems equally plausible, however, to assume
that there was then, as now, a single variable species.
In this connection the paleobotanical history of Cyril-
la seems to have some bearing on the lack of speciation in
this genus. The effects of the climatic deterioration dur-
ing late Tertiary on plants such as Cyrilla is in striking
38 JOAB L, THOMAS
contrast to its effects on some of the more northern ele-
ments that were not as sensitive to low temperature. Some
of these latter examples remained north as the climate
became colder. Various races and species were destroyed,
divided, or isolated by the subsequent glaciation. Many of
these divided populations were apparently able to survive
in refugia for long periods of time, during which they were
geographically isolated from other such populations of the
same species. Moreover, environmental conditions were
obviously changing, giving rise to various differences in
selection pressure toward gene combinations that were best
adapted for survival under conditions prevalent in a given
refugium. These conditions, which are thought to be favor-
able for speciation, were in large part avoided by the more
subtropical elements which would tend to retreat early in
the face of a cooling climate.
The indication that vegetative reproduction may have
developed quite early in the history of Cyrilla could also be
an important factor in interpreting the evolutionary history
of this group. One can only speculate as to what effects
may have had on Cyrilla in the
past, but in view of the effects of this phenomenon on the
group at the present time it seems probable that vegeta-
tive pence has played an important role in the history
of the genus. One of the effects of a reduced rate of gen-
etic recombination will be a tendency toward a more static
group in terms of evolution. The clear evidence that the
wood anatomy and pollen morphology of Cyrilla have un-
dergone so little change since the Upper Oligocene in-
dicates that this group has been rather static for several
million years. The term static is not intended to imply
genetic homogeneity or complete evolutionary stagnation.
So long as the capacity for some genetic recombination is
retained, the group will have some degree of evolutionary
potentiality. On the other hand, the evolutionary history of
such a group should be quite different from one in which
there is a rapid rate of sexual union and recombination. The
latter situation would more likely give rise to a dynamic,
rapidly-evolving group.
GEOGRAPHIC DISTRIBUTION
The geographic distribution of the Cyrillaceae is totally
within the New World. Cyrilla racemiflora, by far the most
wide-spread species, occurs over an area which includes most
of the range of the entire family. This species is distributed
as follows (Map 11) : along the Coastal Plain of southeastern
MONOGRAPH OF CYRILLACEAE 39
United States; in Oaxaca, Mexico and British Honduras; in
the Greater and Lesser Antilles and Trinidad; in British
Guiana, Venezuela, northern Brazil, and western Colombia.
There is one species of Purdiaea which occurs in Venezuela,
Colombia, and Peru; and ten species which occur only in
Cuba and the Isle of Pines. The third genus, Cliftonia, is
quite narrowly distributed, occurring only on the Coastal
Plain of Georgia, Florida, Alabama, and Mississippi. Thus
the family is distributed in the sub-tropical or warm-temper-
ate areas surrounding the Caribbean Sea and the Gulf of
Mexico, with northward extentions on the Coastal Plain of
southeastern United States, and southward extensions on the
mountains of northern South America.
There is little evidence, however, on which a definite center
of origin for the group can be postulated. The fossil record of
the Cyrillaceae indicates that the group is a very old one, and
that it has had representatives in North America since the
Upper Cretaceous. The occurrence of Cyrilla pollen and wood
in deposits of Oligocene age in Brandon, Vermont, indicates
that this genus has been distinct for several million years,
and that it was once distributed considerably north of its
present range. Of the three genera in the Cyrillaceae, Pur-
diaea shows the highest degree of morphological similarity
to the Ericales, and seems to be the most primitive. In Pur-
diaea there are fairly clear lines of specialization which seem
to indicate that the one South American species is more pri-
mitive than the Cuban species. Thus a very tenuous thread of
evidence suggests that the group may have originated in
South America, but this is certainly in the realm of specula-
tion.
However, the center of diversity of the group at the pre-
sent time seems to be in eastern Cuba. Nine of the eleven
known species of Purdiaea are endemic to Oriente Province,
and in this same area one finds the highest degree of local
variation in Cyrilla racemiflora. eis
According to Croizat (1952) the Cyrillaceae did not origi-
nate in the New World, but rather came from South Africa.
The evidence for this view is apparently based on a sup-
posed relationship between the Cyrillaceae and two genera
of uncertain affinities, Crypteronia and Heteropyzxis (treat-
ed as the Crypteroniaceae and the Heteropyxidaceae by
the above author). Croizat contends that these three groups
are surviving relicts of a larger group which originated in
South Africa and was subsequently divided into three major
parts as follows : the Cyrillaceae migrated to the West Indies,
40 JOAB L. THOMAS
the Crypteroniaceae migrated to Malaysia, and the Hetero-
pyxidaceae remained in Africa. In the words of the above
author: “We believe, then, that Cyrillaceae, Crypteronia-
ceae, and Heteropyxidaceae belong to the same archetypal
node, or ‘genorheitron’, and that the mainsprings of their
dispersal are to be accounted for by migrations effected to
Malaysia and a now crumbled range in West Indies from
South Africa. Migrations of the sort are conventional.”
So far as I can determine, the relationship between the
Cyrillaceae and the two taxa mentioned above is, at best, a
superficial resemblance. These groups have flowers borne in
slender racemes with the flower parts in fives. The fruits
are similar also, being small, globose, and with a persist-
ent style. A more critical examination of Crypteronia, how-
ever, reveals that the flowers and fruits of this group are
quite unlike those of the Cyrillaceae. Moreover, the leaves
of Crypteronia are opposite, whereas those of the Cyril-
laceae are alternate. I have not seen material of Heteropyzis,
but the description of this genus does not indicate a close
relationship with the Cyrillaceae.
In view of the well-known relationship between the flora
of eastern United States and eastern Asia, particularly in
regard to the more primitive woody taxa, it would not be
surprising if fossil or even living representatives of the
'yrillaceae were found in eastern Asia. At present, however,
there is no evidence that members of the Cyrillaceae have
ever occurred in the Old World. ~ .
ASEXUAL REPRODUCTION AND VARIATION
IN CYRILLA RACEMIFLORA
variation from one population to the next, though often
striking from the point of view of the extremes, is usually
MONOGRAPH OF CYRILLACEAE 41
graded, and characteristically there are numerous interme-
diate forms. Different ecological variants show very clear
preferences for a certain type of environment, but this is
obvious in the extreme habitats only. In the intermediate
habitats the various forms are usually thoroughly inter-
mixed, and the overall picture is rather complex. This com-
plex intergradation becomes very pronounced in areas such
as the mountains of eastern Cuba, particularly the Sierra
de Moa, where there is a high degree of habitat variation
over a relatively small geographical area. Here one finds
several ecological variants, the extremes of which are quite
distinct morphologically and often form large uniform popu-
lations occupying discrete ranges or micro-ranges. For this
reason several of these variants have been described by earli-
er authors as separate species or subspecies. A careful exam-
ination of these populations in the field, however, particularly
the various intermediate populations, shows that they com-
pose a single, large, intergrading system of variation. A
few collections of the extreme forms give a false picture of
the situation as a whole.
The common occurrence of complex variation patterns in
conjunction with apomixis has long been recognized. The
variation patterns themselves as well as the various factors
involved in the development of this variation are both multi-
form and complex. In general the degree of intergradation
within a system of variation is directly related to the amount
of gene exchange within and between populations. In some
instances barriers to gene exchange become quite rigid,
thereby insuring variation patterns of a discontinuous na-
ture (Gustafsson, 1947b). This is particularly striking in
plant groups in which apomixis has become obligate. For
example, in the genus Alchemilla (Rothmaler, 1936, 1941),
the species of certain sections have apparently lost all capac-
ity for sexual reproduction. The variation pattern is char-
acterized by numerous distinct populations or groups of
— Each of these forms a discrete unit, termed
‘microspecies” by some authors. These units are readily
separable from other such units, and thus form a discontinu-
ous pattern of variation. Moreover, according to Turesson
(1943) these “microspecies” are not rigidly uniform, but
are rather p and segregate, in res-
ponse to different environmental conditions, into what he
refers to as “agamotypes.” Other apomictic genera, which
in certain areas exhibit a variation pattern similar to that
described above, include Crataegus, Hieracium, Poa, Tarax-
42 JOAB L, THOMAS
acum, and several others of the familiar and “difficult” gen-
era (Gustafsson, 1947).
A similar situation is found in the genus Parthenium in
which two species, P. argentatum and P. incanum, are facul-
tatively apomictic. Rollins (1944, 1950) has shown that,
in Parthenium argentatum particularly, numerous different
apomictic populations are readily dstinguishable from other
such populations. There is a limited amount of intergrada-
tion between some of these populations, but they remain
relatively stable, even though the species is only facultatively
apomictic. However, it was clearly shown here that the
relationship between these apomictic populations and the
species as a whole is of a different order from the relation-
ships between the various species in the genus. Moreover,
the variation pattern found in the apomictic populations of
these two species, although discontinuous, is explainable in
terms of the variation found in the sexual members plus the
effects of polyploidy and introgressive hybridization between
the two species.
Similarly, the role of vegetative reproduction is undoubt-
ly of great significance as part of the mechanism involved
in the origin of a variation pattern such as that found in
Cyrilla racemifiora. As described above, this species propa-
gates and spreads extensively by vegetative shoots which
arise from horizontally situated, underground roots. By this
method large clones are developed which usually remain
interconnected by means of the root system. Self-incompat-
ibility has not been demonstrated experimentally in Cyrilla,
but evidence from field observations points clearly in this
direction. The plants produce abundant flowers and fruits
each year, but a high percentage of the fruits is partheno-
carpic. In certain areas, however, a few plants can be found
which produce a much higher percentage of seeds. In fact,
Copeland (1951), in studying the morphology of the repro-
ductive organs of Cyrilla, reported that the fruits of Cyrilla
always contain two seeds. Careful examination of the areas
in which there is a high percentage of seed development has
shown that there are always at least two different indivi-
duals growing in close proximity. Usually this is found in
the vicinity of two adjacent clones. Individual clones that
MONOGRAPH OF CYRILLACEAE 43
clone. Apparently self-pollination is sufficient to stimulate
the development of fruits, but fertilization of the ovule is
accomplished only by pollen from a different individual. It
is conceivable that an occasional self-pollination could result
in the fertilization of an ovule and the development of seeds,
but certainly the available evidence points to a high degree
of self-incompatibility.
The formation of large clones will tend to produce more
uniform local populations, although in self-incompatible spe-
cies, which tend to have more heterozygous populations, this
will not be so pronounced as in species capable of self-ferti-
lization. And, as a clone of a self-incompatible species
increases in size, the plants toward the center become in-
creasingly seed sterile due to the lack of pollen from a
different individual. The overall effect of mutations and
recombinations will also be somewhat different in species
capable of both sexual and asexual reproduction, in contrast
to species reproducing by sexual methods only. The pres-
ence of asexual reproduction will tend to preserve mutant
forms in two ways: 1. The “swamping effect” of sexual
reproduction will be lessened, due to a decreased rate of
sexual union and recombination; and 2. Homozygotes that
would be eliminated for one reason or another in a purely
sexual system could be maintained almost indefinitely and
even spread by asexual means.
Thus the combination of these factors found in Cyrilla
would tend to produce wide-ranging, heterozygous, and ge-
netically variable populations, capable of sexual reproduc-
tion but free from many of its limitations. Moreover, this
combination of factors will tend to produce, in a vari
environment, local populations with a relatively high degree
of adjustment to the habitat. In a genetically heterogeneous
group of this type, gene combinations that are well adapted
to the local environment can be “mass produced” by vege-
tative means. In this manner different populations will
segregate according to habitat preference. In a region
characterized by a high degree of ecological variation one
would expect to find a correspondingly high degree of local
variation among the populations occupying these different
sites. On the other hand, with sexual reproduction still
operative, there will be a certain amount of interbreeding
between populations. This will tend to maintain the genetic
heterogeneity of the group and also give rise to various
intermediate forms which will, in some cases, occupy inter-
mediate habitats. Thus if the various ecological segregates
44 JOAB L. THOMAS
show morphological differences, the group would then be
quite polymorphic, and the different morphological forms
would be more or less segregated according to habitat, with
occasional intergradations on the periphery.
TRENDS OF SPECIALIZATION IN PURDIAEA
There are several trends of specialization in Purdiaea
which indicate that the South American species, P. nutans, is
the most primitive species in the genus. The characteristics
of P. nutans which are considered to be primitive are the
following: leaves with uniformly pinnate venation; racemes
which show a zone of transition from a vegetative to a re-
productive shoot; and flowers with large, scarious, unequal
exterior sepals.
The first character listed, leaves with uniformly pinnate
venation, is found in P. nutans only. In all other species
there are two or more pairs of prominent, lateral ate
which run more or less longitudinally. In Purdiaea cui
these longitudinal, lateral veins arise all along the san tn
and are only slightly more prominent than the smaller,
pinnate lateral veins. These specialized, lateral veins become
quite prominent in other species of Purdiaea, forming an
almost linear sequence of increasing prominence as follows:
Purdiaea nipensis, P. stenopetala, P. stereosepala, P. moaen-
sis, P. velutina, P. shaferi, P. ekmanii. Correlated with this
increase in prominence is a tendency for these specialized
veins to originate near the base of the midvein. Thus, in
Purdiaea there is a series of venation patterns beginning
with a simple, pinnate venation in Purdiaea nutans, and
ending in a highly modified pattern with several prominent,
lateral veins that originate near the base of the leaf and are
almost parallel.
The largest break in this line of specialization occurs be-
tween Purdiaea nutans and P. cubensis, where the venation
pattern changes from simple pinnate to modified pinnate as
described Seach However, the difference between the two
diaea cubensis differ from this sire in being longer and
slightly more prominent. The smaller, lateral veins, and in-
deed most other aspects of the leaves of Purdiaea cubensis,
are very similar to those of P. nutans.
There is a similar line of specialization in the transition
MONOGRAPH OF CYRILLACEAE 45
from a vegetative to a reproductive shoot. In Purdiaea nu-
tans there is a definite zone at the base of the racemes where
a gradual transition from a vegetative shoot to a reproduc-
tive shoot occurs. In this zone there is a change in the
morphology of the shoot axis, and often a transition from
leaves to large bracts to smaller bracts. A similar zone is
found in P. cubensis. A zone of transition is also evident in
some collections of Purdiaea nipensis and P. moaensis, al-
though the zone is much smaller and more obscure than in P.
cubensis. In other collections of these species, however, the
SOUTH | CUBA
AMERICA
PINAR DEL RIO ORIENTE _ PROVINCE
|'SLE OF PINES gp
| P. EKMANII oe
| P. MICROPHYLLA
| P. VELUTINA
| P. PARVIFOLIA
| P. MOAENSIS
| P. aoe
|
| P. he
|
|
| eeaatt
| & cusensis
tae
P. NUTANS |
i
Fic. 23. aay relationships and present geographical distribution in the genus
Purdiaea. Further explanation in text.
t iti tative to a reproductive shoot is
abrupt. In a rather species of Pusdiges: this transition is
usually quite abrupt.
The texture of the sepals also shows ai gare! line of
specialization. In Purdiaea nutans and P. cubensis the
sepals are scarious and quite delicate; whereas in P. moa-
ensis, P. velutina, and particularly P. stereosepala, the
sepals are more foliaceous, or even coriaceous in the latter
Species. (Again, Purdiaea nipensis, with Apo ad mae 7
is
cubensis and the other Cuban species. Generally, there is
46 JOAB L, THOMAS
a correlation between the texture of the sepals and the size
relationship of the exterior sepals . The species with more
scarious sepals also have very unequal exterior sepals; and
species with more coriaceous sepals tend to have equal ex-
terior sepals. There are some exceptions to this, particular-
ly in Purdiaea velutina and P. moaensis. The texture of the
sepals is about the same in these two species, but in the
former the exterior sepals are clearly unequal, whereas in
the latter they are approximately equal.
A secondary line of specialization is indicated in Purdiaea
parvifolia and P. microphylla. As the names suggest, these
species are characterized by very small leaves. They also
have smaller stems, shorter racemes, and smaller floral parts
than other species in the genus. These species occur in ex-
posed and usually drier sites in the mountains, and have
apparently become specialized, through reduction, for these
habitats.
As is true of most plant groups, species of Purdiaea which
show a high degree of advancement i in one structure do not
il ng degree of ad tin
every other structure. The degree of correlation is high,
however, and _the trends in specialization are rather clear.
SYNOPSIS OF THE GENERA
Purdiaea: Sepals unequal, the exterior sepals consider-
ably larger than the interior sepals; petals lavender or
violet ; stamens 10, anthers inverted in bud, becoming erect
at anthesis, dehiscing through terminal pores; fruit 3-5-rib-
bed, style elongate, unbranched.
Cliftoma: Sepals equal ; petals spatulate, white or pink-
ish-white; stamens 10, anthers erect, dehiscing through
longitudinal slits, filaments broad and petaloid below the
middle, oe terete and subulate above; fruit distinctly
style sub-sessile, 3-5-lobed, massive.
Cyrilla: Sepals equal, petals white, thickened and glandu-
lar medially and below the middle; stamens 5, anthers de-
gs through longitudinal slits, filaments terete, subulate;
longitudinally bisuleate or trisuleate, style short, 2-3-
MONOGRAPH OF CYRILLACEAE 47
Purdiaea Planchon in Hook. Lond. Journ. Bot. 5: 250,
1846. Type: P. nutans Planch.
Costaea A. Rich. Fl. Cub. Fanerog. 2:75, 1853. Type: C.
cubensis A. Rich.
Alloiosepalum Gilg, Notizbl. Bot. Gart. Berlin 1, 102:97,
base of the raceme, those toward the apex of the raceme becoming
smaller and ovate to reniform. Sepals scarious to foliaceous, persis-
tent; increasing in length considerably during the maturation of the
fruit; glabrous to velutinous; insert in a quincuncial manner; un-
equal, the exterior sepals considerably larger than the interior sepals,
the first sepal equal to or as much as twice as long as the second and
third sepals. Petals 5, pink to violet; ovate to oblong; apex acute,
acuminate, or rounded; glabrous or rarely puberulent on the dorsal
surface; coalescent at the point of attachment to an hypogynous disk.
Stamens 10, the anthers oblong, caudate, versatile, with poricidal de-
hiscence; distinctly 2-lobed, the two lobes free above the point of at-
tachment of the filaments; the two thecae of each lobe becoming con-
fluent at anthesis; the anthers extrorse in bud, becoming inverted and
introrse at anthesis by the inflection of the apex of the filament. Fil-
aments subulate, adnate basally to the base of the petals at the point
Ovary
bose, glabrous to velutinous, the indumentum usually denser near the
apex; 1- to 5-seeded, the seed composed of a straight, cylindrical
embryo, surrounded by a mass of cellular endosperm; no seed coat
present, the ovary wall serving as the functional seed coat; style 1,
persistent, attenuate.
A. Leaves pinnately veined ............
A. Leaves with two or more lateral ve
lel to the leaf margin.
B. Leaves extremely thick and coriaceous, 0.4-0.6 mm. thick at ma-
turity, venation quite prominent on both sides, the lateral veins
forming a loose prominent network -----:-s-mrrer 11. P. ekmanii.
B. Leaves not over 0.2 mm. thick.
48 JOAB L. THOMAS
C. Leaves 0.5-1.3 em. long, 0.4-0.8 em. broad, are ipa ont ia
minate
C. Leaves 2 cm. long or longer.
D. Exterior sepals less than 2 mm. broad at anthesis, leaves with
revolute margins 5. P. parvifolia.
D. Exterior sepals more than 2 mm. broad at anthesis, leaves usually
without revolute margins.
E. First and second sepal approximately equal in length.
F. Young shoots velutinous aed the base of the peduncle, Lato
ca. 2 mm. in diameter at the bas 8. P.m
F. Young shoots glabrous or eheanuie so a tlow ‘the base of the saree
peduncle 1.0 mm. or less in diameter at the base.
G. First sepal rounded at the apex, thickened at the base, usually
asymmetrical, with the inner margin flattened, the outer margin
flaring outward above the middle; finely pubescent; particularly be-
low the middle Ae 5
G. First sepal with an acute or rarely obtuse apex; only slightly
thickened basally; symmetrical; finely pubescent on a conspicuous
dark area near the base, glabrous or nearly so above this area ........
P. stenopetala.
E. First sepal clearly longer than the second si
H. First leaves of a shoot developing into cdtepayila, densely clothed
with appressed, silky trichomes ..... .. 9. P. velutina.
H. First leaves of a shoot oleiccame no i categhgils. pak
I. First sepal elliptic-oblong; leaves with fine, reticulate venation
clearly visible 2. P. cubensis.
I. First sepal ovate, leaves without fine, reticulate venation.
J. Peduncle 2.5-3 mm. in diameter at the base, densely clothed with
silky, velutinous trichomes. Leaves broadly elliptic-obovate with
6-8 prominent lateral veins ............--scecsessesesesneeeesenseenee 10. P. shaferi.
J. Peduncle ca. 1 mm. in diameter at the base, finely pubescent to
puberulent. Leaves obovate-oblanceolate to spatulate, 2-4 promin-
ent lateral veins 3. P. nipensis.
1, Purdiaea nutans Planchon in Hook. Lond. Journ. Bot.
5: 251, 1846. Type: not seen.
Alloiosepalum weberbaueri Gilg, Notizbl. ‘Bot. Gart. Ber-
lin 11, 102:97, 1931.
rdiaea weberbaueri (Gilg) Mattick, Notizbl. Bot. Gart.
Berlin 12, 393 :398, 1935.
Shrub or small tree up to 6 or 7 meters tall; bark light gray or
brown; stems glabrous. Leaves dark green, shining above, pale be-
neath; 2-5 em. long, 1-2 em. broad, obovate, oblong, or oval; apex
obtuse or rounded, mucronulate; base cuneate, narrowly sessile; mid-
vein prominent beneath, depressed above, secondary veins much weak-
er, uniformly pinnate, bending upward slightly at the margin; smaller
reticulate veins distinct beneath, less so above. Racemes 2.5-5.5 em.
‘The otiginal deseription was based on material in the Hooker Herbarium collected
vee ee ee (Colombia) .
MONOGRAPH OF CYRILLACEAE 49
long, suberect to pendulous; a zone of transition from a vegetative to
a reproductive shoot near the base of the peduncle: rachis attenuate,
ca. 1.0 mm. in diameter at the base, tapering to 0.5 mm. in diameter
at the apex; villous, tomentose, or glabrous: pedicel 2-5 mm. long,
those at the base. of the raceme longer than those at the apex; villous
to glabrous, the ling to that on the peduncle;
the articulation at the point of attachment belie the Sagcieiee usually
swollen: bracts lanceolate or up
mm. long near the base of the raceme, ion mm. long ede the
apex; glabrous or rarely pubescent along the midvein near the base;
borne on a small protuberance of the peduncle, distinctly articulate at
the base. Sepals scarious, very = variable in size; the first
sepal 14-17 mm. long, 7-10 mm. broad at anthesis, up to 19 mm. long,
12 mm. broad at maturity, conaderaly longer than the second sepal;
broadly ovate; apex acute or occasionally obtuse or rounded, mucron-
ulate; glabrous or nearly so: sar sepal 8-12 mm. long, 6-7 mm.
broad at anthesis, up to 13 mm. long, 8 mm. broad at maturity; apex
obtuse or acute, occasionally mucronulate; glabrous or nearly so:
third sepal 7-9 mm. long, 3-4 mm. broad at anthesis; asymmetrical,
the inner margin flattened and cilliate, the outer margin rounded and
glabrous; interior sepals 4.5-5.5 mm. long, 1-1.5 mm. broad at anthesis,
up to 6.5 mm. long, 2.5 mm. broad at maturity; lanceolate; margin
strongly ciliate. Petals 9-11 mm. long, 3-4 mm. broad at anthesis;
apex obtuse or acute, glabrous; violet in color. Anthers 3.5-5 mm.
long, rather variable in size in a single flower; bright yellow: fila-
50 JOAB L. THOMAS
ca. 5.5 mm. long at anthesis, becoming 7 mm. long at maturity; glab-
rous; attenuate.
DISTRIBUTION: Northern South America, dense forests or “elvan
woodlands” of higher elevations.
REPRESENTATIVE SPECIMENS SEEN: VENEZUELA. State of Amazonas:
Cerro Huach i, Rio C ; along South Escarpment at
1500 m. alt., Maguire, Cowan, and Wurdack 29803 (US, NY); vicinity
Summit Camp at 1500 m. alt., Maguire, Cowan, and Wurdack 30011
(us, NY); Summit Camp at 1500 m. alt., Maguire, Cowan, and Wur-
dack 30211 (Us, NY, MO) ; Cerro de la Neblina, Rio Yatua; West Cum-
bre Camp at 5800 feet alt., Maguire, Wurdack, and Bunting 36956
(Ny) ; along west escarpment, 1700-1800 m. alt., Maguire, Wurdack,
and Bunting 37083 (Us, NY, F). Summit of Mount Duida; alt. 7100
feet, Tate 611 (US, NY); alt. 5500-6000 feet, Tate 425 (Ny). Rio Ven-
tuari; Serrania Paru, Rio Paru, Cano Asisa, Cowan and Wurdack
31074 (Us, NY, UC). State of Zulia: Helechal Perija; alt. 2000 m.,
Gines 2022 (US).
COLOMBIA, Department of Huila: Cordillera Oriental; Rio Ambrica,
near confluence of Quebrada la Trinidad, Fosberg and Holdridge 19567
(us, NY). Without locality; Bruckmiller (ny).
Department of Amazonas: C entre
y Taulia, Raimondi 2054 (GH) ; Molinopampa, Weberbauer 4332 (cH).
Purdiaea nutans is the only species of Purdiaea which oc-
curs in South America. It is found in dense forests or “elvan
woodlands” of higher elevations, usually 1500-2000 meters
above sea level. It is sparsely distributed, seldom if ever oc-
curring in large stands or populations. The most distinctive
and probably the most primitive species in the genus, Pur-
nutans is most closely related to P. cubensis. It is easily
distinguishable from the latter species, and indeed, all other
species of Purdiaea by several characters. Purdiaea nutans
is the only species in which the leaves have uniformly pin-
nate venation. Its large flowers are distinctive, being the
largest in the entire Cyrillaceae. The broadly ovate first sepal
and the pendulous racemes also serve to distinguish this
species from Purdiaea cubensis.
A relatively wide-ranging species , Purdiaea nutans shows
a minor degree of variation in several characters. The
leaves, bracts, and sepals are usually somewhat smaller on
specimens collected at high elevations than on specimens
collected at lower elevations in the same geographic area.
The petals, also, are variable in size, but in contrast to the
above structures, a wide range of variation can be found on
a single plant. Occasionally a size range comparable to that
known from the entire species can be found on a single spe-
cimen. Similarly. the degree of pubescence on the rachis and
the pedicel is rather variable, but this variation has no ap-
MONOGRAPH OF CYRILLACEAE 51
a. es
parent geographic or altitudinal signifi P
varying from glabrous to pubescent to tomentose are found
throughout the range of the species, with the exception of
the two collections from Peru. The apex of the first sepal is
also rather variable in shape, varying from acute to obtuse
or rounded. ‘
Purdiaea weberbaueri (Gilg) Mattick was described from
specimens collected in Peru and was segregated from P.
nutans on the basis of its glabrous racemes and more acute
exterior sepals. However, it falls well within the variation
pattern of P. nutans and is included in that species in the pre-
sent treatment. Several collections of P. nutans have been
made in recent years in Colombia and Venezuela. These col-
lections, which were not available when Purdiaea weber-
bauri was described, have made possible a much better under-
standing of the variation pattern in the present species. As
discussed above, the degree of pubescence on the racemes and
the shape of the sepal apex are among the more variable
characters of Purdiaea nutans. The variation in these char-
acters is graded, with many intermediates between the ex-
treme forms. Several specimens of Purdiaea nutans are in-
distinguishable from the type of P. weberbauri. There are,
in fact, collections of P. nutans which accord with the ori-
ginal description of P. weberbauri better than the type col-
lection of that species. In the latter collection the first three
sepals have ciliate margins and the third sepal is finely pu-
berulent on the inner edge of the ventral surface. There
are specimens of P. nutans in which the first three sepals are
glabrous. Future collections of Purdiaea nutans from Peru
will undoubtedly show more variation in the degree of pube-
scence in the plants from that area.
2. Purdiaea cubensis (A. Rich.) Urban, Fedde, Repert.
Spec. Nov. 22:366, 1926. Type: not seen.*
Shrub up to 4 meters tall, ste
sears. prominent, triangular.
somewhat crowded toward the ends of branches, 3-7
cm. broad, obovate to elliptic; apex rounded or obtuse 3 base narrowly
sessile: midvein prominent on both surfaces; 4-6 longitudinal lateral
veins arising along the length of the midvein and running more or
4 The original description of Costaea cubensis was based on material collected in
“Sierra de Marcos Guerra” by José Maria Valenzuela.
52 JOAB L. THOMAS
less parallel to the margin of the leaf, prominent above, less so be-
neath; smaller lateral veins freely branching, pinnate, forming inter-
connections between the midvein and the longitudinal lateral veins.
Racemes 5-12 em. long, erect: rachis glabrous or puberulent; subulate:
pedicel 3-4.5 mm. long, glabrous or puberulent. Bracts oblong, 7-10
mm. long, 2-3 mm. broad near the base of the raceme, becoming ovate,
2-4 mm. long, 1-2.5 mm. broad near the apex of the raceme; glabrous
or nearly so, occasionally with a finely ciliate margin; articulate at
the base. Sepals light pink to white; scarious; very unequal: first
sepal oblong to ovate-oblong, 12-14 mm. long, 6-8 mm. broad at anthe-
sis, up to 16 mm. long, 10 mm. broad at maturity; apex obtuse, rarely
ein glabrous or finely pubescent near the base, often with a ciliate
gin: second sepal ovate, 6-8 mm. long, 2-5 mm. broad at anthesis,
re 10 mm. long, 5-6 mm. broad at maturity; apex obtuse, mucronulate,
glabrous or finely pubescent toward the base; margin ciliate: third
sepal approximately the same length as the second sepal at anthesis,
often slightly larger than the second sepal at maturity; interior mar-
in flattened, strongly ciliate; exterior margin rounded, glabrous or
finely ciliate: fourth and fifth sepal lanceolate, 4-6 mm. long at anthe-
sis, up to 10 mm. long at maturity; pubescent, margin ciliate; mid-
vein conspicuous. Petals 5-7 mm. long, 2-4.5 mm. broad, ovate to ovate-
oblong; apex abruptly acuminate, glabrous; deep pink or violet.
Anthers 2-2.7 mm. long, weakly caudate: filaments 2.5-3 mm. long;
narrow, slightly subulate. Ovary subglobose to globose, 1-1.5 mm. in
diameter at anthesis; 5-lobed, glabrous or pubescent: style subulate,
4.5-5 mm. long at anthesis, up to 7 mm. long at maturity. Fruit 2-2.5
mm. in diameter, subglobose to globose; strongly 5-angled; glabrous
or pubescent.
DISTRIBUTION: Pine barrens of Pinar del Rio Province, Cuba and
the Isle of Pines.
MONOGRAPH OF CYRILLACEAE 53
REPRESENTATIVE SPECIMENS SEEN: CUBA. Proy. Pinar del Rio: Wright
2204 (UC, GH, MO); Carabia 3940 (NY); Pinares near Loma, Marcos
Guerra, N. W. of S. Diego, Léon and Charles 4887 (NY, MT, LS); Gal-
alon to San Pedro Del Caimito, Shafer 11950 (Us, NY); Pinar del Rio
City, between Laguna Maguina and Laguna de Junco, Ekman 17905
(Ny); Herradura, Van Hermann 5851 (sv, 2 sheets); Sabanas de
Santo Tomas, Acufia 11210 (sv); between Mina de Oro and Playa
del Soldado, Killip 43900 (us, LS); pinelands between Pinar del Rio
and Surnidero, Roig 3150 (US, GH). Santa Catalina; Van Hermann,
3242 (uC, SV); Caldwell and Baker 7073 (F); cerro de Cabras; Léon
17758 (LS); Britton, Britton, and Cowell 9796 (NY); Marie-Victorin,
Léon, Alain, and Carabia 17758 (MT); Léon 3395 (NY, MT, LS); on
Guane Road, Britton, Britton, and Gager 7289 (US, NY, F). La Cajal-
bana; Acuiia 16439 (sv); Ekman 70480 (Ny); La Palma, Alain and
Acuiia 1169 (Ls).
ISLE OF PINES: Vicinity of Los Indos; Marie-Victorin and Léon
17864 (MT); Jennings 326 (US. NY, GH); Britton, Britton, and Wilson
14202 (GH, F, US, NY, MO); Killip 44042 (Ny, LS); Léon and Marie-
Victorin 17864 (LS); Marie-Victorin and Alain 88 (LS); Calvius and
Elfameli 7978 (sv); Léon, Marie-Victorin, and Carabia 18860 (Ls).
Purdiaea cubensis, a very distinctive species of the pine
barrens and savannas of western Cuba and the Isle of Pines,
is seldom if ever confused with any other species of the
genus. In common with all species of Purdiaea from Cuba,
the leaves of P. cubensis have a venation pattern character-
ized by two or more pairs of prominent lateral veins which
arise from the midvein, curve abruptly, and run longitudinal-
ly, parallel to the margin of the leaf. In the leaves of Purdi-
aea cubensis, however, there is a fine, densely reticulate pat-
tern of smaller veins, which is distinctly visible on both sur-
faces of the leaf. This characteristic reticulum is not found
in the leaves of other species of Purdiaea from Cuba. A sim-
ilar reticulate pattern is found in the leaves of Purdiaea
nutans, but in that species the lateral veins are pinnate, and
do not run parallel to the margin of the leaf. The large, de-
licate oblong-elliptic exterior sepals also serve to distinguish
Purdiaea cubensis from all other species in the genus.
Purdiaea cubensis is most closely related to P. nutans and
is discussed under that species. Among the Cuban species,
Purdiaea cubensis seems most closely related to P. nipensis.
This relationship is obscure, however, and Purdiaea nipensis,
in contrast to P. cubensis, is clearly related to other Cuban
species. 3
In spite of its bicentric distribution, Purdiaea cubensis ex-
hibits only minor morphological variation. The material
from the Isle of Pines was segregated by Marie-Victorin
(1948) as Purdiaea cubensis var. albosepala, but there seem
54 JOAB L. THOMAS
to be no constant characters by which the material from
these two areas can be distinguished. According to the above
author, the plants from the Isle of Pines have white sepals,
whereas those from Pinar del Rio have pink or rose-colored
sepals. Very few specimens have label data giving the color
of the sepals, but in at least two collections from the Isle of
Pines (E. P. Killip 44042 and 43900) the sepals are described
as varying from pink to white. In a few other collections
from this area (notably Jennings 326) the sepals have re-
mained distinctly pink on the dried material, although sepal
color in Purdiaea usually fades upon drying. I have not seen
P. cubensis in the field, but in all of the species of Purdiaea
which I have seen in the field, sepal color was rather variable.
In Purdiaea velutina, for example, the color of the sepals
was found to vary from white to deep pink in plants growing
only a few feet apart.
The average leaf size in Purdiaea cubensis is usually
slightly larger in the collections from the Isle of Pines than
in collections from Pinar del Rio, but there is a high degree
of overlap in the measurements, and several collections can
be assigned to the above variety only if the area from which
they were collected is known. For these reasons it seems
that the material of Purdiaea from the Isle of Pines does not
warrant varietal status, and variety albosepala is not recog-
nized in the present treatment.
There are a few collections of Purdiaea cubensis from
Pinar del Rio which have noticeably smaller than average
leaves ( Van Hermann 3243 and Caldwell and Baker 7073.)
There is also some variation in the shape of the exterior
sepals. One specimen in particular (Roig 3150) has sepals
that are narrower and more pointed than those found in
other collections of this species. These are apparently ecolo-
gical variants, although there is little information available
on the effects of different ecological factors on any of the
species in this genus.
3. Purdiaea nij Marie-Victorin & Léon, Contrib.
Inst. Bot. Univ. Montréal, 63:55, 1948. Type: Léon,
Marie-Victorin, Clément, and Alain 19837
Purdiaea nipensis Marie-Victorin var. alaini Marie-
Victorin, Contrib. Inst. Bot. Univ. Montréal 63:56, 1948.
Shrub with light gray bark, young shoots puberulent to glabrous;
light brown, becoming gray with age. Leaves highly variable in size,
2.5-10 em. long, 1-1.5 em. broad ; obovate, obovate-spatulate, or obovate-
oblanceolate; apex acute, obtuse, or te
MONOGRAPH OF CYRILLACEAE 55
base attenuate or cuneate; midvein and 2-6 longitudinal lateral veins
prominent beneath, weak above; 2 longitudinal lateral veins often
arising near or slightly above the middle of the leaf; smaller reticu-
late veins distinct beneath, less so above. Racemes 3.7-7 cm. long,
erect to pendulous: rachis pubescent or puberulent, attenuate; ca. 1
mm. in diameter at the base tapering to 0.3 mm. in diameter at the
apex; zigzag in shape, often strikingly so toward the apex: pedicel
1-2 mm. long, 0.3-0.5 mm. in diameter at anthesis, pubescent: bracts
lanceolate, ca. 6 mm. long near the base of the raceme, becoming ovate-
lanceolate to ovate, ca. 2 mm. long toward the apex of the raceme;
pubescent below the middle, glabrous above. First sepal 9-11 mm.
long, 4.5-6 mm. broad at anthesis, up to 15 mm. long, 9 mm. broad
at maturity; considerably longer than the second sepal; ovate to
glabrous or pubescent.
DISTRIBUTION: Mountains and stream valleys of Oriente Province,
Cuba.
PURDIAEA NIPENSIS
56 JOAB L. THOMAS
REPRESENTATIVE SPECIMENS SEEN: CUBA. Prov. Oriente: Sierra de
Nipe; Arroyo seco between Woodfred and La Mina, Léon, Marie-
Victorin, Clément, and Alain 19837 (Type: GH. Isotypes: LS, 2 sheets;
sv; MT); Léon, and Alain 20414 (MT, Type of P. nipensis var. alaini) ;
Cayo del Rey, Gaaiaies de la Loma de Bio, Carabia 4059 (NY, LS); En
el charrasco, Cayo Rey, Lopez 1839 (LS); Ekman 9485 (ny); Bosque
de La Plancha, Léon and Alain 19110 (LS); Cajo La Plancha, sur la
limonite granuleuse, Léon and Alain 19111 (LS, MT); Chemin de la
‘oodfred, Léon and Alain 19147 (MT, Ls) ; Léon, Marie-Victorin,
and Alain 19837 (tS); Alto de la Bandera, Léon, Marie-Victorin, Clé-
ment, Alain 19754 (LS); Falda Norte Loma Mensura, M. Lopez 2824
(sv, Ls); Charrascales de Arroyo Potrero, Lopez 2824 (SV, Ls); Marie-
Lage Carabia 4059 (MT); South of lumber camp, alt. 600-700 m.,
C. V. Morton and Acufia 3051 (us); along trail Piedra Gorda to
Woodfred, 400-500 m. alt., Shafer 3110 (us, NY); Minas de la Nicaro,
Ocujal, Acufa, Alonzo, tind Pifia 18781 (sv). Sierra de Cristal;
Bosques Reciedus de El Prado, Alain, Acuiia, and Lopez 5391 (SV, LS);
Alain, Lopez 4694 (LS). Sierra de Maestra: Pico de La Bayamesa,
4800 ft., Lopez 2304 (SV, LS); between Pico Turquino and La Baya-
mesa, alt. 1350 m., Morton and Acuia 3755 (US); Banks of Arroyo
Corjo, near Nagua, Pico Turquino, Ekman 14738 (NY, LS). Vicinity
of Toa; Orillas de un arroyo, Km. 20 al Sur de Sabanilla, Via Azul,
Alain and Morton 5159 (Ls) ; Nuevo Pinar, Alearraza arriba, Clément
5107 (48); same locality, Clément and Carlos 5107, (LS). Without
locality; Wright 2205 (GH, UC).
species, known from three rather isolated mountain
ranges in eastern Cuba, is the most probable connecting link
between Purdiaea cubensis and the other Cuban species of
Purdiaea. Although the relationship between these two spe-
cies is rather distant there are two points of similarity which
seem significant. In the leaves of both species two or more
of the longitudinal lateral veins consistently arise near the
middle of the leaf. This occurs sporadically in other species
but usually the longitudinal lateral veins arise near the base
of the leaf. The other point of similarity is the very unequal
exterior sepals, with the first sepal almost twice as long as
the second sepal. This extreme size difference between the
two exterior sepals is not found in any of the other species
of Purdiaea from Oriente Province (although it is found in
P. cubensis from Pinar del Rio and P. nutans from South
America). In general aspect, and in most characteristics,
however, Purdiaea nipensis is more like the other Cuban
species, all of which are endemic to Oriente Province. It is
most closely related to P. stenopetala, but the two species
are readily distinguishable because of the longer and very
unequal exterior sepals i in P. nipensis.
nipensis is one of the more variable species in
the genus, and future collections and field observations in
MONOGRAPH OF CYRILLACEAE 57
eastern Cuba may necessitate the establishment of one or
more geographical varieties. The material from the Sierra
de Maestra has longer and more obovate leaves, and some-
what more acuminate sepals than the material from the
Sierra de Nipe. Similarly, in the collection from extreme
eastern Cuba, south of Sabanilla (Alain and Morton 5159)
the leaves are even longer and more elliptic and the sepals
are more foliaceous than those of collections from other parts
of the range. The collections from the Sierra de Maestra and
Sabanilla are few in number, however, and are insufficient
to show the variation pattern of the plants from these areas.
Purdiaea nipensis var. alaini Marie-Victorin, on the other
hand, occurs in the Sierra de Nipe, apparently sympatric
with other representatives of the species which are not as-
signed to this variety. The variety does differ from the type
in having slightly broader leaves and more acuminate sepals,
but this difference falls well within the range of variation
found in other collections from this area, and there are many
intermediates between the two forms. For this reason var.
alaini is not retained in the present treatment.
NoTES: Some confusion has arisen as to which sheet of
the type collection represents the holotype. In the specimen
citations following the original description of the species,
Marie-Victorin (1948) cited the type as follows: “Arroyo
Seco, entre Woodfred et la Mine, Sierra de Nipe. 6 Avril,
1941. Léon, Victorin, Clément and Alain 19837. (TYPE dans
Vherbier de Colegio de la Salle, La Habana, Cuba)”. To my
knowledge there is but one specimen on which the label data
conforms to this citation. This specimen, now in the Gray
Herbarium, is stamped “EX HERBARIO DE LA SALLE; HABANA,
cuba”. In the herbarium of the College of La Salle in
Havana there are two sheets bearing collection number
19837, but the name of one of the collectors, Clément, is not
listed. One of these sheets, marked as TYPE, has the locality
designated as “Cayo de las Mujeres, falda Oeste de la Loma
Mensura, Sierra de Nipe, Mayari, Oriente”, and is dated
April 6, 1941. The other sheet is dated April 4, 1941, and
has the locality designated as: “Camino de la Mina — Wood-
fred, Sierra de Nipe, Oriente”. The other two isotypes cited
above bear a full complement of collectors, Léon, Victorin,
Clément, and Alain; both are dated April 6, 1941, and the
locality data, though not the same, apparently refer to the
same area. The locality is apparently the same for all five
of these specimens, and so far as I can tell the material on
the five sheets is identical. It seems logical to assume, there-
58 JOAB L. THOMAS
fore, that the sheet in the Gray Herbarium does represent
the holotype since it is the only sheet that conforms in hab-
itat data to the original citation, and since this sheet appar-
ently was in Havana at the time of the citation.
1860. Type: Wright 34
Purdiaea ophiticola Marie-Victorin, Contrib. Inst. Bot.
Univ. Montréal, 63 :56, 1948.
Shrub with light gray bark, stems glabrous, smooth, light brown
becoming gray with age. Leaves dark green above, pale beneath,
2.5-8 cm. long, 1-2 em. broad; obovate, obovate-oblanceolate, or spat-
ulate; apex obtuse, truncate, or slightly emarginate and mucronulate;
base attenuate, narrowly sessile; midvein and 2-4 longitudinal lateral
veins running parallel to, and very near, the leaf margin; smaller
secondary veins more or less distinct beneath, less so above. Racem
5-6 cm. long at mid-anthesis, up to 7.5 em. long at maturity, usually
more or less pendulous. Rachis glabrous, puberulent or pubescent;
attenuate, ca. 1.5 mm. in diameter at the base, tapering to ca. 0.5 mm.
in diameter at the apex, ridged. Pedicel 1-1.5 mm. long, 0.5 mm. in
diameter, glabrous or pubescent, distinctly articulate at the point of
attachment to the rachis and the receptacle. Bracts ovate-lanceolate
or lanceolate, 4-5 mm. long near the base of the raceme, becoming
ovate to reniform, 1-1.5 mm. long near the apex of the raceme; articu-
late at the base; glabrous or pubescent, the i pondi:
to that of the peduncle. Sepals highly variable in size, scarious in bud
becoming more foliaceous at maturity; the first sepal 5-7.5 mm. long,
3.5-4 mm. broad at anthesis, up to 12 mm. long, 7 mm. broad at ma-
turity; ovate or ovate-deltoid; apex acute or obtuse; base truncate or
cordate; finely pubescent on a conspicuous, dark area near the base
of the sepal, glabrous, or nearly so above this area: second sepal ap-
proximately the same size as the first sepal with a narrower and more
acute apex; usually pubescent on both surfaces below the middle,
glabrous or nearly so above: third sepal equal to or slightly shorter
than the second sepal; apex acute, base truncate; asymmetrical, the
4. Purdiaea stenopetala Grisebach in Goett. Abh. 9:45,
1
pubescent on both surfaces, densely so below the middle; the two sepals
more or less opposite and enclosing the fruit. Petals 5-6 mm. long,
with a weakly ciliate margin near the base; deep violet in color. An-
thers 2-2.7 mm. long, slightly caudate; filaments 2.5-3 mm. long, subu-
late. Ovary subglobose, ca. 1.5 mm. in diameter at anthesis; 5-angled,
glabrous or pubescent. Style persistent, subulate, ca. 3 mm. long at
anthesis, becoming 5-6.5 mm. long at maturity; glabrous, exserted.
Fruit subglobose to ovoid, usually strongly 5-keeled, 2.5-3.2 mm. in
diameter at maturity.
DISTRIBUTION: Mountains of Oriente Province, Cuba.
MONOGRAPH OF CYRILLACEAE 59
REPRESENTATIVE SPECIMENS SEEN: CUBA. Prov. Oriente: C. Wright
341 (isotypes: MO, 2 sheets; GH, 2 sheets; NY, 2 sheets; UC). Vicinity
of Moa; sur la limonite humide de la plage, Marie-Victorin 21433 (MT,
type of Purdiaea ophiticola Marie-Victorin; isotypes: MT, GH, LS);
Acuiia 12528 (US, SV); Playa Vaca, Acuna 12527 (US, SV); Acuna
12529 (us, SV); Bucher 19 (NY); Bucher 1173f (Sv); Bucher 11044
(sv); Petit ruisseau de chemin Aviation, Clément 3557 (MT, Ls) ; Clé-
ment 3559 (MT, 2 sheets; Ls, 2 sheets); Léon, Marie-Victorin, and
Clément 20669 (MT, LS); Pinares de Moa, Marie-Victorin and Clé-
ment 2592 (LS); entre la vallée du Rio Cayoguan et la vallée 4 l’est,
vers la mine Delta, Marie-Victorin and Clément 3666 (GH, MT, Ls);
Clément 3672 (MT, LS); Sobre la meseta, Alt. 400 m., Marie-Victorin,
Clément, and Alain 21786 (LS). Rio Cabanas; Charrascos, Léon,
Marie-Victorin, and Clément 20736 (LS); plateau entre le Rio Cabana
et le Rio Moa, Marie-Victorin and Clément 21786 (MT, 2 sheets; GH).
Cayo Chiquito; Clément, Alain, and Chrysogone 3802 (mT) ; Clément,
Alain, and Chrysogone 3803 (LS). Charrascal del Coco; Lugares
secos, Léon, Alain, Clément, and Chrysogone 22598 (Ls) ; Léon, Alain,
Clément, and Chrysogone 22615 (LS) ; Léon, Clément, Alain, and Chry-
sogone 4718 (LS). Rio Jicotea; Léon, Clément, Howard 20143 (MT,
Ls); Léon, Clément, and Howard 20144 (mt, 2 sheets; Ls); Pinares,
Léon, Clément and Howard 20174 (LS, MT); Rio Yagrumaje; Alain,
Clément and Chrysogone A. 953 (LS); Clément 3611 (Mt, Ls); Pin-
ares, Clément, Alain and Chrysogone 6854 (LS) ; Charrascos, Marie-
Victorin, Clément, and Alain 21787 (LS). Cananova; La Esmeralda,
Clément, Alain, and Chrysogone 6926 (LS); Cerro de Miraflores, Léon
21098 (MT, LS); Léon 21146 (MT, LS); Léon 21154 (MT, Ls). Yamuri
Arriba; to Bermejal, Shafer 8456 (US; NY, 2 sheets) ; Sierra de Moa;
alt. 800 m. approx., Alain 3430 (LS).
In reproductive as well as vegetative structures, Purdiaea
stenopetala is the most variable species in the genus. The
leaves vary in size from 2.5-8 cm. and in shape from obovate
ee.
60 JOAB L. THOMAS
to obovate-oblanceolate to spatulate, with the apex varying
from obtuse to truncate to slightly emarginate. The obovate
leaves usually have an obtuse apex, and as the leaves become
more spatulate the apex becomes more abruptly truncate or
emarginate. The variation within the complex is graded,
and it is possible to find leaves of varying size and shape
within one geographic area.
The sepals are also variable, the first sepal ranging from
5-7.5 mm. in length at anthesis. Moreover, the exterior se-
pals often nearly double in length between anthesis and
maturity. The ontogenetic increase in size is not always so
extreme, however, and as a result the sepals are even more
variable in size at maturity than at anthesis. Failure to
recognize this variation in the ontogeny of the sepals has,
in the past, led to some confusion in the taxonomy of this
variable group.
The pubescence on the rachis also varies in Purdiaea sten-
opetala. The extreme forms may be glabrous or pubescent,
but again there is gradation with many intermediates be-
tween the extremes.
Marie-Victorin (1948) divided this group into two spe-
cies: Purdiaea stenopetal ted only by the type col-
lection (Wright 341); and P. ophiticola, a new species,
represented by all of the other collections in this complex.
The latter species was differentiated from the former on the
basis of the two most variable characteristics in the complex:
sepal size and leaf shape. Oddly enough, there is little if any
No character or combination of characters has been found
by which this complex can be divided into discontinuous
groups. Thus, until more is known about the nature of the
variation in this group it seems that the entire complex
should be considered a single, variable species. Precise field
observations and population studies of this species are badly
needed for a better understanding of the variation in this
interesting group.
The species most closely related to Purdiaea stenopetala
are P. nipensis and P. parvifolia. The relationship is dis-
MONOGRAPH OF CYRILLACEAE 61
cussed under those species. Purdiaea stenopetala also re-
sembles P. stereosepala in the shape and venation of the
leaves and in general aspect.
5. Purdiaea parvifolia (Marie-Victorin) Thomas, comb.
nov. Type: Shafer 8179
Purdiaea ophiticola var. parvifolia Marie-Victorin, Con-
trib. Inst. Bot. Univ. Montréal, 63:57, 1948.
Shrub 3-4 meters tall, bark light gray; young stems pubescent or
velutinous, the indumentum usually closely appressed. Leaves dark
Rachis attenuate, ca. 1.0 mm. in diameter at the base, tapering to 0.2
mm. in diameter at the apex; pubescent or velutinous. Pedicel 1-1.5 mm.
long, ca. 0.3 mm. in diameter, pubescent or velutinous, the indumentum
corresponding to that of the peduncle. Bracts ovate-lanceolate, ca.
3.5 mm. long at the base of the raceme, becoming ovate to ovate-deltoid,
ca. 1.0 mm. long toward the apex of the raceme: velutinous below the
middle, margin ciliate. Sepals 1.8 mm. broad at anthesis, up to 7 mm.
long, 3 mm. broad at maturity; ovate-lanceolate; apex acute, obtuse,
or rarely somewhat rounded; glabrous or puberulent toward the base
near the margin; margins ciliate: second sepal approximately the same
length or only slightly shorter than the first sepal; narrower with a
more acute apex; pubescent toward the base: third sepal usually sym-
metrical, ca. 2/3 as long as the first and second sepal in bud, the three
sepals approximately equal shortly after anthesis; interior sepals lan-
ceolate, apex acute or acuminate; 3-3.5 mm. long, 0.7-1 mm. broad at
anthesis, 3.5-4 mm. long, 1-1.5 mm. broad at maturity; pubescent or
velutinous on both surfaces; margin ciliate. Petals oblong, 3.5-4 mm.
long, 2.5-3 mm. broad; apex obtuse or acute; usually puberulent near
the base on the dorsal surface, particularly in bud. Anthers 1.5-1.8 mm.
long, weakly caudate: filaments 2-2.3 mm. long, subulate, glabrous.
Ovary globose, ca. 1.2 mm. in diameter at anthesis, weakly 5-ribbed;
glabrous or pubescent; style 1.5-2 mm. long at anthesis becoming 2-2.5
mm. long at maturity. Fruit globose, 5-ridged, pubescent or glabrous,
the indumentum concentrated along the ridges.
DisTRIBUuTION: Mountains of northern Oriente Province, Cuba.
U
along the Piloto Road, ca. 3000 ft. alt., on red, sandy soil, Thomas 522
(GH); Thomas 523 (GH); Summit, Thomas 524 (GH); Thomas 525
62 JOAB L. THOMAS
PURDIAEA
PARVIFOLIA
a
=
(GH); Thomas 526 (GH); Camp San Benito; Shrub 6-15 ft., very
common in higher altitudes (2500-3100 ft.), Shafer 4057 (US, NY, GH);
Vicinity of Toa; La Magdalene, Charrascos de Pefia Prieta, Alt. ap-
prox. 750 m., Alain 3618 (LS, SV) ; Bosque charrascoso, cumbre del Pico
Galano, Sierra del Frijol, La Alagria, Alt. approx. 1200 m., Alain 3711
(is).
This species was originally described as Purdiaea ophiti-
cola var. parvifolia Marie-Victorin, but now that abundant
flowering and fruiting material is available it is readily
apparent that this taxon is a distinct species. Although
Purdiaea parvifolia is closely related to P. stenopetala, it
is easily separated from that species on the basis of several
morphological characters. In Purdiaea parvifolia the leaves
are smaller, more obscurely veined, and have revolute mar-
gins; the stems are more densely pubescent; and the exterior
sepals are shorter, narrower, and more lanceolate. The
petals of Purdiaea parvifolia also differ from those of P.
stenopetala, being shorter but relatively broader, and puber-
ulent on the dorsal surface near the base. Purdiaea parvi-
folia also resembles P. microphylla and is discussed under
that species.
In contrast to most of the species in the genus, Purdiaea
parvifolia is seldom if ever found as scattered individuals
growing along stream banks. In the Sierra de Moa, cer-
tainly, it occurs as large populations, extending over several
acres. _ Though it is abundant in these areas, it does not
occur in dense stands, but grows in a very rich, mixed vege-
tation. The soil in this area is dark red with a high iron
content. The entire area is quite moist, but Purdiaea parvi-
folia is usually found in the more exposed and relatively
drier places.
MONOGRAPH OF CYRILLACEAE 63.
There is little variation within a given population of this
species, but there is a small degree of clinal variation parti-
cularly in leaf size. As would be expected, the leaves are
somewhat smaller on specimens collected at higher elevations
than on those collected at lower elevations, but the difference
between the most extreme forms is not particularly striking.
There is also some variation in size and shape of the external
sepals, although this seems to have little or no correlation
with elevation. Future collections of flowering and fruiting
material of Purdiaea parvifolia from the Sierra de Cristal
and in the vicinity of Toa will possibly show more significant
geographic variation in this species.
The material available from these regions differs some-
what from the material from Sierra de Moa in the size and
shape of the leaves; but there are only one or two collections
from each area, and all of the material is sterile.
6. Purdiaea microphylla Britton and Wilson, Bull. Tor-
rey Club 42:389, 1915. Type: Shafer 8265
abruptly below the middle to a narrowly sessile base; mid-vein prom-
inent beneath, obscure above, 2-4 longitudinal lateral veins, faintly
visible beneath, usually not showing above in mature leaves (venation
more prominent in immature leaves). erect, 1-2 cm. long:
rachis pubescent to puberulent, attenuate; ca. 0.8 mm. in diameter at
the base tapering to 0.2 mm. in diameter at the apex: pedicel 1 mm.
long, 0.4 mm. in diameter, pubescent. Bracts 0.5-1 mm. long 0.4-0.8 mm.
broad, deltoid to reniform, apex obtuse or rounded; base obtuse, sessile;
pubescent on both surfaces. Sepals unequal, standing apart above the
middle, not enclosing either the flower or the fruit; first sepal 2.5-2.8
mm. long at anthesis,
at anthesis, usually slightly shorter than the first sepal; obovate, apex
obtuse, acute, or acuminate; tl
pubescent on the periphery, glabrous near the middle: third sepal
=
nate or acute: base obtuse: interior sepals 1.6-1.8 mm. long, 1.4-1.5
mm. broad at anthesis, the two often unequal; ovate, apex acute;
conspicuously thickened medially and basally; pubescent on both sur-
faces; margin ciliate. Petals ca. 3.5 mm. long, 3.0 mm. broad at anthe-
sis, obovate-oblong, apex truncate; glabrous; violet in color. Anthers
ca. 2 mm. long, slightly caudate: filaments 2.5-3 mm. long, subulate.
64 JOAB L. THOMAS
Ovary subglobose, 0.7-0.9 mm. in diameter at anthesis, more or less
sharply 5-ridged; pubescent to hirsute, the indumentum often forming
longitudinal striations. Style persistent, attenuate, ca. 1.5 mm. long,
glabrous. Fruit subglobose, 1.5 mm. in diameter, 5-ridged, not enclosed
by the persistent sepals.
DISTRIBUTION: Mountains of northern Oriente Province, Cuba.
SPECIMENS SEEN: CUBA. Proy. Oriente: Vicinity of Sierra de Moa;
Camp La Gloria, Shafer 8265 (type: NY; isotypes: NY, LS); Shrub 4
ft., rocky bank of river, vicinity of Camp San Benito, 900 m. altitude,
Shafer 4095 (US; NY, 2 sheets; F) ; Charrasco, en las cumbres de la
Sierra de Moa, Alain 3387 (Ls
Because of its extremely small leaves, Purdiaea micro-
phylla is a very distinctive species. It is closely related to
P. parvifolia which also has small leaves, but the two are
easily separated even in sterile material, by the larger, revo-
lute, obovate-spatulate leaves in the latter species. The
leaves of P. microphylla are ovate, with an abruptly acumin-
ate apex, and are much smaller than those of P. parvifolia.
The flowers of Purdiaea microphylla also afford several
characters by which this species can be distinguished from
other species in the genus. The sepals are open or slightly
in
reflexed in bud, and remain so after anthesis. Thus neither
the bud nor the fruit is enclosed by the sepals. The sepals
of Purdiaea microphylla are the smallest in the genus, and
show the least amount of differentiation between the ex-
terior and the interior sepals. In all other species of Purdiaea
the interior sepals are quite different from the exterior se-
pals. In the present species however, the interior sepals are
ovate, with a length-width ratio approximately the same as
of the exterior sepals. Moreover,Purdiaea microphylla
is the only species in which one of the interior sepals is
MONOGRAPH OF CYRILLACEAE 65
noticeably larger than the other. The unusual petals of
Purdiaea microphylla are almost square, being only slightly
longer than they are broad, with an abrupt truncate apex.
The variation within this species is very poorly known,
there being but one collection, of a single sheet, with flower-
ing material (Alain 3387). The variation in the leaves of the
available collections is within the range of variation found
on a single specimen. Although this species apparently oc-
cupies the same general area as Purdiaea parvifolia, I was
unable to locate it during my brief stay in the Sierra de Moa.
Future field work in this area will probably reveal that the
two species are ecologically, if not geographically, isolated
to acertain extent.
7. Purdiaea stereosepala Thomas, sp. nov. Type:
Thomas 516
Shrub or small tree 4-5 meters tall; stems smooth, glabrous, light
brown, becoming gray with age; leaf scars prominent, more or less
diamond shaped. Leaves dark green above, pale beneath; obovate to
obovate-spatulate; apex emarginate, truncate or occasionally rounded,
mucronulate; base attenuate; 4-6.5 mm. long, 1.5-2 mm. broad: mid-
vein prominent beneath, impressed or slightly prominent above; 6
prominent lateral veins running longitudinally, more or less parallel
to the margin of the leaf; the first two arising at the base of the mid-
vein and extending ca. 1/2 the length of the leaf; the next two arising
just above the first two and extending ca. 2/3 the length of the leaf;
the fifth and sixth arising alternately, well above the other four, ex-
tending the entire length of the leaf, usually connecting with the mid-
vein at the apex: shorter pinnate lateral veins forming interconnections
between the midvein and the longitudinal lateral veins. Racemes 6-7
cm. long, erect or occasionally bending near the apex: rachis attenu-
ate, 1-1.2 mm. in diameter at the base, tapering to 0.3 mm. in diameter
at the apex, villous or velutinous, the trichomes ca. 0.5 mm. long:
pedicel ca. 2 mm. long, 0.2-0.5 mm. in diameter. One to 3 large bracts
near the base of the raceme, 5-7 mm. long, 2-2.5 mm. broad; ovate with
66 JOAB L. THOMAS
pubescent, the exterior edge rounded and glabrous or nearly so; inte-
rior sepals lanceolate to lorate, apex rounded, 5-7 mm. long, 1-1.5 mm.
broad; thickened basally; pubescent on both surfaces, margins ciliate.
Petals 6-6.5 mm. long, ca. 3 mm. broad, ovate oblong; apex obtuse;
dorsal surface pubescent on the lower 1/3, glabrous above, ventral
surface glabrous. Anthers oblong, 3 mm. long, caudate, the cauda
acuminate, ca. 0.4 mm. long: filaments 2.5 mm. long, subulate. Ovary
globose, 5-ribbed, velutinous; style persistent, attenuate, 4.5 mm. long
at anthesis, becoming 5-5.5 mm. long at maturity. Fruit globose, 5-
ribbed, velutinous above the middle, ca. 3 mm. in diameter.
Frutex vel arbor, 4-5 m. alta; ramis planis glabrisque; foliis 4-6.5
em. longis, 1.5-2 em. latis, ebosaie vel obovato-spatulatis, apice sive em-
arginatis sive truncatis, interdum rotundatis cum mucronulo in medio;
venis lateralibus utrinque ternis, cum margine foliorum plus minusve
aes racemis 6-7 cm. longis; rhacide sursum attenuato, villoso;
bracteis racemi basin versus ovatis, magnis, supra reniformibus min-
prea sepalis rigidis, per anthesin scarioso-foliaceis, post anthesin
foliaceis, basi incrassatis, margine sese tangentibus; sepalis exteriori-
bus per anthesin ca. 8 mm. longis, 6 mm. latis, post anthesin 11 mm.
longis, 9 mm. latis, aequalibus, ovatis, pubescentibus; sepalo tertio
asymmetrico, margine interiori pubescent. margine exteriori glabro;
sepalis lanceolatis vel loratis, apice rotun-
datis, basi incrassatis, 5-7 mm. longis, 1-1.5 mm. latis; petalis ovato-
oblongis, ca. 6-6.5 mm. longis, 3 mm. latis anthera oblonga; filamentis
subulatis; ovario globoso, 5-loculato; stylo pertinaci, subulato.
DISTRIBUTION: Known only from the type locality.
@ PURDIAEA EKMANII
A P. STEREOSEPALA
@ P. SHAFER!
T
Sasi Sec Se
SPECIMENS Proy. Oriente: Vicinity of Moa; Rocky
shores of Rio aa Thomas 516 (Type: GH).
This distinct new species is the fifth Species of Purdiaea
known from the vicinity of Moa, and is the ninth species
from — Oriente Province, Cuba. It is closely related to
Purdiaea moaensis and P. velutina, and more distantly re-
lated to P. stenopetala and P. shaferi. Purdiaea stereosepala
MONOGRAPH OF CYRILLACEAE 67
is readily distinguishable from other species by several char-
acters. As alluded to by its name, the sepals are rigid and
thickened. The exterior sepals are coarse and foliaceous, and
meet along their lateral margins and often at the apex. The
first sepal is asymmetrical, flaring outward on the side to-
ward the rachis, slightly flattened on the side away from
the rachis.
Purdiaea stereosepala also differs from closely related
species by its smooth, glabrous vegetative shoots, and well-
differentiated, villous rachis. This characteristic is also
found in P. shaferi, but in the latter species the external
sepals are scarious, and very unequal in length — quite un-
like those of the present species. Also, the indumentum on
the peduncle is different in the two species. In Purdiaea
stereosepala the trichomes are unicellular and erect, where-
as in P. shaferi the trichomes are multicellular, quite long,
and usually somewhat appressed.
The pattern of leaf venation in the present species, with
p inent, longitudinal lateral veins arising well above the
base of the midvein, suggests some relationship with Pur-
diaea nipensis; although the leaves of P. stereosepala are
more coriaceous, more obovate, and have a more broadly
sessile base.
8. Purdiaea moaensis Marie-Victorin, Contrib. : Inst.
Bot. Univ. Montréal 63:60, 1948. Type: Marie-
Victorin, Clément, and Alain 21788
Shrubs or small trees up to 4 meters tall, 4-5 em. in diameter; bark
light gray or brown; young stems pubescent to velutinous toward the
broadly sessile; elongate-elliptic,
late; apex obtuse or rounded, mucronulate; 6-12 cm. long, 1-2.3 em.
broad, the upper leaves of a shoot glabrous, the lower leaves often
creasingly greater angles
interconnections
the lateral veins arise. Racemes rigidly erect, 7-9 cm. long at mid-
anthesis, becoming 12-13 em. long at maturity; transition from vegeta-
tive to reproductive shoot fairly abrupt, rarely a short zone of transi-
tion evident. Rachis robust, ca. 2 mm. in diameter at the base tapering
to ca. 1.0 mm. in diameter at the apex; ridged, densely clothed with
68 JOAB L. THOMAS
long, velutinous trichomes up to 1.5 mm. long. Pedicel short, ca. 1.5
mm. long, 0.5 mm. in diameter, velutinous, the articulate base and apex
obscured by the indumentum. Bracts lanceolate, up to 10 mm. long,
. broad near the base of the raceme, becoming smaller and
rounded, 2-3 mm. long, 1.2-2 mm. broad near the apex of the raceme.
Sepals prominently veined, the three outer sepals approximately equal
in length; first sepal 6-8 mm. long, 5-6 mm. broad at anthesis, 9-11 mm.
long, 7-7.5 mm. broad at maturity; ovate, apex acute or obtuse; pubes-
cent on both surfaces, particularly toward the base; margin ciliate:
second sepal slightly narrower than the first sepal; ovate, apex obtuse,
rarely acute, margin ciliate, dorsal surface pubescent below the middle,
becoming puberulent above; ventral surface glabrous or nearly so
above the middle, becoming puberulent to pubescent below. Third sepal
narrow, somewhat asymmetrical, the interior edge flattened basally,
puberulent, the exterior edge rounded, glabrous; apex acute, margins
ciliate: interior sepals lanceolate; apex acute to acuminate; somewhat
shorter and much narrower than the exterior sepals; dorsal and vent-
ral surfaces densely pubescent, a ridge of conspicuously longer tri-
chomes evident along the midvein on the dorsal surface; margin strong-
ly ciliate. Petals oblong, apex rounded or obtuse, 6-7 mm. long, 2-2.5
mm. broad, violet in color. Anthers bright yellow, oblong, 3-3.5 mm.
long, 0.5-0.7 mm. broad, distinctly caudate, the cauda bearing a small,
spheroid, glandular tip. Ovary 5-lobed, subglobose, densely clothed
with long silky trichomes. Fruit rounded to ovoid, velutinous, particu-
larly near the apex: style persistent, 5-6.5 mm. long at anthesis, be-
coming ca. 8 mm. long at maturity, exserted.
DISTRIBUTION: Vicinity of Moa, Oriente Province, Cuba.
SPECIMENS SEEN: CUBA. Prov. Oriente: Vicinity of Moa; Rio Yag-
— Punta ohpeig, Marie-Victorin, Clément, Alain 21788 (type:
MT; isotypes: SV); Acuha 12532 (US, sv); Alain, Clément, Chry-
sogone 3866 i con braiteas solas, Clément 3640 (LS, MT); d’en
haut, arbusto muy alto, casi arbol, Clément 3641 (LS, MT); Thomas
502 (GH); Thomas 505 (GH).
This species is known from only one locality, a small river
MONOGRAPH OF CYRILLACEAE 69
near the town of Moa. The plants are fairly abundant along
the rocky shores of this stream, however, and future collect-
ting will undoubtedly show that the species is more widely
distributed.
Purdiaea moaensis is closely related to P. velutina, and
the two species are occasionally confused, particularly in
sterile material. In both of these species the first leaves pro-
duced on a shoot develop into cataphylls, densely clothed
with long, silky, appressed trichomes. These peculiar struc-
tures are not found in any of the other species in the genus.
In Purdiaea moaensis the cataphylls are smaller and less
conspicuous than those of P. velutina, and in the former spe-
cies the cataphylls are usually shed shortly after flowering
begins. In Purdiaea velutina the cataphylls are very con-
spicuous structures and usually persist during most of the
flowering period. In both species the cataphylls are closely
spaced on the stem, and in early stages are somewhat strobi-
loid in appearance. Marie-Victorin (1948) in discussing
Purdiaea velutina described this species as having an “. ..
énorme bourgeon d’inflorescence ou les bractées velues com-
posent un espéce de strobile argenté . . .”.
Purdiaea moansis also resembles P. stereosepala and P.
shaferi in having very prominently veined, coriaceous leaves,
and densely pubescent racemes. The present species is usu-
ally distinguishable from related ones, however, by its elon-
gate-elliptic to elliptic-obovate leaves. It is distinguishable
from Purdiaea velutina and P. shaferi in having exterior
sepals that are approximately equal in length; and is distin-
guishable from P. stereosepala in having scarious exterior
sepals and densely pubescent vegetative shoots.
The leaves of Purdiaea moaensis, although distinguishable
from those of other species in the genus, are quite variable
in size and shape, even on a single individual. On one fairly
sizeable shrub (Thomas 505) I found leaves varying in shape
from elongate-elliptic to elongate-spatulate. On this same
shrub the mature leaves varied in length from six to eleven
cm. The flowers, on the other hand, are relatively uniform
on an individual plant, and show little variation from one
plant to the next in the few collections of flowering material
available.
9. Purdiaea velutina Britton and Wilson, Bull. Torrey
Club 42:389, 1915. Type: Shafer 4474
Shrub or small tree up to 4 meters tall, 3-6 cm. in diameter; bark
light gray, scaly; stems velutinous, particularly toward the ends; ro-
bust, growing in a zigzag pattern. Leaves dark green, shining above,
70 JOAB L. THOMAS
pale beneath; 4-9.5 em. long, 2.2-3 cm. broad, obovate-oblanceolate to
spatulate; apex broad, rounded, mucronulate or slightly emarginate;
more or less broadly sessile: midvein prominent on both surfaces,
broadening considerably near the base; 4-6 longitudinal secondary
veins arising at or near the base of the midvein, running more or less
parallel to the leaf margin; smaller pinnate lateral veins prominent
above the middle, forming interconnections between the midvein and
the prominent longitudinal lateral veins. The first leaves of a shoot
developing into a dense cluster of velutinous cataphylls. Racemes 3-5.5
n diameter, often partially enclosed by the subtending bract;
velutinous, the ind ding to that of the rachis. Bracts
near the base of the raceme lanceolate to ovate-lanceolate, ca. 5 mm.
long, those near the ase Me the ae Coors to reniform, ca. 2.5 mm.
long. Sepals ; white, pink, or rose-
colored, unequal: first a 11-12 mm. long, 5-6.5 mm. broad at an-
thesis, becoming 14-17 mm. long, 6-8 mm. broad at maturity; ovate,
ovate-elliptic or elliptic; apex acute, rarely slightly obtuse; dorsal
surface clothed with fine, silky trichomes below the middle; ventral
surface finely pubescent toward the base, glabrous or nearly so above;
margins sparsely ciliate: second sepal clearly shorter than the first,
7-9 mm. long, 4-5 mm. broad at anthesis, becoming 9-11 mm. long, 5-6
mm. broad at maturity; ovate, apex acute; more densely pubescent
than the first sepal, especially on the dorsal surface: third sepal slight-
ly longer and narrower than the second sepal, somewhat asymmetrical,
the interior edge flattened, particularly near the base; puberulent or
pubescent near the flattened margin on the dorsal surface, pubescent
or velutinous on the lower half of the ventral surface; margins ciliate:
interior sepals lanceolate with an acuminate apex; 3-4 mm. long, 0.8-1
mm. broad at anthesis, becoming 5-8 mm. long, 1.0-1.5 mm. broad at
maturity, dorsal and ventral surfaces velutinous with a ridge of con-
spicuously longer trichomes along the full length of the midvein on
the dorsal surface; margin strongly ciliate. Petals ovate to ovate
oblong, ca. 7.5 mm. long, 3.5-4 mm. broad, violet in color. Anthers ob-
long, 4-4.5 mm. long. Ovary sa velutinous, 5-lobed; style ca.
7 mm. long at anthesis, becoming up to 9 mm. long at maturity, ex-
serted.
. rome ai Mountains and lowlands of northern Oriente Province,
REPRESENTATIVE SPECIMENS SEEN: CUBA. Prov. Oriente: Vicinity of
Toa; Rio Yamaniguey to Camp Toa, on compact red iron ore, Shafer
4474 (type: NY) ; Vicinity of Moa; Charrascos, Rio Jaragua, Clément,
Alain 4020 (ts); Clément, Alain, and Chrysogone 4020 (LS); Rio
Cayoguan, zone riparienne, prés du pont de la mine Delta, Marie-
Victorin and Clément 21789 (us; MT, 3 sheets; GH); Marie-Victorin,
Clément and Alain 21789 (Ls); Marie-Victorin, Clément and Alain
21790 (MT) ; Marie-Victorin, Clément, Mr. and Mrs. Bucher 2609 (Ls) ;
MONOGRAPH OF CYRILLACEAE 71
along shores of Arroyo Jicotea, between Punta Gorda and Moa, Thom-
as 506 (GH); Thomas 507 (GH) ; Thomas 508 (GH) ; Camp San Benito;
margin of rocky river, small tree to 15 ft., Shafer 4092 (US, F, NY);
Shafer 4094 (US, NY, F); Camp La Gloria; tree 10-12 feet, Shafer
8261 (NY).
PURDIAEA
VELUTINA
6
} .
| Som
ese
rg y
ra If
This species is known from both the mountains and the
lowlands of northern Oriente Province. It usually occurs as
widely-scattered individuals along the rocky banks of small
rivers and arroyos.
Purdiaea velutina is closely related to P. moaensis and
Jc These: relationshi
more distantly to P. 'p ae
closely related.
The present species is rather uniform in its distinctive
characters such as the structure of the sepals, the venation
pattern in the leaves, and the structure of the cataphylls.
However, there is some variation in several characters. The
leaves are rather variable in shape and even more variable
in size. The mature leaves may be obovate-oblanceolate to
spatulate and range from four to nine centimeters in length.
The degree of prominence of the small, lateral veins which
arise above the middle of the leaf is also rather variable. As
in most species of Purdiaea the sepals, particularly the third
sepal, vary in the degree of pubescence. The sepals also show
72 JOAB L. THOMAS
a rather diverse range of color, often varying from white to
bright pink in plants that are only a few yards apart.
Purdiaea velutina is a very beautiful shrub, and would
likely be a popular ornamental in areas with a warm climate
if the plant were better known. The dark green, shining
leaves, the silvery, strobiloid cluster of cataphylls, the long
racemes of flowers with lavender petals, bright yellow sta-
mens, and persistent sepals which vary in color from white
to rose, are features which make this an unusually attractive
shrub.
10. Purdiaea shaferi Britton and Wilson, oo Torrey
Club 42:389, 1915. Type: Shafer 42.
Shrub 2-3 meters tall, stems glabrous, light brown, ase gray
with age. Leaves 4.5-9.5 cm. long, 2.5-4.5 em. broad, broadly elliptic-
obovate; apex mucronulate, rounded or ees base broadly ses-
sile: midvein and 6-8 longitudinal, lateral veins prominent on both
surfaces; a rather coarse reticulum of smaller veins prominent be-
to puberulent above. Racemes ca. 5 cm. long, erect; rachis robust, 2.5-3
mm. in diameter at the base, tapering to 1.0 mm. in diameter at the
apex, densely velutinous, the trichomes silky, 0.5-0.8 mm. long: pedicel
1.5 mm. long, 0.7 mm. in diameter; velutinous. Sepals scarious, very
unequal: first sepal broadly ovate to ovate-oblong, 12-14 mm. long,
7-8 mm. broad at anthesis; apex rounded or acute, apiculate; base
cuneate-attenuate; pubescent toward the base, the trichomes appressed,
silky: second sepal 8-9 mm. long, 4.5-5 mm. broad at anthesis; ovate,
apex acute; base cuneate-attenuate: third sepal approximately the
same length as the second sepal; asymmetrical, the interior edge flat-
tened: interior sepals lanceolate, 5-6 mm. long, 2 mm. broad; velutin-
ous, margin ciliate, apex acuminate. Petals elliptic, ca. 5 mm. long,
2.5-3.2 mm. broad, mucronulate, glabrous. Anthers 2.5-3 mm. long,
caudate, the cauda small and pointed: filaments ca. 3 mm. long, strong-
ly subulate. Ovary globose-ovoid, hirsute; style attenuate; fruit un-
known.
DISTRIBUTION: Known only from the type collection.
SPECIMENS SEEN: CUBA. Prov. onsen Vicinity of Baracoa; Pine-
lands back of town, iShsfor 4285 (type: NY
This poorly known species has hie been re-collected since
the original collection by Shafer in 1910. Purdiaea shaferi
is quite similar in vegetative structures to Purdiaea ekmanii.
This relationship i is discussed under the latter species. The
present species also resembles Purdiaea moaensis and P. vel-
utina in several characters including the silky trichomes on
the rachis, the rigidly erect, robust stems, and the very
prominent, longitudinal, lateral veins in the leaves. Purdiaea
feri is also similar to P. velutina in having unequal ex-
MONOGRAPH OF CYRILLACEAE 3
terior sepals. The present species is easily distinguished
from Purdiaea moaensis and P. velutina, however, by its
large, broadly ovate leaves, glabrous stems, and particularly
the absence of cataphylls, which are so characteristic of the
former species. The one available specimen of Purdiaea sha-
feri is in bud, just prior to anthesis, and there is no evidence
of any cataphylls on the specimen. This is the stage at which
the cataphylls are most prominent in Purdiaea velutina and
P. moaensis.
11. Purdiaea ekmanii Marie-Victorin, Contrib. Inst.
Bot. Univ. Montréal 63:61, 1948. Type: Ekman
15
Shrub, apparently small; branches robust, light gray to brown,
glabrous. Leaves crowded toward the ends of branches, giving a ros-
and fruits unknown.
DISTRIBUTION: Mountain tops, Sierra de Cristal and Sierra de Moa,
Oriente Province, Cuba.
SPECIMENS SEEN: CUBA. Prov. Oriente: Sierra de Cristal; In low,
rocky arthrostylidium thickets which cover the top, 1100-1325 meters,
Ekman 15981 (Photograph of type: NY; isotypes: LS, NY); Alain and
Lopez 4824 (Ls). Sierra de Moa; Charrascos, alt. 800 m. approx.
diaea, it is possible that P. ekmanii and P. shaferi are geo-
graphical varieties of the same species. The evidence for
this is insufficient at present, however, and a critical treat-
ment of this group must await further collections and field
wor!
Cliftonia Banks ex Gaertner f. Fruct. 3:246, 1805.
Type: C. monophylla (Lam.) Britton ex Sarg.
Mylocaryum Willd. Enum. Hort. Berol. 454, 1809.
Walteriana Fras. ex Endl. (in synonymy) Gen. 1413, 1841.
74 JOAB L. THOMAS
eautit or small Eos ‘ekotaee bagi polternate, entire, estipulate,
glabro ‘i Flowers in
ater or axillary Sone pea obovate-spatulate or spatulate,
one subtending each pedicel: bracteoles lanceolate or oblanceolate, two
borne on each pedicel: sepals 5 (6-7) imbricate in bud, small, deltoid,
persistent. Petals as many as the sepals, spatulate to obovate-elliptic:
stamens 10 in two whorls of 5, the outer whorl opposite the sepals, the
inner whorl shorter and opposite the petals: filaments flattened and
petaloid below the middle, terete above: anthers versatile, dehiscing
through longitudinal slits: ovary borne on a glandular, hypogynous
disk; ovary 3-5-locular, each locule containing a single ovule: stigma
massive, subsessile: fruits winged, indehiscent; endosperm cellular,
embryo straight, elongate.
The genus is composed of a single species, Cliftonia monophylla.
Cliftonia monophylla (Lam.) Britton ex Sargent, N.
‘Amer. Silva 2:7, 1892. Type: not seen, presum-
ably in the Lamarck Herbarium, Paris
Ptelea monophylla Lam. Illus. 1:336, 1789.
Cliftonia nitida Gaertn. f. Fruct. 3:247, 1805.
Mylocaryum ligustrinum Willd. Enum. Hort. Berol. 454,
1809.
‘ec ligustrina (Willd.) Sims ex Spreng. Syst. 2 :316,
25.
Evergreen shrubs or small trees usually 4-5 meters tall, 6-12 cm.
in diameter, branching near the base, rarely becoming 12 meters tall,
30 cm. in diameter. Bark reddish brown or gray, thick and spongy on
the trunks of idee trees. Young stems dark red, smooth, becoming
gray and scaly after 1-2 years. Leaves simple, alternate, entire, estip-
ulate, short-petioled, sub-coriaceous or coriaceous; light green, shining
above, pale, glaucous beneath; oblanceolate-elliptic or rarely obovate-
oblanceolate, apex acute or slightly emarginate, 3-6 cm. long, 1-2 em.
terminating
Pedicels 3-5 mm. long, 0.3-0.4 mm. in diameter, articulate at the point
of attachment to the rachis. Bracts 2.5-5 mm. long, 0.5-1.0 mm. broad,
— or occasionally pinkish, usually turning brown above the middle;
texture only slightly coarser than that of the
paar chossteepatnlale or spatulate, apex rounded, slightly concave
poe thickened basally, midvein obscure; articulate at the base, cad
sg ogemarnnced well before anthesis. Two bracteoles borne someaheriit
the pedicel, 1-2 mm. long, 0.2-0.4 mm. broad, lanceolate or rarely
see aaa usually shedding shortly after anthesis. Sepals 5 (rarely
6-7), white or pinkish, imbricate in bud, 0.5-0.7 mm. long, 0.6-0.8 mm.
broad, deltoid, apex acute or rounded; persistent, membranaceous at
anthesis, becoming foliaceous at maturity. Petals 5 (6-7), white or
pinkish, imbrieate in bud, spatulate to obovate-elliptic, occasionally
MONOGRAPH OF CYRILLACEAE 75
weakly clawed near the base, 3.5-5.5 mm. long, 1.5-2.5 mm. broad.
distinctly 3-veined, the veins running longitudinally. Stamens 10 in
two whorls, the outer whorl 3.5-4 mm. long, opposite the sepals, in-
serted lower on the receptacle than the inner whorl; the inner whorl
2.5-3 mm. long, opposite the petals; filaments laterally expanded and
petaloid below the middle, narrowing abruptly, becoming terete and
subulate above; the expanded lower portion 0.6-1.0 mm. broad; a single
conspicuous vein running the length of the filament; anthers versatile,
attached slightly below the middle, dehiscing by means of longitudinal
slits. Ovary borne on a small, slightly concave, glandular disk which
secretes a sticky fluid at anthesis; ovary ovoid or oblong, 3-5-angled,
3-5-locular, each locule containing a single, pendulous ovule; stigma
massive, sub-sessile, 2-5-lobed, the lobes corresponding to the number
of locules. Fruit 2-5-winged, dry, indehiscent, usually devoid of seeds,
but as many as 5 may develop in a fruit. Seeds consisting of an elong-
ate embryo with short cotyledons, surrounded by a mass of cellular
endosperm; seed coats lacking.
DISTRIBUTION: Coastal Plain of Southeastern United States.
REPRESENTATIVE SPECIMENS SEEN: UNITED STATES. Alabama: Bald-
win County: Route 31, 4 miles east. of Bay Minette, Coker (NY); Es-
cambia County: 2 miles south of Atmore, Blanton 1 (F, MT, NY, UC,
us); Mobile County: Spring Hill, along a water course, Graves 540
(mo). Florida: Bay County: Lynn Haven, St. Andrew’s Bay, Banker
3477 (NY); Calhoun County: Rt. 71, 1 mile north of Blountstown, S.
C. Hood, 1566 (FLAS); Clay County: Green Cove Springs, W. A. Mur-
rill (rLAS); Escambia County: Bayou Marcus Creek, west of Pensa-
cola, R. K. Godfrey 54590 (FLAS, UC) ; ‘Franklin County: swamps and
ponds in the pine barrens, Apalachicola, Chapman Herbarium 2004a
(us) ; Gadsden County: ad. fluv. “Little R.” prope Quincy, Rugel (Mo,
Ny); Gulf County: north of Port Saint Joe, Small, DeWinkeler, and
Mosier 11229 (NY); Holmes County: 2 miles S. W. of Bonifay, Ford
CLIFTONIA }MONOPHYLLA
76 JOAB L. THOMAS
3537 (FLAS); Jefferson County: Tyty Bay, Harper 131 (us); Liberty
County: Demaree 10143 (Mo, UC); Okaloosa County: near Laurelhill,
Palmer 38632 (Mo, Uc); Santa Rosa County: east bank of the Black-
water River, across from the town of Milton, Webster and Wilbur 3582
(us); Wakulla County: 35 miles S. W. of Tallahassee, along Road 319,
Deam 57745 (NY); Walton County: near De Funiak Spring, Curtiss
6381 (MO, NY, UC, US); Washington County: east of Holmes Creek
and north of Highway 280, Ford 3764 (FLAS). Georgia: Appling Coun-
ty: S.W. of Baxley, Steyermark 63344 (F); Chandler County: 10
miles N. W. of Rosemary Church, Hermann 10123 (F, NY, US); Clinch
County: 5.2 miles N.E. of Homerville, McVaugh 5294 (uc); Coffee
County: Seventeen Mile Creek, Harper 692 (F, MO, NY, Us); Early
County: along Temple Branch, 5 miles north of Blakely, Thorne 6818
(tT); Emanuel County: pine barrens near Graymont, Harper 811
(0, Ny, Us); Irwin County: along Alapaha River, 3 miles W. of
Irwinville, Hermann 10054 (F, MO, NY, Us); Liberty County: swamp
in and about the Alatamaha River, Small ( F, NY); Randolph County:
along west side of Nochaway Creek, Thorne and Muenscher 7898 (NY);
Tatnall County: sandhills 3 miles W. of Reidsville, Cronquist 4938
(FLAS, MO, NY, US); Toombs County: thickets along creek, near Lyons,
Palmer 38281 (F, MO, MT, US); Ware County: 2 miles east of Millwood,
Pyron and MeVaugh 1503 (r); Wayne County: 1 mile south of Rt.
#341 on State Road #23, Duncan 1990 (uc); Worth County: vicinity
of Poulan, Pollard and Maxon 539 (NY, US);. Mississippi: Harrison
County: along ravines in pine woods, Pass Christian,A. B. Langlois
(F, NY, Uc); Jackson County: pinelands east of Pascagoula, Small,
Mosier, and Matthaus 12808 (NY).
This species occurs only on the Coastal Plain of southern
Georgia, Alabama, Mississippi, and northern Florida. In
this area Cliftonia monophylla is found growing side by side
with Cyrilla racemiflora, but the former species has a much
narrower ecological tolerance, growing only in very wet,
sandy, acid soils. In contrast to Cyrilla racemiflora, Cliftonia
monophylla shows very little intra-specific variation from
one population to the next. There is a very minor degree of
variation in the size and shape of the stamens and petals,
but a range of variation comparable to that found through-
out the range of the species can usually be found on a single
individual. Thus in general aspect and in most characteris-
ties Cliftonia monophylla is rather uniform throughout its
limited geographic range. In view of the age of the genus,
the narrow phic distributi , and the lack of any ap-
preciable variation, it seems likely that Cliftonia is only a
relict form in the present-day flora.
Cyrilla Garden ex L. Mantissa Plant. 1:5, 1767. Type:
. racemiflora L.
Ttea L’Hérit. Stirp. Nov. 1:137, 1785.
Andromeda Marsh. Arbust. Amer. 9, 1785.
MONOGRAPH OF CYRILLACEAE T7
Shrubs or trees, leaves alternate, entire, estipulate, short-petioled,
dark green above, paler beneath. Flowers white or pinkish white,
borne in slender, axillary racemes. Pedicels ascending in bud, laterally
spreading at anthesis, borne in the axils of persistent bracts: 2 lance-
olate, persistent bracteoles on each pedicel. Sepals 5, ovate-lanceolate
to deltoid, acute, thickened medially and basally, coriaceous. Petals 5,
bl at late to oblong-elliptic, thickened and glandular on the
inner surface below the middle, inserted on a small, glandular disk
at the base of the ovary. Stamens 5, opposite the petals; anthers versa-
tile, dehiscing by means of longitudinal slits; filaments terete, subulate.
Ovary superior, sessile, 2-4-loculed: ovules pendulous, anatropous, 3
in each locule: style short, persistent, 2-4-lobed. Fruit dry, indehiscent,
often devoid of seeds; not more than one seed in each locule.
devoid of a seed coat; embryo cylindrical, radicle superior.
The genus is composed of a single species, Cyrilla race-
miflora.
Cyrilla racemiflora L. Mant. 1:50, 1767. Type: in the
Linnaean Herbarium. (Photograph of type: GH)
Itea Cyrilla L’Héritier, Stirp. Nov. 1:137, 1785.
Andromeda plumata Marshall, Arbust. Amer. 9, 1785.
Itea caroliniana Lam. Encyclop. Méthod. 3:315, 1789.
Cyrilla caroliniana Michx. F1. Bor.-Am. 1:158, 1803.
C. antillana Michx. l.c.
C. polystachia Raf. Autik. Bot. 8, 1840.
C. parvifolia Raf. 1.c.
C. fuscata Raf. l.c.
C. parvifolia Shuttl. Bull Torrey Club 23:101, 1896. (A
later homonym.)
C. perrottetii Brig. Ann. Conserv. Jard. Bot. Genéve 232,
C. brevifolia N. E. Brown, Trans. Linn. Soe. Ser. 2, 6:22,
1901
C. cubensis P. Wilson, Mem. Torrey Club 16:77, 1902.
C. racemiflora L. var. parvifolia Sargent, Jour. Arnold
Arb. 2:166, 1921. Based on C. parvifolia Shuttl.
C. arida Small, Bull. Torrey Club 51:383, 1924.
C. nipensis Urb. Fedde Repert. 22 :365, 1926.
C. nitidissima Urb. l.c.
iflora L. var. subglob Fernald, Rhodora 46 :46,
78 JOAB L. THOMAS
beneath; often clustered toward the ends of branches; 2-15 cm. long
including the petiole, 0.4-4.5 cm. broad; obovate, obovate-oblong, oblan-
ceolate, or oblanceolate-elliptic; apex acute, obtuse, rounded, or emar-
ginate; often apiculate; midvein prominent beneath, prominent or
depressed above; numerous lateral veins freely branching, forming a
dense reticulate pattern, usually prominent beneath, either prominent
or obscure above. Flowers in slender, glabrous, erect or pendulous ra-
cemes, borne near the ends of branches, in the axils of leaves or leaf-
scars of the previous growing season: rachis 4-26 cm. long, ca. 1.0 mm.
in diameter, articulate at the point of attachment to the rachis, as-
sli’ in bud, laterally spreading at anthesis, persistent. Bracts
lanceolate, concave, thickened basally, persistent, 0.4-4.1 em. long, 0.2-
0.6 mm. broad, longitudinally traversed by a single, obscure vein.
bud, spreading after anthesis, alternate or opposite, 0.4-1.3 mm. long,
0.2-0.4 mm. broad, lanceolate, persistent, somewhat thickened basally.
Sepals 5 (rarely 6), free or occasionally coalescent laterally at the
extreme base; persistent, ovate-lanceolate to deltoid, apex acute,
medially and basally thickened, 0.7-1.8 mm. long, 0.4-0.9 mm. broad;
imbricate in bud. Petals 5 (rarely 6), white or pinkish white, inserted
on a small, glandular disk; oblong-lanceolate to oblong-elliptic, slightly
concave; apex acute, obtuse or rounded; membranaceous laterally and
above the middle, thickened and glandular on the inner surface medial-
ly and below the middle. Stamens 5, opposite the petals, 1.4-2.5 mm.
long, shorter than the petals; filaments terete, subulate; anthers versa-
tile, attached near the middle on the dorsal side; 2-lobed, the lobes free
below the point of attachment to the filament, united above; usually
a small apical protuberance between the two lobes; pollen shed through
longitudinal slits. Ovary superior, sessile, 2-3, rarely 4-locular, ovules
pendulous, anatropous, 3 in each locule; stigma 2- to 4-lobed, corres-
ponding to the number of ques Fruit dry, indehiscent, subglobose,
globose, ovoid, ‘id-ob] bisulcate or
trisuleate, 1.6-4.9 mm. in pain _ devoid of seeds, but with as
many as 4 seeds developing; not more than one developing in each
locule; seeds pendulous, elongate, devoid of a seed coat; endosperm
cellular, abundant; embryo elongate, cylindrical, the cotyledons short,
inferior, the radicle superior.
DISTRIBUTION: Coastal Plain of Southeastern United States; Middle
America; Northern South America.
REPRESENTATIVE SEEN: UNITED STATES. Alabama. Es-
cambia County: McCullough, Blanton 16 (MT, NY, UC, US); Mobile
County: Spring Hill, Bush 43 (Ny, Us); Conecuh County: 3 miles
west of Evergreen, Crawford and Harvill 5738 (TEx) ; Russell County:
2 miles W. of Phoenix City, Duncan 9627 (Mo); Lee County: Auburn,
Earle and Baker (MO, NY, US); Baldwin County: Fairhope, Jack 3040
(A); Montgomery County : Vicinity of Montgomery, G. McCarthy s.n.
(A); Barbour County: 3 miles north of Eufaula, Wiegand and Man-
ning 1869 (GH). Florida. Wakulla County: en W. W. Ashe s.n.
(Ny); Seminole County: from Wagner to Oviedo, M. F. Baker (FLAS) ;
Okaloosa County: near Crestview Blanton 6576 (Mo, us); Franklin
MONOGRAPH OF CYRILLACEAE 79
County: near Apalachicola, Curtiss 1774 (F, MICH, NY, US); Duval
County: near Jacksonville, Curtiss 4280 (NY, SV, UC, US); Levy Coun-
ty: Gulf Hammock, J. H. Davis (FLAS); Gulf County: 3 mi, N. o
Indian Pass, Ford 3245 (FLAS); Holmes County: east bank of Chocta-
whatchee River at bridge of Highway #2, Ford 3473 (FLAS) ; Jefferson
County: swamps, A. S. Hitchcock sn. (F, MO); Gadsden County: 3
mi. s.w. of Quincy, G. M. Hocking s.n. (FLAS); Escambia County:
Bayou San Marcus, west of Pensacola, Kral and Godfrey 6029 (MT);
Baker County: McClenny, Lighthipe 588 (FLAS, NY) ; Osceola County:
Lake Wilson, McFarlin 5660 (A, MICH, TEX); Alachua County: S. of
Cox Bridge, Murrill 407 (us); Columbia County: Lake City, Nash
2218 (F, GH, FLAS, MICH, MO, NY, SV, US); Calhoun County: along
stream, Rock Bluff, O’Neill 6158 (Mo); Indian River County: Indiap
River, E. Palmer 325 (F); Leon County: small creek near Tallahassee,
E. J. Palmer 35198 (A, NY); Highlands County: between Avon Park
and Sebring, J. K. Small, J. W. Small, J. B. DeWinkeler 11486 (GH,
MICH, MO, NY, TEX). GEORGIA. Upson County: S. E. of Woodbury,
Cronquist 5502 (FLAS, GH, NY, UC, US); Tattnall County: 7 miles W.
of Claxton, Duncan 2727 (FLAS, GH, UC, US); Early County: 10 mi. S.
of Blakely, Hardin 147 (FLAS, MICH, TEX); Sumpter County: Leslie,
Harper 1392 (A, F, MO, NY, US); Thomas County: near Thomasville,
80 JOAB L. THOMAS
Small (F, NY); Baker County: along Ichawaynochaway Creek, across
bridge from Field Station, Thorne and Ford 1953 (Us); Wilcox Coun-
ty: 12 miles N. W. of Abbeville, Wilbur and Webster 2714 (GH, US) ;
Bulloch County: 6 mi. N. W. of Statesboro, Wilbur and Webster 2764
(GH, Us). LOUISIANA. Washington Parish: 10 mi. W. of Bogalusa,
Brown 5631 (A); St. Tammany Parish: vicinity of Covington, Arséne
12234 (US); near Akita Springs, Pennell 4194 (NY); Tangipahoa
Parish: 1 mile W. of Robert, D. S. Correll and H. B. Correll 9302 (GH,
NY). MISSISSIPPI. Stone County: south of Wiggins, L. G. Brenner
(mo) ; Jackson County: Ocean Springs, Demaree 28118 (TEX); Harri-
son County: Rt. 49, Bayhead, N. of Saucier, Hood 4646 (FLAS) ; Jones
County: Ellisville, Tracy 3373 (NY); Forrest County: 2 miles S.E. of
Hattiesburg, Webster and Wilbur 3377 (GH, Us). NORTH CAROLINA.
Perquimans County: along river, Hert Ford, Bartley and Pontius 491
(Ny); Anson County: 7 mi. s. of Lilesville near Jones Creek, Boyce
1037 (GH); Craven County: Riverdale, Brown 2359 (TEX); Halifax
County: Weldin, Canby (Fr); Martin County: low pocosin woods near
Jamesville, Correll 1857 (a); Hyde County: moist woods near Swan-
quarter, Correll 1765 (A) ; Harnett County: Buie’s Creek, Foust (NY);
Moore County: 1/2 m. S. of High Falls, Fox 2539 (GH) ; Gates County:
on Rt. 158, 2 1/2 miles E. of Chowan River, Fox and Whitford 350,
(uc); Onslow County: Euelaville, Godfrey 4467 (US); New Hanover
County: shrub savannah at Carolina Beach, Godfrey and Shank 4208
(GH, US) ; Robeson County: near aoa ing: Heller 14015 (MO) ; Greene
County: Snow Hill; L. F. and F. R. Randolph 774 (GH); Richmond
County: 3 mi. N. of Hoffman, Wiegand and Manning 1869 (GH); Pitt
County: 1 mi. E. of Grimesland, Wiegand and Manning 1868 (GH);
Columbus County: 1.5 mi. N.E. of Nakina, Wilbur 4210 (GH) ; Pam-
lico County: 2 mi. E. of Hobucken, Wilbur 2897 (GH); Bladen County:
east rim of White Lake, Woods (uc). SOUTH CAROLINA. Aiken
County: lakes near Graniteville, Eggert (Mo, us) ; Georgetown County:
10 miles northwest of Georgetown, Godfrey and Tryon 736 (GH, MO,
NY, UC, Us) ; Clarendon County: 3 mi. S.W. of Manning, Godfrey and
Tryon 932 (GH, NY, US); Charlestown County: 6 mi. N.W. of McClel-
lanville, Godfrey and Tryon 1130 (GH, NY, US); Lexington County:
Manning 1870 (GH). TEXAS. Hardin County: Saratoga, Bail
(us); Newton County, 5 miles _ of Kirbyville, Cory 49791 (Ny, UC,
us) ; Tyler County: Woodville, G. L. Fisher (F, Us); Jasper County:
Jasper, Stark 8570 (FLAS, TEX, uc, us); Polk County: in Big Thicket,
B. C. Tharp (MIcH, Mo, TEX). VIRGINIA. Southampton County:
Franklin, Eggleston 4917 (mo, NY); Dinwiddie County: woods west
of Winfred’s Mili, Fernald and Long 13676 (GH); Norfolk County:
eastern side of Great Dismal Swamp, north of Wallaceton, Fernald
and Long 13676 (GH, MO); Nansemond County: east of Cox Landing,
Fernald and Moore 15113 (GH, MO, US).
MEXICO. Oaxaca: near pc W. Nelson 794 (US). BRITISH
HONDURAS. El Cayo District: Rio Privacion, Bartlett 11788 (F, MICH) ;
MONOGRAPH OF CYRILLACEAE 81
Mountain Pine Ridge, Lundell 6883 (MICH); Temash River, Schipp
1356 (F, GH, MICH, MO, NY). CUBA. Prov. Oriente. Sierra de Maestra:
summit of Pico Turquino, Acufia 6750 (NY); Loma del Gato, Acufta
9834 (SV); alto de Comejin, Ekman 9353 (NY); alto de la Valenquela,
Lopez 2256 (LS, SV, US) ; Cordillera de la Gran Piedra, Lopez 2766 (Ls,
sv); Aserradero de Fresneda, E. E. Smith 256 (sv); Jiquarito Mt.,
Taylor 502 (Nx); Vicinity of Moa: Pinares, Acufia 12523 (SV, US);
cerca de Cayo Fortuna, Alain 3311 (LS); arroyito, en el camino de
Cayo Chiquito, Clément 3645 (LS, MT); Rio Yagrumajes, Clément and
Alain 3872 (Ls); Rio Cayoguan, cerca del puente de la Mina Delta,
Clément 23303 (LS); Charrascal del Coco, Léon, Clément and Alain
22620 (LS); chemin de Cayoguan, Léon, M.-Victorin and Clément
20831 (LS, MT); entre le Rio Cabofios et le Rio Moa, M.-Victorin and
Clément 21780 (MT); Bosque de Centeno, M.-Victorin, Clément and
Alain 21784 (LS, MT) ; vicinity of Camp San Benito, Shafer 4067 (NY);
alluvial valley of Rio Yamaniguey, Shafer 4236 (NY, US); Camp La
Gloria, Shafer 8032 (GH, NY, US); Sierra de Nipe: Pinar Mayari, Ar-
royo Naranjo, Carabia 3621 (NY); Cayo del Rey, Rio Canapu, Carabia
4087 (MT, NY); in carrascales, Ekman 2168 (NY); cabezades del Ar-
royo Guaro, Léon 19171 (LS); Cerro de Miraflores, Léon 21135 (LS,
MT); chemin de la mine Woodfred, Léon and Alain 19171 (Ls, MT);
Cayo de Monte, al pie de la Mensura, Léon, M.-Victorin, Clément,
Alain 19925 (LS); Vicinity of Baracoa: banks of Rio Janco, Léon
11755 (Us); Cabuebaje, Finca la Maria, Thomas 501 (GH). Prov.
Pinar del Rio: Loma de Cajalbana, Le Palma, Alain and Clément 1461
(1s) ; banks of Rio de las Vueltas, foot of Cajalbana, Léon and Charles
4931 (LS, NY); Sierra del Rosario, Alain and Ponce de Léon 103, (LS) ;
dry sandy soil, Herradura, Shafer 436 (NY, SV); along stream, vicinity
of Herradura, Britton, Britton, Earle and Gager 6490, (NY); Sierra
Rangel, Loma Pelada, Léon 12534 (MT); Arroyo del Sumidero, Shafer
and Léon 13627 (MO, NY, US); Bahia Honda, Léon 17088 (LS, MT).
ISLE OF PINES. near Nueva Gerona, Curtiss 490 (F, MO, NY, SV, US) ;
Rio del Indos, Jennings 436 (GH, NY, US); savanes de Los Indos, M.-
Vietorin and Alain 93 (GH, MT); Arroyo del Hatillo, Léon and M.-
Victorin 17825 (LS); San Juan, Roig and Cremata 1814 (Ny).
WEST INDIES. JAMAICA. St. Thomas: crest of Gossamer Peak,
Mazon 9249 (GH, US); Prospect Hill, Thompson 7947 (F, NY); Port-
land: southeast slope of Caledonia Peak, Proctor 15549 (GH); St.
Andrews: New Haven Gap, vicinity of Cinchona, Britton 192 (NY);
near Cinchona, Harris 9117 (F, GH, NY, US); mist forest on track to
Old England, West and Arnold 690 (FLAS, GH); Gordon Town, Hart
890 (F, US); 2 miles northwest of Hardwar Gap, Webster and Wilson
4937 (GH); Trelawny: 4 miles west of Troy, Maxon 2879 (NY, Us);
Crown Lands, near Troy, Harris 8725 (F, NY). HAITI. Massif du
Nord, Pors-Margol, Ekman 2813 (Us); Massif de la Hotte, western
group, near Dutreuil, Ekman 10764 (Us); road from Camp No. 1 to
82 JOAB L. THOMAS
La Barriere Couchant, Nash and Taylor 1107 (Ny); Riviere Glacu,
Holdridge 2226, (us). DOMINICAN REPUBLIC. Prov. Monti Cristi:
Mancion, Lagunas de Cenobi, Ekman 12877 (GH, US). Prov. Santiago:
road to Loma Bajita, Jicome, Valewr 726 (F, GH, MICH, MO, NY, Us);
en los alrededores del Pico del Gallo, J. Jiminez 2515, (Us). PUERTO
RICO, in sylvis montanis ad Sta. Isabel, Sintenis 6167 (F, GH, MO, NY,
SV, Us); El Duque, Sierra de Naguabo, Shafer 3657 (F, GH, MO, NY,
US); Rio de Maricao, Britton and Cowell 4220 (Ny, US); Maricao,
Sargent 502, (us); San Sebastian, Sargent 353 (us); El Verda, Lu-
quillo Mountains, Blomquist 11907 (TEX); near Mayaguez, Heller 4582
(F, GH, MICH, MO, Ny, sv); Mt. Ategrillo, Britton, Stevens and Hess
2611 (NY, US). GUADELOUPE. Soupriére, Questel 1666 (us) ; Source
du Galion, Chemin Soupriére, Stehle 387 (us). DOMINICA. Solfatara
area near Soufriére, Howard 11782 (GH, US); Soufriére, Lloyd 16
(xy). ST. VINCENT. in elfin woodland, Letchwood Valley, Beard
614 (GH, MO, NY, UC).
SOUTH AMERICA. BRITISH GUIANA. Mt. Roraima: summit, Tate
401 (NY); Ipelemorita, Arapos River, Jenman 35 (us); Camp River,
Jenman 1861 (NY); Pomeroon District, Mora Landing, Moruka River,
Cruz 1882 (F, GH, MO, NY, US); upper Mazaruni River, Cruz 2080 (GH,
MO, NY, UC, US); Ituni Road, Mackenzie, Demerara River, Fanshawe
5242 (NY); Rockstone, Gleason. 486 (GH, NY, US); Kaieteur Plateau,
Maguire and Fanshawe 23106 (F, NY, us). COLOMBIA. Rio Kanan-
ari, Cerro Isibukuri, Schultes and Cabrera 15058 (us); Rio Kuduyari,
Cerro Yapoboda, Vaupés, Schultes and Cabrera 14472 (us). VENE-
ZUELA. State of Bolivar: Sarven-tepui, Wurdack 34099 (NY); Plato
arriba da Auyantepui, Vareschi and Foldats 4954 (Ny); Sororopan-
tepui, Steyermark 60147 (F) ; Mount Roraima, Steyermark 58763 (F,
NY, US); Cerro Guaiquinima, Rio Paragua, Maguire 32867 (NY);
Amazonas: Ma:
NY); Cerro Yapacana, Rio Orinoco, Maguire, Cowan and Wurdack
30664 (F, NY); Cerro Huach i, Rio C Maguire,
Cowan and Wurdack 30216 (Ny, UC, US); Serrania Paru, Rio Paru,
Cano Asisa, Rio Ventuari, Cowan and Wurdack 31231 (ny). BRAZIL.
Prope San Carlos, ad Rio Negro, Spruce 2999 (GH, NY); Rio Curi-
curiari, middle course, Rio Negro, Schultes and Lopez 9718 (GH, US) ;
Mt. Roraima, on savannas, Tate 231 (NY); Mt. Roraima, it,
Tate 387 (Ny).
An extraordinary of variation is found in the genus
Cyrilla. It is so variable, in fact, that on the basis of various
extreme forms, eleven species and several varieties have
been described by previous authors, However, on studying
the material in the field over a wide geographic and ecologi-
cal range, and on bringing together for examination most
MONOGRAPH OF CYRILLACEAE 83
of the major collections, it has become apparent that the
genus Cyrilla should be considered as a single species, C.
racemiflora.
Several aspects of the nature of the variation in Cyrilla
are discussed above under the section on asexual reproduc-
tion and variation in Cyrilla racemiflora. The following
discussion of the variation in this group is primarily des-
criptive.
There are several patterns of variation within this species,
but there are no real gaps or discontinuities separating one
pattern from the next. Instead, the different patterns of
variation are connected by intermediate forms. Studies were
made of many of the complex variation patterns that occur
within and between populations of this species throughout
most of its geographical range. These studies have been
based on mass collections in a few instances, but primarily
on herbarium material. The use of mass collections or simi-
lar sampling methods for studying the variation within a
given population was found to have certain limitations due
to the widespread occurrence of vegetative reproduction in
the group. Within a population of Cyrilla, particularly a
small population, there will usually be relatively few biologi-
cal individuals. Thus in examining the variation found in
such a population one is more often than not dealing with
the ecological variation within a single clone. Moreover,
large populations of Cyrilla are exceptional. It usually occurs
as scattered groups of bushes which frequently can be shown
to be a single clone. This sporadic occurrence of clones is
particularly characteristic along stream banks and wooded
areas. The larger populations are usually found in open
fields, cut-over hillsides, or along highway right-of-ways —
areas where young sprouts from the roots will not be shaded
or crowded out by other competition. Populations under
these conditions are often somewhat weedy in nature.
The occurrence of large clones proved to be of some advan-
tage, however, in studying the variation found in a single
individual. The variation that may occur within a single
genotype can be assessed more accurately by studying a
large clone than by studying a single plant. The mass col-
lections used are of two types: those in which material was
collected from a number of plants that were known to con-
stitute a single clone; and those in which no two collections
were made within 60 feet of each other. In the latter type
it was still not possible to be certain that each sample col-
lected represented a different individual, thus limiting their
84 JOAB L. THOMAS
value as random samples of a population. Familiarity with
the plant under field conditions made it possible in many
instances to select different individuals by their appearance,
but certainly samples of this type are not random. Never-
theless, with the large number of herbarium specimens avail-
able, in addition to special collections made during two
summers of field work, the sampling at worst should be of
significance.
In studying the variation in Cyrilla, primary emphasis has
been placed on the size, shape, and texture of the leaves.
There were several reasons for this: 1) most of the species
which have been described have been segregated on the basis
of leaf size or shape; 2) leaves are the most variable part
of the plant; and 3) leaf length proved to be a very con-
venient basis on which to compare other, less variable char-
acteristics.
Geographically, three areas were of particular interest in
studying the variation in the group: eastern Cuba, northern
South America, and southeastern United States. These areas
will be
EASTERN CUBA
The highest degree of local variation in Cyrilla was found
in the mountains and stream valleys of northern Oriente
Province, particularly in the Sierra de Moa. Here, within an
area of 10 square miles, one can find individuals that closely
resemble those found throughout the range of the species.
A glance at the divergence in leaf size and shape, shown in
Fig. 24, will give some idea of the degree of variation found
in the leaves of plants from this region. The leaves shown
in this plate were taken from specimens collected within
a small geographical area in the Sierra de Moa. They repre-
sent, as nearly as was posible to determine, average, mature
leaves. In each case, the leaves shown with the adaxial side
up were selected as being average for the specimen, and
those with the abaxial side up were selected to show some of
the variation that occurs among the mature leaves of a given
specimen. arrangement of the leaves was intended pri-
marily to show two obvious trends of variation from the
average leaves shown at the top to the small form in row A,
and the large form in row B.
The effects of the environment are rather clear in the
extreme forms. Moisture and shade seem to be the most
important ecological factors influencing leaf size in Cyrilla.
The large-leaved forms, shown in the lower third of row B
are invariably found growing along stream margins and
MONOGRAPH OF CYRILLACEAE
a
es Of 1 | Bi, By RE BO
LEAF LENGTH mw.
in the pine flatwoods. Further explanation in text.
leaf length were found to differ less between the two pop-
ulations than was expected, and the standard deviation was
quite high in each case. For the population in the swamp,
the mean leaf length was 44.6 mm. with a standard deviation
of 15.7; whereas in the collections from the pine flatwoods,
the mean was 28.06 mm. with a standard deviation of 10.5.
There is clearly a large overlap in leaf size between the two
populations, but they are significantly different.
These two populations also differed significantly in flow-
ering time, the large-leaved population being at mid-anthe-
sis, while the small-leaved population was not yet in flower.
The flowering time in Cyrilla extends over a relatively long
period of time, beginning at the base of the raceme and pro-
gressing toward the apex. Because of this it is possible to
recognize several arbitrary stages during the time of flower-
ing. In comparing the populations under consideration, four
arbitrary stages of flowering were recognized between the
two extremes. In this manner flowering stage and leaf length
102 JOAB L. THOMAS
can be compared and contrasted in the intermediate popu-
lation from the artificial levee. One aspect of this relation-
ship is shown in Fig. 39 in which flowering stage is plotted
along the ordinate and leaf length along the abscissa. The
int di and di 1 of the two characters are quite
apparent in this diagram. Plants in late stages of flowering
have small leaves, and conversely plants in early flowering
stages often have large leaves. Similar results are obtained
by using leaf length-width ratio as the abscissa (Fig. 38).
In this area it is apparent that the two different forms
occupying two different habitats were, to a certain extent,
thrust together by the intervention of man in dredging the
Intercoastal Waterway. The interbreeding of two popula-
tions that resulted from this has given rise to a highly vari-
able, int diat pulati In terms of plant generations,
this population was relatively young and there had not been
sufficient time for any appreciable blending of the various
extremes. The fact that the intermediate population tended
to be graded toward the two extremes indicates that some
backcrossing had occurred, but the different forms were
intermixed for the most part.
In other areas, however, one can find more uniform pop-
ulations in various stages of intermediacy between the two
extreme forms. One such population was found in Walton
County, Florida, approximately 3 miles west of Santa Rosa,
near Highway 98. In this area the plants showed some vari-
ation in flowering time and leaf size, although the variation
here was considerably less than that found in the plants
from the intermediate population described above. The pop-
ulation as a whole appeared more like the small-leaved form,
although it was clearly intermediate between the extremes,
and a few individuals were found which appeared more like
the long-leaved form. Eastward from this area the popula-
flowering period ; those to the west had larger leaves and
A similar situation was observed in a population near the
Alabama-Florida line on United States Highway 90. This
population was also intermediate, but in this area the varia-
MONOGRAPH OF CYRILLACEAE 103
tion pattern was more like the large-leaved form. Likewise,
the variation pattern in the plants from this area was gr:
from west to east in a manner similar to that described
above. The intergradation was not so evenly graded in this
area as in the population near Santa Rosa, possibly because
the former population was not continuous, but rather broken
up into numerous smaller populations. Nevertheless, the
tendency toward larger leaves and earlier flowering time
was apparent in plants growing west of this area.
The habitat seems to be highly significant in determining
the over-all character of these intermediate populations.
The extreme variation found in the population on the arti-
ficial levee along the Intercoastal Waterway indicates that
the effects of the habitat must be primarily that of selecting
well-adapted gene combinations, and secondarily that of eco-
logical modification of a given genotype.
The other named species of Cyrilla from the United States
is Cyrilla arida Small. This species was described from
plants collected from Highlands County, in south-central
Florida. According to Small (1924) this species “ .. . differs
from C. parvifolia in the narrower sepals and petals and the
notched anthers, as well as in the vine-like branches.”
At the time this species was described, it was thought to
be geographically isolated from the Cyrilla in northern Flo-
rida, but subsequent collections have revealed that scattered
populations of Cyrilla occur in several areas between central
and northern Florida, (Map 11). Apparently there was also
some confusion as to the habitat of “Cyrilla arida’. It was
said to grow high up on dunes of dry, white sand. I have
made an intensive search in the vicinity of the type locality,
between Avon Park and Sebring, Florida, and have found
no plants of Cyrilla growing out on the sand dunes. In this
area, as elsewhere in Florida, Cyrilla occurs only in low
moist areas.
It seems likely that the designation of “Cyrilla arida” as
a distinct species was due in part to this confusion in habitat
and distribution. The petal and sepal shape, as well as all
other characteristics of this “species”, fall well within the
variation pattern found in plants of Cyrilla from northwest
Florida. The notched anthers, as mentioned above, occur in
collections of Cyrilla throughout its geographic range. Plants
with small, “vine-like” branches are also found in other
areas. Moreover, this character is not found in all of the
collections from Highlands County, Florida. In short, the
small-leaved representatives of Cyrilla in south-central Flo-
104 JOAB L. THOMAS
rida are not significantly different from the small-leaved
representatives in northern Florida, and the latter inter-
grade with the large-leaved forms in the same area.
CYRILLOPSIS
The genus Cyrillopsis Kuhlmann, which was included in
the Cyrillaceae by Mattick (1935) and Uphof (in Engler
and Prantl, 1942) is excluded from the present treatment.
This is done primarily on the basis of the structure of the
fruit, the pollen, and the wood anatomy. Although no mature
fruits of Cyrillopsis have been seen, it is readily apparent
from the immature fruits that the structure is unlike that of
members of the Cyrillaceae. The fruits of Cyrillopsis are
elongate, flat, and asymmetrical in shape, and contain seeds
with well-developed seed coats. Seed coats are not developed
by members of the Cyrillaceae. The pollen grains of Cyrillop-
sis are also very different from those of the Cyrillaceae. As
mentioned above in the section on pollen morphology, the
pollen grains of Cyrillopsis are similar to those of some mem-
bers of the Celastraceae, but show none of the structural fea-
tures that are characteristic of members of the Cyrillaceae.
Anatomically, Cyrillopsis differs from members of the Cyr-
illaceae in having much more advanced wood structure. As
shown in Fig. 42 and 43 the vessel elements of Cyrillopsis are
short and broad. The end walls of the vessel elements are
completely perforate, and fall into the category of group III
or group IV vessels according to the classification of Bailey
and Tupper (1918). The imperforate tracheary elements
are composed of libriform fibers and very thick walled fiber-
tracheids. No tracheids have been observed in the wood of
Cyrillopsis.
The present genus also differs from members of the Cyr-
illaceae by several other characters, including its exserted
stamens with non-versatile anthers and filaments that are
imbricate in bud; its persistent, cup-shaped disk on which
the anthers, but not the petals, are borne, and its leaf vena-
tion with alternating strong and weak lateral veins.
Superficially Cyrillopsis resembles the genus Itea, of the
Saxifragaceae. The resemblance is so close, in fact, that an
occasional sheet of Itea virginica is found misidentified as
Cyrillopsis paraensis. However, the morphology of the flower
and particularly the fruit is unlike that of members of the
105
MONOGRAPH OF CYRILLACEAE
5D ae Se
106 JOAB L, THOMAS
racemiflora. 44. Pine flatwoods near Covington,
ez 2
ny
near Po: ula, Louisiana.
MONOGRAPH OF CYRILLACEAE 107
Saxifragaceae, but does show certain similarities to members
of the Celastraceae. Also, the leaf venation and the pollen
morphology indicate a possible relationship with the Celas-
traceae. The wood anatomy, on the other hand, is more ad-
vanced than that found in most members of the Celastraceae.
Additional collections of plants with mature fruits, and
further study are needed before this genus is assigned to the
proper family, but provisionally I would suggest that Cyr-
illopsis should be included in the Celastracaee.
SPECIES EXCLUDED
Cyrilla aquatica Roxb. Pl. Corom. 2:47, 1795 = Limnophila racemosa
Benth. in Wall. Cat. n. 3907.
Cyrilla indica Koen. ex Wight and Arn. Prod 1:364, 1834 = Vahlia
oldenlandoides Roxb. Hort. Beng. 86: Fl. Ind. 2:89, 1832.
Cyrilla goo Koen. ex Wight and Arn, l.c. = Vahlia olden-
g pralees Roxb. 1
S seratas Nutt. in Am. Journ. Sc. 5:290, 1822 = Ardisia
eee Torr. and Gray ex D.C. Prod. Syst. Veg. 8:124, 1844.
Cyrilla spinosa Spreng. Nov. Prov. Hort. Hall. 15, 1831 = Bursaria
spinosa Cav. Icon. Descrip. Plant. 4:30, 1797.
Cyrilla viscosa Koen. ex Wight and Arn. Prod. 1:364, 1834 = Vahlia
viscosa Roxb. Hort. Beng. 86: Fl. Ind. 2: 89, 1832.
Cyrilla pulchella L’Hérit. Stirp. Nov. 147, 1785 = Achimenes coc-
cinea Pers. Syn. Plant. 2:165, 1807.
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____ "ann W. Spackman. 1949. A preliminary study of
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BRITTEN, J. 1905. Note on the history of Cliftonia. Jour. Bot. 43:
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BRITTON, N. L. 1899. Index to recent American botanical literature.
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Brown, N. E. 1901. Report on two botanical collections from Mt.
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CoPpELAND, H. F. 1953. Observations on the Cyrillaceae particularly
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Eviort, S. 1821. A Sketch of the Botany of South Carolina and
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ErRDTMAN, G. 1952. Pollen Morphology and Plant Taxonomy. Stock-
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FERNALD, M. L. 1950. Gray’s Manual of Botany (eighth edition).
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Gite, E. 1892. The Cyrillaceae. In Engler and Prantl, Die Natiir-
lichen Pflanzenfamilien (first edition). 3. 5:179-182.
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MONOGRAPH OF CYRILLACEAE 109
Hitcucock, A. S. 1935. Manual of the grasses of the United States.
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KUHLMANN, J. G. 1925. C ibuiga hecimento de algu-
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ee G. H. M. 1951. Taxonomy of Vascular rok New York,
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MARIE-VICTORIN, FRERE. 1948. Nouvelles études taxonomiques sur la
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MATTICK, FR. 1935. Die Gattung Purdiaea Planchon sb Viren eae
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MONOGRAPH OF CYRILLACEAE 1i1
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APPENDIX
TO COLLECTION NUMBERS OF LEAVES USED IN FIG, 24
Top leaves — Thomas 515
A-1. M. Lopez Figueiras 1902
532
7. Léon, Marie-Victorin,
Clément 20753
8. Léon, Marie-Victorin,
Clément 20832
9. Montero 21148
10. Ankerman 11790
13. Howard 6013
14. Shafer 4109
15. Shafer 4054
B-1. Morton, Acufia 3205
2.
Chrysogone 6798
894
6. Marie-Victorin,
Clément, Alain
21610
seen 1s 3884
8 Thom:
9. ieee
Léon 21312
10. Thomas 503
11. Clément, Alain
Chrysogone 3829
12. Thomas 512
COLLECTION NUMBERS OF FRUITS USED IN FIG. 29
2. Clément 2486.
Victorin 20753. i
71. 6. Léon, Mari
Victorin and Clément 20831.
20831. 9. Acufia 13174. 10. Shafer 4181.
B-1. Morton and Acufia 3 2. Clément, Alain and Chrysogone
6798, A894. 3. Shafer 4054. 4. Clément 2486. 5. Léon and Alain
19172. 6. Thomas a ae fers Lopez Figueiras 1902. 8. M. Lopez
C-1. ‘Thomas 503. Svein and Léon 21312. 3. Thomas
521. 4, Shafer 3575. * Thomas 529. 6. Clément 4426.
D-1. Morton and Acufia 3250. 2. Clément 3547. 3. Acufia 13173.
4. Shafer 8083.
KEY TO COLLECTION NUMBERS OF LEAVES USED IN FIG. 32
Collection Altitude (Meters)
A-1. Tate 231 1280
2. Cowan, Wurdack 32131 2000
3. Pinkus 155 2194
4. Card 2000-2200
112 JOAB L. THOMAS
6. Cardona 2042 1800
7. Steyermark 59790 2200
8. Wurdack 34214 2200-2300
B-1. Maguire, Politi 28478 1676
2. Williams 14242 127
3. Steyermark 58353 1820-2075
4, Tamayo 3117
5. Pinkus 77 1280
6. Maguire, Cowan, Wurdack 30178 1800
7. Maguire, Politi 27523 b 1500
8. Steyermark 58140 1700-1980
C-1. Steyermark 60112 2255
2. Cardona 929 1740
3. Maguire, apron Bunting 37037 1700
4. Steyermark 587! 2255-2620
5. Tate 1045 (not ee numbered) 2164
6. Steyermark 58803 2560-2620
7. Tate 401 summit of Roraima
8. Tate 387 A
D-1. Cardona 2696 2100
2. Phelps, Hitchcock 402 1700
3. Maguire, Cowan, Wurdack 30640 1000
4. Maguire, Cowan, Wurdack 30946 1250
5. Schultes, Cabrera 14472 450
6. Gleason 486
7. De La Cruz 3467
8. Spruce 2999
KEY TO THE COLLECTION NUMBERS OF STAMENS USED IN FIG. 27
1. Howard 6013. 2. Léon, Clément, and Howard 20117. 3. Léon,
Clément, Alain, Chrysogone 4693. 4. Léon, Alain 19178. 5. Léon,
Howard, Clément 20117. 6. Clément, Alain, and Chrysogone 4117.
7. Carabia 4087. 8. Morton and Acufia 3250. 9. Marie-Victorin,
Clément, Alain 21610. 10. Léon 11755. 11. Clément, Alain 3872. 12.
Clément, Alain, and Chrysogone 3864. 13. Léon 19171. 14. Clément
3645. 15. Clément 2000. 16. Montero 21135. 17. Clément, Nestor,
24. Alain 3350. 25. Clément, ae and Chrysogone 7407. 26. Léon,
Clément, and Nestor 5441. 27. Clément, Alain, and Chrysogone 3872.
28. Clément, Alain, and Chrysogone 4002. 29. Thomas 510. 30.
Thomas 521. 31. Thomas 520. 32. Thomas 532. 33. Thomas 527.
34, Thomas 528. 35. Thomas 515. 36. Carabia 4087.
KEY TO THE COLLECTION NUMBERS OF PETALS USED IN FIG. 28
A-1. Thomas 515. 2. Clément and Alain 3872. 3. Thomas 503. 4.
Thomas 512. 5. Léon, Clément, — and Chrysogone 3872. 6.
Clément, Alain, and Chrysogone 4002. Alain, and Chry-
sogone 3872. 8. Clément 3645. 9. Wine 510. 10. Marie-Victorin,
MONOGRAPH OF CYRILLACEAE 113
and Alain 21610. 11. Thomas 509. 12. Léon 19171. 13. Thomas
504.
B-1. Thomas 511. 2. Thomas 520. 3. Thomas 515. 4. Carabia
4087. 5. Clément, Nestor, and Chrysogone 7407. 6. Morton and
Acufia 3250. 7. Thomas 504. 8. Clément and Alain 3872. 9. Thomas
518. 10. Alain 3350. 11. Clément 3643. 12. Clément, Alain, and
Chrysogone 3829. 13. Léon and Alain 19171.
C-1. Thomas 521. 2. Clément 2000. 3. Carabia 4087. 4, Clément,
Alain, and Chrysogone 4117. 5. Montero 21135. 6. Thomas 532. 7.
Léon, Howard, and Clément 20117. 8. Léon 11755. 9. Howard 6013.
10. Léon, Clément, Alain, and Chrysogone 4693. 11. Howard 5857.
12. Léon, Clément, and Howard 20117. 13. Léon and Alain 19173.
KEY TO THE COLLECTION NUMBERS OF PETALS USED IN FIG. 34
A-1. Irwin 103. 2. Steyermark 59684. 3. Maguire and Politi 27523.
4. Maguire and Fanshawe 23106. 5. Lasser 1476. 6. Cardona 1133.
9. Pinkus 91. 10. Phelps and Hitchcock 402. 11. Tate 1189. 12.
Maguire 32867. 5. Maguire, Cowan, and Wurdack 30946. 6. Wur-
dack 34214. 7. Maguire 32807. 8. Phelps and Hitchcock 42. 9. Ma-
guire, Wurdack, and Bunting 36936. 10. Maguire, Cowan, and Wur-
41. Pinkus 100. 12. Maguire, Cowan, and Wurdack
615. uf Cardona 929. 8. Pinkus 190. 9. Steyermark 60112. 10. Tate
693. 11. Maguire and Maguire 35122. 12. Wurdack 34099. 13.
Steyermark 58826.
NT USED IN
MORPHOLOGICAL STUDIES
Parts Studied
Species and Geographical 5
iN nm
Collection Area Ps g 8 g 5 £
pa a Se cag 7
AE ees elie = Me = ae = =
Cyrilla racemiflora
x et x
470 N. Carolina xx =
472 N. Carolina x 5 ae 3 =x
483 S. Carolina pee
492 Florida ox
500 Cuba Ce ine eee 3
503 Cuba > ei eae
i =e
114 JOAB L. THOMAS
PLANT MATERIALS USED IN
MORPHOLOGICAL STUDIES
Parts Studied
Species and Geographical
Collection Area ae
Te SB lace es
22226
Harvard Wood Collection
H-19068
H-24138
H-24139
H-24671
H-24691
Purdiaea cubensis
Acufia 16439 Cuba 2x
microphylla
Alain 3387 Cuba = x
moaensis
Thomas 505 Cuba
Lopez 2563 Cuba
nutans
Maguire, Cowan, Wur-
dack 30211 Colombia
parvifolia
Thomas 525 Cuba oe
stereosepala
Thomas 516 Cuba > a ee
velutina
Marie-Victorin, Clém-
ent 21789 Cuba ke
— < x
tonia monophylla
Thomas 379 Florida x x
380 = Florida Pare * x
Thomas 576 Mississipp: Meme x
Godfrey 52959 Florida x
Harvard Wood Col
H-24134
H-24135
H-24136
Cyrillopsis i
Ducke 20629 Brazil x x
Ducke 33826 x
Harvard Wood Collection
mature wood
pollen
Hn K KK
CONTRIBUTIONS FROM THE GRAY HERBARIUM
OF HARVARD UNIVERSITY
Edited by
Reed C. Rollins and Robert C. Foster
NO. CLXXXVII
THE MAIDENHAIR FERNS (ADIANTUM) OF
COSTA RICA
BY
Epira ScaAMMAN
A REVIEW OF THE GENUS DENNSTAEDTIA
IN AMERICA
By
Rotta Tryon
THE LEPTOSPORANGIUM OF THE
NEW ZEALAND FERN
ANARTHROPTERIS DICTYOPTERIS
Br
KENNETH A. WILSON
OBSERVATIONS ON THE LEAVES OF
PELLAEA ANDROMEDIFOLIA
THE GRAY HERBARIUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASS., U.S. A.
1960
CONTRIBUTIONS FROM THE GRAY HERBARIUM
OF HARVARD UNIVERSITY
Edited by
Reed C. Rollins and Robert C. Foster
NO. CLXXXVII
THE MAIDENHAIR FERNS (ADIANTUM) OF
COSTA RICA
By
EpItH SCAMMAN
A REVIEW OF THE GENUS DENNSTAEDTIA
IN AMERICA
By
Rotta TRYON
THE LEPTOSPORANGIUM OF THE
NEW ZEALAND FERN
ANARTHROPTERIS DICTYOPTERIS
By
KENNETH A. WILSON
OBSERVATIONS ON THE LEAVES OF
PELLAEA ANDROMEDIFOLIA
By
ALICE F. TRYON
PUBLISHED BY
THE GRAY HERBARIUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASS., U. S. A.
1960
Issued June 30th
THE MAIDENHAIR FERNS (ADIANTUM) OF
COSTA RICA
EpiItH SCAMMAN
Costa Rica, although a small country, is very rich in ferns,
and contains a greater number of species than any other
country in Central America. The high mountains of the in-
terior and the rushing streams and rivers flowing down the
slopes to both Atlantic and Pacific Oceans afford a great
variety of habitats and climates, ideally suited for an abun-
dance of fern life.
Many species from Mexico and Guatemala in the north ex-
tend into Costa Rica, as well as more tropical species native
to Panama and South America on the southern border. Be-
side the species known to be more or less common and widely
distributed throughout Central America along the coasts at
lower elevations, many rare ferns endemic to the mountain
regions and great central plateau have been discovered and
described by early botanists.
The purpose of my four trips to Costa Rica, usually during
the months of February and March, in 1951, 1953, 1955, and
1956, was to add to the collection of ferns from that country
already in the Gray Herbarium. Accordingly, I spent more
of my time in the interior, visiting regions of historic inter-
est to botanists, such as La Palma, La Hondura, Vara Blanca,
Las Nubes, Valley of the Reventazén, slopes of Trazt and
Barba. Staying in Cartago during my first season, I tried to
go everywhere the crowded buses would take me into the
little villages and then following the cart-roads up into the
hills. The Pan American Highway, recently extended over
the high cerros and so-called “paramos”, offered an oppor-
tunity to collect in regions formerly reached only by horse-
ck.
It was my good fortune during my first trip to meet Dr.
Leslie R. Holdridge of the staff of the Interamerican Institute
of Agricultural Sciences at Turrial The Holdridges kindly
offered me their home in San José as headquarters for my
later visits to Costa Rica, and the days spent in 1955 and
and Sarapiqui rivers proved to be most fruitful. Weekend
pienics and all-day collecting trips tee Sos omit gi in
their jeep, wherever I desired to go, enal me to visit many
vine toestuat to thusiast. My collections could
4 EDITH SCAMMAN
As comparatively little has been written recently about the
ferns in any single Central American country, it has seemed
worthwhile to present the results of a study of a single genus
of wide distribution as a guide to future collectors and fern
lovers who may visit that beautiful land
Costa Rica is divided into seven epee and I have ar-
range the specimens cited under t hem — Limén on the
Atlan and Pt the Pacific, and
Heredia, Alajuela, San 5 osé and Cartan kt in we interior of the
The habitat and altitude given for each species refer only
to the specimens seen from Costa Rica. Only in cases where
no specimen has been seen from there have the habitat and
altitude been noted from another country mentioned.
For inspiration, encouragement, and valuable assistance in
the rag gg sain of this paper I am greatly indebted to Dr.
Rolla M. Tryon of Harvard University, and for the help
afforded by his pe Meant on the Ferns of Peru. He has also
generously provided me with the use of many of the draw-
ings already prepared by Mrs. Bernadette Velick for his
Peruvian work. Other excellent drawings were made espec-
ially for this paper by Mr. John Gumppenberg.
ADIANTUM
The Adiantums are terrestrial ferns of shady ravines and
rocky river banks in moist forests. Although the genus is
found in other parts of the world, Tropical America is the
home of the majority of species of Maidenhair. Of the
twenty-eight species enumerated here from Costa Rica,
many grow along the banks of streams and rivers at low
elevations, while a few are found only in cloud forests in the
mountains at high altitudes.
mes are thick and short-creeping or slender and long-
creeping ; stipes and rachises are firm, dark, usually polished,
erect or less often pendulous ; blades are 1- to 6-pinnate; pin-
nules vary greatly in size and shape, are usually glabrous,
sessile or stalked, often articulate at the base of the stalk and
sometimes deciduous. The genus is especially characterized
by the separate or continuous marginal sori, with the spor-
angia borne = a under side of the modified and reflexed
marginal ind
KEY TO SPECIES
a. Ultimate segments (pinnae — pinnules) borne on long slender
stalks, blade once or bipinna‘ «b.
b. Blade once pinnate, eg ane ts taking root at the apex, pinnae
broadly lunate te fan-shaped, gree a several oblong
Soci bared ttielmenaeea ects
ADIANTUMS OF COSTA RICA 5
c. Sterile margins entire or with obscure, rounded “teeth”, veins
often ending in the indefinite sinus, segments not articulate
ec. Sterile ara sharply serrate, each vein ending in a tooth,
men’ iculate .. 23. A. deflectens.
b. Blade once re Sees not rooting, ultimate segments ovate-
acuminate, glaucous beneath, several oblong to linear sori on
eacl 3. A. St
a. Ultimate segments sessile to short-stalked, or if rather long-stalked
then the blade tri- to quadripinnate . .. .
d. Ultimate segments definitely to rather lonieatathed: the dark
color of the stalk stopping abruptly at the segment base, usual-
ly articulate by a sharp clean break .... e.
e. Rachis and pinna-rachises glabrous, atropurpureous to ebeneous
£: Uitimate segments glaucous eect small (about 0.5-1.5
long), coriaceous, cuneat rhizome long-creeping
ak
f. Ultimate segments green beneath; or if in no. 27 sometimes
—— then trapeziform and large (about 3-5 em. long)
g. ‘Sori tone on the convex outer edge of the fertile segments
h. Seema porn flabellate, the sterile ones with mar-
gins with indefinite teeth, sori oblong-| -lunate to linear,
rhizome scales blackish or with narrow Letgpy’ margins
. Braunii.
h. Segments mostly trapeziform — to ee uae sori
roundish to short-oblong ... - i.
ij. Rhizome scales small, distinctly bicolorous ( center
blackish, margins brown) and long-ciliate;
of sterile segments serrula
. Rhizome scales large, qonbtiorotiss preht brow ait
only slightly ciliate; margins of sterile segments sub-
entire or only faintly denticulate .....- 26. A. princeps.
. Sori borne on the upper and outer edges of the segments;
apex of fertile segments sterile and sie e to acuminate
Wee ser 27. A. trapeziforme.
. Rachis and pinna-rachises densely panernied brown, blade
scandent, its rachis and pinna-rachises strongly flexuous; seg-
ents suborbicul; te, puberulent beneath .... 28. A. Feei.
d. Ultimate segments sessile to very short-stalked, or definitely
stalked egies color of the stalk passing into the segment-
aa
id
i Paper glee S reduced to the apical ultimate segment - he Tee
. Sterile margins of segments sharply serrate, each vein end-
ing in a tooth; rhizome slender, long-creeping, its scales
brown 7. A. Capillus-Veneris.
k. Sterile margins of with each vein ending in a
sinus between the ge Oi or indefinite teeth ... .1-
EDITH SCAMMAN
1. Pinnae subsessile, the lower as Well as the upper with the
inner upper pinnules overlying the rachis, inner upper
pinnule of lower pinnae divided into two ultimate seg-
ments 19. A. concinnum.
1, Pinnae, or at least the basal or lower ones, definitely stalked,
their inner upper pinnules not or hardly overlying the
rachis .
m. Rhizome: sak very ebb ois stipes closely clus-
tered, ultimate segments cuneate to Sgr ge late,
sori orbicular to pc ese i ... 20. A. Raddianum.
m. Rhizome slender, rather anger dig stipes at inter-
vals, ultimate segments mostly orbicular to suborbicu-
sibs sori oblong to lunate, occasionally with yellow
21. A. Poiretii.
j. Blade nO a more = less — terminal pinna, or lacking a
truly apical segment’
n. Blade pedate or tite peat the ultimate branches once
UDI ALES TY OE UE ch ie Fe a 16. A. patens.
n. Blade pinnate ....o0.
0. Blades commonly 4- to 5-pinnate at the rare leaves very
large, segments deeply incised-lobed .... 15. A. pectinatum.
o. Blade once pinnate or opie a
a Blade once pinnate .
- One long sorus iets an edge of the pinna, occasion-
ally this interrupted ....r.
r. Veins somewhat anastomosing toward the edge; pinnae
cordate, stalked, the fertile lanceolate-ovate ... :
1. A. Wilsonii.
¥ ee all free; pinnae sessile or with a very short
Lsomepioe era opposite, nw lowermost with a
road truncate subcordate base, the fertile long-
2. A
. Several short, oblong-arcuate or javb-oblange Sort “along
each margin of the pinna.... 8
or portions evenly serrate, rhizomes often long-
creeping, with spreading scales, the stipe bases usual-
ly well. 5. A. petiolatum.
t. Stipe and rachises clothed with numerous wide-
spreading subulate-filiform scales, pinnules glaucous
beneath; indusium with a broad membranous border
t. Stipe and rachiss gabrous, puberuent to shoré-caly;
indusium usually with a narrow membranous bord
Se eis
ADIANTUMS OF COSTA RICA 7
u. Terminal pinna larger than the ion laterals and
usually broadest at the base... .
yv. Segments green, sterile ones ey biserrate
4, A. obliquum.
y. Segments glaucous .... Vv’.
v’. Sterile segments evenly Serrate ....-.--s-+cesseeeee
5. A, petiolatum.
v’. Sterile segments biserrate .... 6. A. caryotideum.
u. Terminal pinna about the same size as the laterals
and usually narrowed at the base... . W.
w. Pinnules nearest the long acuminate terminal
segment of the pinna greatly reduced, less than
x. Sori several, roundish to oblong, on the upper
and outer edges of the fertile pinnules . .. . y.
y. Pinna-rachises puberulent, rhizome moder-
ately stout, op Berio 2 pinnules long-
triangular or trap
calbreyeri.
y. Rachis and pinna-rachises Say or Sates’!
sealy....Z.
z. Rhizome long-creeping, the stipe-bases dis-
pinna and acute ....... .. 11. A. tetraphyllum.
z. Rhizome coun vere sane often knot-
ted, the stipe bases adjacent, fertile pin-
nules with the sterile — mos’ ight
A, fructuosum.
the
OUtET CAGE -n-r-nesseeseeeeer ... 14, A, pulverulentum.
w. Pinnules nearest the terminal segment usually
not greatly as about half as long as the
aa. Rhizome cord-like, ey long-creeping, sterile
pinnules finely and evenly serra’ serrate .... bb.
bb. Pinnules herbaceous and glaucous, midvein
margins
distinct, i
flat, oblong sori usually on the lower as well
as the upper edge of Samer hese
oe ‘latifotinm.
8 EDITH SCAMMAN
1. Adiantum Wilsonii Hook. Sp. Fil. 2: 6, pl.72A. 1851
This species is one of the few Adiantums, and the only one in Costa
Rica, with veins somewhat anastomosing, only uniting toward the
edges. The fronds are always simply pinnate with a large terminal
pinna and from 2-6 subsessile lateral ones on each side, these ovate-
acuminate, and somewhat rounded at the base. The midrib is median
and distinct, and the sori linear and continuous on both margins nearly
to the apex.
Mexico to Panama, to Colombia; West Indies.
In wet forests at low elevations from near sea level to 200 m.
Specimens seen: LIMON: La Colombiana Farm of United Fruit Co.,
Standley 36785, 36864 (Us); Ontario Farm, Reventazén, Lankester
950 (US). HEREDIA: Plains of Rio Hondo, Cook & Doyle 575 (us).
ALAJUELA: Llanuras de San Carlos, Brade 473 (NY).
2. Adiantum macrophyllum Sw. Nov. Gen. Sp. Prod. 135. 1788
A beautiful fern, distinctive and widely distributed. The blade is
once pinnate, with large, equal-sided, opposite pinnae, the lower-most
ones with a broad truncate to subcordate base. The young fronds are
reddish in color and are striking when seen in the forest and on
of streams.
Mexico to Panama, to Bolivia and Brazil; West Indies.
Generally distributed, usually on banks of rivers in wet forests at
100 to 1400 m. on both the Atlantic and Pacific slopes. Among the
many collections seen, all of the provinces of Costa Rica are repre-
sented.
Representative specimens: LIMON: Tsaki, Talamanca, Tonduz 9458
(GH, NY, US) ; Foréts de Shirores, Talamanca, Pittier & Tonduz 9201
(Us) ; Jiménez, Llanos de Santa Clara, J. D. Smith 5076 (GH, NY, US);
La Lola, cacao finca, near Rio Madre de Dios, Seamman 7066 (GH).
HEREDIA: Confluence of Rio Puerto Viejo and Sarapiqui, Pittier 7500
(us) ; Finea La Selva, Rio Puerto Viejo, Scamman & Holdridge 7450
(GH). : San Ramén, Tonduz 17598 (us); Alajuela, J. D.
Smith 6869 (US). SAN JOSE: Vicinity of El General, Skutch 3898 (GH,
US). CARTAGO: Peralta, Lankester 591 (GH, Us); El Munieco, south of
Navarro, Standley 32479 (us); Dulce Nombre, Standley 35901 (US);
Rio Reventazén, Turrialba, Seamman 6103, 7632 (GH); Rio de las
Vueltas, Tucurrique, Tonduz 12809 (US). GUANACASTE: near Hacienda
Tilaran, Standley
TARENAS: Rio Naranjo, Mazon 657 (us).
: 3. Adiantum Seemannii Hook. Sp. Fil. 2:5, pl. 81 A. 1851
This attractive fern was discovered by Seemann, on one of his voy-
ADIANTUMS OF COSTA RICA 9
a pinna, X 1. Fig. 5. A. petiolatum: a pinna, X 1. Fig.
pinna, X %. Fig. 7. A. villosum: A, base of a pinna, X 1; B, apex of a pinna, X
%-
10 EDITH SCAMMAN
In humid forests and deep ravines at low altitudes from 150 to 700
m. on both the Atlantic and Pacific slopes
Specimens seen: 1901-1905 Wercklé (us). LIMON: Talamanca, Ton-
duz 8580 (us); Los Diamantes, Rubber Plant Station, Scamman 6099
(GH). HEREDIA: La Emilia, Llanuras de Santa Clara, J. D. Smith
6866 (GH, US). CARTAGO: Rio Reventazén below Turrialba, Skutch
4673 (GH, Us) ; Turrialba, Maxon 149 (US), Scamman 6098, 7073, 7633,
7939 (GH). GUANACASTE: Naranjos Agrios, Standley & Valerio 46537
(GH, US); Arenal, May 21, 1927, Valerio (us); Matambé, Nicoya
Peninsula, Cook & Doyle 692 (US). PUNTARENAS: Vallée du Diquis,
Pittier 11952, 11968 (Us)
4, Adiantum obliquum Willd. Sp. Pl. 5: 429. 1810
This species is rather similar to the following one, A. petiolatum,
and often confused with it, especially as both may occur in two forms
—once pinnate and bipinnate. Throughout the range the once pinnate
form is more common; it has the blades broadly linear to elongate-
triangular with few pairs of alternate pinnae, these ovate-lanceolate
and usually more acute at the tip than those of A, petiolatum. In
both species the terminal pinna of the bipinnate blades is larger than
the few laterals and broadest at the base. The segments of A. obli-
quum are green on both sides, not glaucous.
Mexico to Panama, to Bolivia and Brazil; West Indies.
In forest ravines and on the banks of streams from sea level to 600
m. This species seems to be rare in Costa Rica and the few collections
which have been seen are mostly bipinnate.
: HEREDIA: Finca La Selva, Scamman & Holdridge
7937 (GH). CARTAGO: Turrialba, G. P. DeWolf 218 (GH). GUANACASTE:
Tilaran, Standley & Valerio 45012 (GH); Carrillo, Pittier 1167 (US).
PUNTARENAS: Jestis Maria, Lankester 615 (Us).
5. Adiantum petiolatum Desv. Berl. sci 5: 326. 1811
‘Adesnbues Kaulfussii Kze. Linnaea 21: 221.
The pinnae and pinnules of this species are ase and glaucous be-
neath and are usually oblong in shape rather than ovate-acute; the
ones margins are evenly serrate. It is much more common than A.
m.
Mexico to Panama, to Bolivia and Brazil; West Indies.
In wet forests and on rocky river banks, usually from 50 to 300 m.
Representative specimens: LIMON: Tsaki, Talamanca, Tonduz 9485
(us); Hamburg Finca, Rio Reventazén below Cairo, Standley & Va-
lerio 48687 (us); Siquirres, Wercklé 595 (us); La Lola near Rio
de Dios, Seamman 7068, 7070 (GH); Los Diamantes, Rubber
Plant Station, Scamman 7071 (GH). HEREDIA: Rio Toro Amarillo,
Llanuras de Santa Clara, J. D. Smith 6867 (GH, US) ; Rio Hondo, Plains
of Santa Clara, Cook & Doyle 504 (US); Confluence of Rio Puerto
Viejo and Sarapiqui, Pittier 7499 (us); Finca La Selva, Rio Puerto
Viejo, Seamman & Holdridge 7451, 7452, 7933 (GH). ALAJUELA: San
GUANACASTE: Nicoya, peed gg 677 (US); Carrillo, Wercklé ara
(GH, Us); Rio Liberia, Dodge & Thomas 7868 (GH). PUNT.
ADIANTUMS OF COSTA RICA 11
Buenos Aires, Pittier 10557 (Us); Between El General and Buenos
Aires, Holdridge 7069 (GH); Vallée du Diquis, Pittier 11966 (US);
Plaines de Surubres de Puntarenas, Cote du Pacific, Pittier 2685 (US).
6. Adiantum caryotideum Christ, Bull. Soc. Bot. Genéve II, 1: 230. 1909
Type from Costa Rica, 1904, Wercklé.
Rhizome long-creeping, nodose; stipe and rachises glabrous, black
ebeneous; blade bipinnate with the terminal pinna than th
few laterals and broadest at the base. The pinnules are on short but
definite stalks and glaucous beneath, with long lunate sori, adjacent
but separate, on the upper and outer edges. The sterile margins are
biserrate.
The length of the petiolules helps to distinguish this species from
bipinnate fronds of A. obliquum and A. petiolatum.
Mexico to Panama; along rivers from sea level to 300 m.
No specimen seen from Costa Rica.
7. Adiantum yillosum L. Syst. Nat. ed. 10, 2: 1328. 1759
In this species, as in the two following, the pinnules nearest the
terminal segment of the pinna are not greatly reduced. The pinnules
are dark green, lustrous and oblong to narrowly rhombic with the
oblong or linear sori usually extending around the acuminate or acute
tip. The margins of the sterile segments are coarsely serrate. The
rhizome is strongly nodose and rather short-creeping.
ico to Panama, to Peru and Brazil; West Indies.
Specimens seen: LIMON: Along Banana River above R. R. Bridge,
W. W. & H. E. Rowlee 465 (Us). ALAJUELA: Cataratas de San Ramén,
Brenes 13562 (NY). GUANACASTE: Vicinity of Tilaran, Standley &
Valerio 45012 (US); Nicoya, Cook & Doyle 658 (us), Matamba, Nicoya
Peninsula, Cook & Doyle 713 (US). PUNTARENAS: Caldera, Jan. 1926,
Lankester (us).
8. Adiantum latifolium Lam. Encycl. 1: 43. 1783
The rhizome is slender and very long-creeping, with the stipes well
ed. The pinnules are herbaceous, glaucous beneath, the fertile ones
oblong-lanceolate, acute or acuminate with the sori usually on the low-
er as well as the upper edge; the sterile segments are often rounde
at the apex, evenly serrate or denticulate. In this species the midvein
1100 m.
Representative specimens: Cocos Island (Pacific) : R. E. Snodgrass
& E. Heller 967 (GH, us); June 11, 1929, A. K. Fisher (GH, us);
Valerio 2224 (us) ; A. Stewart 226 (us). Limon: La Colombiana Farm
of United Fruit Co., Standley 36704 (us); Port Limén Oct. 20, 1911,
A. S. Hitchcock (us); Zent, Tonduz 14561 (us) ; Siquirres, Scamman
6102 (cH); La Lola, near Rio Madre de Dios, Scamman. 7065 (GH).
HEREDIA: Confluent of Puerto Viejo and Sarapiqui, Pittier 7285 (us),
Finca La Selva, Rio Puerto Viejo, Scamman & Holdridge 7449 (GH).
ALAJUELA: Canton de San Carlos, Austin Smith 1482 (NY); La Palma
de San Ramén, Brenes 5790 (NY); Boca ¥ Vallée de San Juan
12 EDITH SCAMMAN
Nicaragua, Pittier 9632 (US). CARTAGO: Turrialba near Rio Reventa-
z6n, Scamman. 6100, 7064 (GH).
9. Adiantum serratodentatum Willd. Sp. Pl. 5: 445. 1810
This species can be recognized by the many small oblong to rhombic
pinnules, usually rounded at the apex, rigid to coriaceous, with the
few sori borne on revolute margins on the upper and outer edges, but
not on the lower. The terminal pinna is little reduced at the base. The
sterile margins are usually finely serrate.
Guatemala to Panama, to Brazil; West Indi
In open grass lands and thickets in low a middle poo Only
one specimen has been seen from Costa Rica, but it is not uncommon
in Panama. PUNTARENAS: Buenos Aires, Pittier 4850 (US
10. Adiantum Kalbreyeri C. Chr. Ind. Fil. 28. 1905
Adiantum pilosum Bak., Ann. Bot. 5: 207. 1891, not Fée, 1850-52.
This differs from all the other bipinnate species of Costa Rica in
its puberulent pinna-rachises.
Costa Rica to Peru.
This species seems a be rarely collected, usually at middle elevations,
in the interior, in wet forests about 900 to 1500 m.
Specimens seen: CARTAGO: Pejivalle, Standley & Valerio 46879 (GH,
us) ; El Mufieco, Rio Navarro, Standley & Torres 51070 (us) ; Navar-
rito, Lankester 735 (us).
1. Adiantum tetraphyllum Willd. Sp. Pl. 5: 441. 1810
It seems to be difficult to separate this species from A. fructuosum
without the rhizomes which many herbarium specimens lack. How-
ever typical A. tetraphyllum has the sterile apex of the pinnules acute
and almost falcate.
Generally distributed throughout tropical America.
In wooded ravines and dense forests from 50 to 1500 m.
a,
ber Plant Station, Scamman 6105 (GH); near Rio Madre de age Pit-
tier 10256 (US). HEREDIA: Finca La Selva, Rio Puerto Viejo, Seam-
man & Holdridge 7453, 7940 (GH); Vara Blanca, Skutch 3633 (US).
ALAJUELA: San Ramén, Tonduz 17599 (Us), Rio Grande, San Ramén,
Brenes 18875 (GH); Zapote on the road to Villa Quesada, Seamman
7637 (GH). CaRTAGO: Tucurrique, Tonduz 12779 (Us); Tuis, Tonduz
11313 (Us); Juan Vifias, Reventazén Valley, Cook & Doyle 211 (Us);
Tq ba, G. P. DeWolf 22 (GH), Scamman 7074, 7636 (GH). GUANA-
CAsTE: Boca Culebra, on the Pacific, Pittier 12019 (us); Los Ayotes,
near Tilaran, Standley & Valerio 45368 (US). PUNTARENAS: Vallée du
Diquis, Pittier 12052 (us).
12. Adiantum fructuosum Spreng. Syst. Veget. 4: 113. 1827
Apart from the difference i in the rhizomes mentioned in the key the
Mexico to Panama, to Peru and Brazil; West Indies.
In moist forests from 100 to 1100 m.
ADIANTUMS OF COSTA RICA 13
pase of a pinna, X 1. Fig. 9. A.
Kalbreyeri: base of a pinna, X 1.
ion of a pinna, X 1. Fig. 12. A. fructuosum:
scales on is, X 4.
PLATE 2. Fig. 8-13. Fig. 8. A. latifoliwm:
serratodentatum: base of a pinna, X 1. Fig. 10. A.
Fig. 11, A. tetraphyllum: apical
a pinna, X %4. Fig. 13. A. decoratum: A, a pinna, xX 1; B,
14 EDITH SCAMMAN
Representative specimens: LIMON: Port Limén, Cook & Doyle 415,
416 (US). HEREDIA: La Concepcién, Llanuras de Santa Clara, J. D.
Smith 6864 (Us). ALAJUELA: Rio Surubres near San Mateo, Feb. 1906,
Biolley (Gu, Us); Zapote on the road to Villa Quesada, ‘Seutencie 7635
ona, Jimenez 390 (US). PUNTARENAS: Between Golfo Dulce and Rio
Terraba, Skutch 5256 (us).
13, Adiantum decoratum Maxon " i cde Am. Jour. Bot. 19: 165.
932
“Stipe conspicuously shaggy S00 very numerous, wide-spreading,
subulate-filiform scales, these extending freely to the rachises through-
out.” So wrote the authors of this species, and this characteristic is
sufficient to distinguish it from all other related ones. The rhizome is
short-creeping and coarsely nodose. The segments are oblique, rhombic-
oblong to lanceolate, membrano-herbaceous, dark green above, glaucous
beneath with oblong adjacent sori on the upper and outer margins.
= sterile pinnules are unequally serrulate at the upper margin and
ernie to Panama.
In moist ravines in forests from 50 to 800 m. on the Pacific slope
(except the specimen from Turrialba).
pecimens seen: ALAJUELA: Capulin, Rio Grande de Tarcoles, Stand-
ley 40174 (US); Gorge of Machuca River, near San Mateo, Biolley 2017
(us), Rio Machuca, Biolley 17392 (GH, US). SAN JOSE: Hills above
Rio Paquita, Dodge & Goerger 9876 (US); Cerro Turrubares, Orotina,
Jimenez 602 (Us); Cangrejal de Aserri oe April, 1906, Biolley
(Us). CaRTAGO: Turrialba, Scamman 6101 (GH). GUANACASTE: Matam-
ba, Niecoya Peninsula, Cook & Doyle 693 an. Nicaves Cook & Doyle
655 (US), Tonduz 13766 (GH, US); Garza, Holdridge 7631 (GH).
14. Adiantum pulverulentum L. Sp. Pl. 2: 1096. 1753
It is a relief in this group of Adiantums to find a species as easily
recognizable as this one. It is characterized by the single long sorus
borne on the upper margin of the pinnules. Rarely a shorter sorus
occurs on the outer edge. The rachis and eset eared are pubescent-
scaly and the pinnules slightly stellate-scaly beneath.
Mexico to Panama, to Brazil; West Indies.
In shaded ravines in forests from 600 to 1300 m.
Specimens seen: 1901-1905, Wercklé — SAN JOSE: San Marcos,
Pittier 7722 (us); Cangrejal de Aserri (Pacific slope) April, 1906,
Biolley (us). carTaco: Juan Vifias, Reventazén Valley, Cook & Doyle
203 (us); Livingston on Reventazon, W. W. Rowlee & H. E. Stork 610
(us) ; Turrialba, Seamman 6097, 7077, 7634 (GH).
15, Adiantum pectinatum Kze. ex Ettingsh. Farnkr. 85, t.45, fig. 14-16
: 1865
This is the largest of the Adiantums, the fronds sometimes becoming
2 m. tall. The ade i broadly deltai-vate to deti-crelar, 4. to
5-pinnate, rarely 6-pinnate. The segments are numerous and small,
deeply incised-lobed with the roundish sori borne at the apex of the
ADIANTUMS OF COSTA RICA 15
184
18B
PLATE 3. Fig. 14-18. Fig. 14. A. pulverulentum: base of a pinna, X 1. Fig.
A. pectinatum: base of a pinna, x 1. Fig. 16. A. patens: pase of a pinna, X 1.
Fig. 17. A. Capillus-Veneris: A, a small pinna, X 1; B, a sterile segment, x 2. Fig.
18. A. Braunii: A, a sterile pinna, X %4; B, apex of a fertile pinna, X *%-
16 EDITH SCAMMAN
lobes on the upper and sometimes outer edge. They have conspicuous
veins and are glabrous or have a few stellate scales beneath.
Costa Rica, where it is rare, to Argentina and Brazil.
In wet forests from low to middle elevations, 80 to 500 m. in Costa
Rica.
Specimens seen: ALAJUELA: Capulin on the Rio Grande de Tarcoles,
Standley 40171 (Gu, US). SAN JOSE: Llano Grande de Puriscal, Jim-
enez 835 (US); San Luis de Turrubares, Valerio 1729 (us); Rio Tur-
rubares, Brade 437 (Ny).
16. Adiantum patens Willd. Sp. Pl. 5: 489. 1810
The frond is pedately tripartite and resembles, in the manner of
division of the blade, A. pedatwm — the beautiful and beloved Maiden-
hair of North America. It is smaller and has fewer branches; the
stipe and rachises are reddish brown and glossy, slightly puberulent.
The lunate or reniform-orbicular sori are borne in rounded sinuses
on the upper and outer edges of the segments.
Mexico to Costa Rica; Venezuela to Colombia, to Bolivia.
Rocky slopes in mountains from 1000 to 1900 m. in the region of
Costa Rica known as the Meseta Central.
Specimens seen: Nov. 1886, J. J. Cooper (GH, Us); 1906, Wercklé
(Us). ALAJUELA: Alajuela, 1887, A. Alfaro (Us). SAN JOSE: San José,
Valerio 205 (us), Brade 154 (NY); San José to Escast, Pittier 48
(us); La Urica, Pittier 913 (Us); Rio Torres, Tonduz 8066 (us); La
Verbena, Alajuelita, Tonduz 8798 (us); Tablazo, Biolley 107 (us);
Santa Maria de Dota, Standley 42375a (us); Pozo Azul de Pirris,
Lankester 1170 (us). CARTAGO: Dulee Nombre, Quiros Calvo 1158
(yy).
nous border, each vein ending in a tooth; a striking characteristic
which distinguishes this well-known and widely distributed species
from many of the Adian‘
Temperate and tropical America, also in parts of Europe, Asia and
Africa.
Mexico and Guatemala from sea level to 1500 m. No specimen seen
from Costa Riea, but as it may be expected to occur there, it is included
in the key. :
ii Prominent nerves, the upper and outer borders deeply and bluntly
lobed, the sterile margins with indefinite teeth. The fertile pinnules
On moist shady banks in the mountains, 1100 m., rare in Costa Rica.
imens seen: SAN JOSE: San José, Brade 16673 (us) ; La Verbe:
prés Alajuelita, Tonduz 8802 (us).
ADIANTUMS OF COSTA RICA 17
19. Adiantum concinnum Willd. Sp. Pl. 5: 451. 1810
The blade is lance-oblong in shape, usually drooping, bi- to tri-
pinnate, with subsessile pinnae, both the lower and upper with the
inner upper pinnules overlying the rachis; sori roundish to reniform.
inner news pinnule of the lower pinnae into two ultimate segments.
Mexico to Panama, to Venezuela and Peru; West Indies.
On moist cliffs and shaded roadside banks, common in some localities,
from 400 to 1400 m,
Representative specimens: Nov. 1886, J. J. Cooper (GH, US).
(GH). SAN JOSE: San José, Rio Tiribi, Biolley 103 (GH, US) ; San José,
Sept. 1887, EZ. S. Hyde (GH, NY, US); El General, Skutch 2913, 3949
(GH, US); Aserri, Scamman 6094 (GH); Finca , Desamparados,
Scamman & Holdridge 7935 (GH); Santiago, A. Alfaro 8068 (GH, NY,
us) AGO: Orosi, Scamman 7063 (GH). @ Delicias
};UAN ACASTE:
below Hacienda Santamaria, Dodge & Thomas 6288, 6291 (GH).
20, Adiantum Raddianum Presl, Tent. Pterid. 158. 1836
Adiantum cuneatum Langsd. & Fisch. Ic. Fil. 25. t. 26. 1810, not
Forst. 1786.
Adiantum Werckleanum Christ, Bull. Herb. Boiss. II, 4: 1093. 1904.
Hab. Costa Rica, Wercklé.
This is a complex species with slight variations, some fronds with
chief differences between this species and the next one— A. Poiretii
—are shown in the key. The blade is usually tripinnate, rarely bi-
pinnate or quadripinnate, the stipes dark reddish brown to blackish,
with the pinnae definitely stalked and the ultimate segments cun cuneate
to broadly cuneate-flabellate.
A specimen of Wercklé’s collection determined by Christ seen in
New York and other specimens so labelled in the U.S. National Her-
barium do not appear to differ fundamentally from forms of A, Rad-
dianum, so they have been included under this species.
Tropical America.
In niggas country on banks of streams, in oak forests from 1700 to
is seen? 1906, Wercklé (US). ALAJUELA: Zarcero, July 13,
i Tiatehep Radoailo:
above Escast,
Standley 32518 (GH, US) ; Between Aserri and Tarbaca, Standley 41396
(us); Vicinity of Santa Maria de Dota, Standley 43119 (us); Near
Quebradillas north of Sits Maria de Dota, Standley 42904 (GH, US);
Tablazo, Brade 147 (NY, US).
re Adiantum Poiretii Wikstr. Vet. Akad. Handl. 1825: 443. 1826
. thalictroides Willd. ex Schlechtd. Adumbr. Pl. pt. 5: 53. 1832.
cele, capt the stipes vary
from light to dark reddish brown to atropurpureous; the ultimate seg-
18 EDITH SCAMMAN
segment, 22.
X 4. Fig. 23. A. deflectens: A, a leaf, X %; B, a sterile pinna, X 114.
ADIANTUMS OF COSTA RICA 19
ments are suborbicular to subcuneate, lobed, and the sori oblong to
lunate.
avico to Costa Rica, to Chile and Argentina; Hispaniola.
i i moist shady banks, rocky hillsides, and slopes
of voleanos at high elevations, from 1500 to 3000 m. in Mexico and
Guatemala. The single cole from Costa Rica does not indicate
the habitat or altitude.
SS en seen: SAN JOSE: 5 miles northeast of Coronado, M. A.
Chrysler & W. E. Roever 5339 (us).
hone Adiantum philippense L. Sp. Pl. 2: 1094. 1753
A ng fern easily recognized by its once pinnate blade oc-
ee rooting at the tip with the oblong-lunate, dimidiate pinnae
on fairly long petiolules and with the uppermost pinna cuneate-
flabellate. The sori are oblong-linear, close together and sometimes
contiguous. The segments are not articulate.
Mexico to Panama, also Cuba, and northern South America. Widely
distributed in Africa, Asia, and Australia.
Among rocks on banks of creeks and streams and near waterfalls,
from 100 to 650 m.
In Costa Rica it has been rarely collected and all the specimens
seen are from the Pacific slope. ALAJUELA: Aguacate, C. Hoffmann
754 (NY, US); Canyon, Rio Grande de TArcoles, Oratina, W. W. & H.
E. Rowlee 200 (us); Machuca River near San Mateo, Lankester 611
(US). GUANACASTE: — of Tilaran, Standley & Valerio 46570
(US). PUNTARENAS: Rio Surubres, Brade 318 (NY), Plaines de Suru-
bres, Puntare: ria ain 2666 (us); Rio Jesis Maria, route de Pun-
tarenas, Pitted 490 (us).
23. Adiantum deflectens Mart. Ic. Crypt. Brasil. 94. 1834
This delicate little fern, somewhat related to A. philippense, is very
distinct from that species since the sterile margins are sharply serrate,
each vein ending in a tooth. The margins of the fertile pinnae have
are articulate with a clean break at the junction with the stalk.
Guatemala, Honduras, Panama; Guianas to Colombia, to Peru and
Brazil.
No has been seen from Costa Rica, but it is included here
as it doubtless may be found there.
24, Adiantum andicola Liebm. Dansk. Vid. Selsk. Skrift. V, 1: 266
(seors 114). 1849
Adiantum glaucophyllum Hook. Sp. Fil. 2: 40. 1851.
Adiantum Cooperi Baker, Journ. Bot. 25:25. 1887.
Blade deltoid, 3- to 5-pinnate, ultimate segments flabellate-cuneate,
rather small, coriaceous, green above, glaucous beneath, veins promi-
nent; sori at apex of lobes, roundish-reniform.
Adiantum Cooperi was described by Baker from a collection from
Costa Rica by J. J. Cooper. He refers to it as “allied to A. glauco-
phyllum Hook., from which it differs by its much larger, fewer, less
crowded segments, not at all glaucescent beneath.” The size size of the
20 EDITH SCAMMAN
Garden Herbarium does not seem distinct enough to warrant calling
it a separate species.
Mexico to Pahanin, to Colombia.
Along shady banks and in oak forests on slopes of volcanos, at
high altitudes from 1700 to 3000 m.
Specimens seen: 1901-1905, Wercklé (us). ALAJUELA: La Ventolera,
southern slope of Volcan Pods, Standley 34699 (GH, US). SAN JOSE:
Cerro de Piedra Blanca above Escast, Standley 32490 (us); Vicinity
of Santa Maria de Dota, Standley 41664 (GH, US); Finca Guayabillos,
July, 1936, Dodge & Goerger (GH); Laguna de la Escuadra, northeast
of El Copey, Standley 42013 (us); Las Nubes, Scamman & Holdridge
7941 (GH). CARTAGO: Slopes of Volcan Irazi, near Hotel Robert,
Williams & Molina 13862 (GH), Scamman 6104 (GH); near Sanatorio
Duran, Scamman 7062 (GH); Cervantes, Lankester 665 (Us); Reven-
tado, Lankester 693 (US); Rio Birris, southern slope of Volean Irazt,
Standley 35418 (us).
25. Adiantum tenerum Sw. Nov. Gen. Sp. Prod. 135. 1788
Adiantum trapezoides Fée, Gen. Fil. 117. 1852. (See Morton, Am.
Fern Jour. 45: 113-115, 1955, for discussion of this and the following
species).
Rhizome short-creeping with many bicolorous, ciliated scales. Stipes
and rachises dark purplish brown, polished, blades deltoid-ovate, 3- to
5-pinnate, with segments quite variable, rhombic-oblong to flabellate-
cuneate, sori retuse-oblong and numerous. The ultimate segments are
jointed at the base of the stalks and definitely articulate by a sharp
ean break, easily falling when dry.
Florida, Mexico to Panama to Venezuela; West Indies.
Cool ledges, shaded banks, old walls, rarely collected in Costa Rica.
In Mexico from 300 to 2000 m.
imens seen: Costa Rica, Nov. 1886, J. J. Cooper (GH, US); 1901-
1905, Wercklé (us).
26. Adiantum princeps Moore, Gard. Chron. n. ser. 4: 197, fig. 43, 44.
1875
Adiantum trapezoides of recent authors, not Fée.
A large fern 1-1.3 m. high, the stipes almost black, very shiny, seg-
ments larger than those of Adiantum tenerum, generally —s in
shape, but variable, articulated at the junction with the The
margins of the sterile segments that are subentire or ony ae
denticulate, and the concolorous brown scales distinguish this species
from A. tenerum.
Mexico (common in western Mexico) to Panama; Colombia.
—— shady river banks from 100 to 1200 m.
seen: HEREDIA: Banks of Rio Virilla, P. H. Allen 572
ig ALAJUELA: Gorges of Machuca, near San Mateo, Biolley 2021
(us), Rio Machuca, Biolley 17387 (us), Lankester 610 (US). SAN
JOSE: El Rodeo, Hunnewell 16484 (GH). GUANACASTE: Vicinity of Ti-
laran, Standley & Valerio 45695 (Us); La Colonia Carmona, Jimenez
389 (US). PUNTARENAS: Puntarenas, Jan. 1926, Lankester (us).
21
ADIANTUMS OF COSTA RICA
Ji
&
oN
A.
26. A. princeps: a pinna, X %. Fig. 27. A. trapezi-
a fertile segment,
24. A. andicola: an upper pinna, X 1. Fig. 25. A.
Fig. 28. A. Feei: A, a pinna, X 1; B,
Fig. 24-28. Fig-
tenerum: a pinna, X %. Fig-
forme: base of a pinna, X %.
x 3.
22 EDITH SCAMMAN
27. Adiantum trapeziforme L. Sp. Pl. 2: 1097. 1753
Blade quadripinnate with large glabrous pinnules, bright green
above, sometimes glaucous beneath; stipe naked, polished, blackish;
the ultimate segments long-stalked, mostly trapeziform, acute to
acuminate at tip; sori numerous, contiguous, on the upper and outer
e.
A striking and beautiful Adiantum with the shining almost black
stalks extending to the base of the green segments, but not articulate.
Mexico to Panama to Brazil ; West Indies.
In forested ravines, on the banks of creeks and rivers from 80-1000
m. on the Pacific slope.
‘im. - UELA: Gorges of Rio Machuca, near San
Mateo, Biolley 2018 (us), Rio Machuca, Biolley 17393 (us), Surubras
Grande de TArcoles, Standley 40130 (GH, US); Rio Grande, A. Alfaro
(GH). GUANACASTE: Garza, Holdridge 7638 (GH); Nicoya, Cook &
Doyle 676, 680 (us).
28, Adiantum Feei Moore ex Fée, Mém. Foug. 7: 29. pl. 24, fig. 1. 1857
Adiantum fleruoswm Hook. Second Cent. Ferns pl. 61. 1861. (See
Morton, Am. Fern Jour. 45: 117, 1955).
This fern is distinct from all other Adiantums by its scandent
blade and zigzag rachis and pinna-rachises spreading at a right angle.
The densely tomentose brown stalks stop abruptly at the base of the
small rigid, puberulent, suborbicular-cuneate segments.
Mexico to Panama, on mountain slopes, 1350 to 2400 m. Very rare.
The only specimen seen or reported from Costa Rica is labelled:
“Costa Rica and Veragua”, (Panama) collected by Warscewicz, no. 32
(yy, Us).
Other species of Adiantum described from Costa Rica are the follow-
ig. As no specimens have been seen and the types have not been
available, these names cannot be placed; they are undoubtedly syno-
nyms of some of the species treat
A. heteroclitum Christ, Bull. Herb. Boiss. II, 4: 1094. 1904. Costa
Rica, 1903, Wercklé.
A. orosiense Christ, Repert. Sp. Nov. 8: 17. 1910. Costa Rica,
Orosi, finca de D. Valverde, 1900 m., Brade 16816; Carrillo, 400 m.
18 jun. 1909, Brade 314.
A. palmense Christ, Bull. Soc. Bot. Genéve TI, 1: 230. 1909. Costa
A. subtrapezoideum Christ, Bull. Herb. Boiss. TI, 4: 1094. 1904.
Costa Rica, Nicoya, Pittier 13768. — Gray HERBARIUM, HARVARD UNI-
_ NOTE. — On a recent visit in Paris where he saw the type specimens
in Christ’s herbarium, Dr. Tryon has determined A. orosiense to be a
Synonym of A. Kalbreyeri, A. palmense of A. andicola, and A. sub-
trape; A. princeps,
A REVIEW OF THE GENUS DENNSTAEDTIA
IN AMERICA oe
¢
ROLLA TRYON
Some twenty species of Dennstaedtia are currently rec-
ognized in the Americas; however, there has been no recent
survey of them and several have not been adequately defined.
The treatment of the genus by Maxon’ is an excellent one but
includes only four (two of his species I place together) of our
ten tropical species. During preparation of an account of
Dennstaedtia in Peru, it become apparent that to treat the
genus adequately it would be necessary to consider all of the
American species. I have found reasons to maintain only
eleven of the species previously recognized; however, these
conclusions are sometimes tentative, for a number of prob-
ems remain which must be studied in the field and with more
adequate collections.
Most of the species of Dennstaedtia are of common occur-
rence and of these I have cited only a selection from the many
specimens examined. I have not cited any specimens of the
well known and distinctive D. punctilobula. A few species
are rare and in their treatment I have cited all of the mater-
ial I have seen. The specimens in the Gray Herbarium and
the United States National Herbarium have formed the
principal basis for this study. I am indebted to Conrad Vv.
Morton for the loan of material and also to Bassett Maguire
of the New York Botanical Garden for making certain speci-
mens available for my study. Mr. Richard van Frank has
taken the photographs. :
I have considered all of the names that seemed pertinent
to the nomenclature of the species and some others that are
rather widely used in the literature or in the herbarium. I
have not listed combinations made under segregates of
Dennstaedtia, none of which is currently considered serious-
ly: Patania Presl, Sitolobium Desv., Adectum Link and Lit-
clobium Newm.
THE GENUS DENNSTAEDTIA
The definition of Dennstaedtia has, through the years,
been increasingly clarified. Species were often. originally
described in such genera as Polypodiwm, Dicksonia, Deparia
and Microlepia. Most of these have been transferred to
7 MAXON, W. R. Pteridophyta in Sci. Surv. Porto Rico and the Virgin Islands,
G: 491-494, 1926.
24 ROLLA TRYON
Dennstaedtia. The genus is particularly close to Microlepia
and the relation of these two genera will be discussed. Other
related genera, with hairs on the rhizome, are Oenotrichia
and Leptolepia of the Old World; they seem distinct from
Dennstaedtia but I have not especially considered them.
In relation to Microlepia, I have surveyed the species of
it and of Dennstaedtia readily available to me, which in-
cluded 12 American species, 54 Old World species and one
pan-tropic one. On the basis of this study it is evident that
the two genera are indeed closely related and evidently in-
tergrade. However, of the 67 species studied, nearly all
clearly belong in either one genus or the other, and the spec-
ies that would tend to unite the genera are few. These major
groups of species are each sufficiently h and
distinct so that it does not seem a justifiable course, or a
practical one, to unite them. I have assessed the genera and
maintained them on the basis of their soral characters and
a thorough revision of all species of Microlepia and Denn-
staedtia may support this course. Such a study may indi-
cate, however, that other and better characters exist and
should be used either to unite the two genera or to maintain
them.
In Dennstaedtia the sorus is marginal, the indusium being
formed of an inner (true) indusium and an outer indus-
ium which is the opposed leaf tissue, modified in form and
re; these are fully joined (connate) beyond the margin
of the segment to form a reflexed saucer-, purse- or cup-
shaped, or globular or cylindrical whole indusium; or they
are joined as far as the margin and extend separately be-
ties with Microlepia Hookeriana. The generic (soral)
characters presented here separate the two species and I am
inclined to regard their similarities as due to convergent
evolution. Each is a 1-pinnate species in a genus of species
DENNSTAEDTIA IN AMERICA 25
with decompound leaves and the sorus has perhaps been
modified in each case by the alignment of adjacent sori
along an entire margin, so that neither species has a sorus
quite typical for its genus. Saccoloma Wercklei is closely re-
lated, in its pubescence, to several American species (nos. 7-
10) and in its sorus especially to one of these, D. arborescens
(Compare figs. 32 and 38 ). The indusium of M. Hookeriana
shows a relation to the form sometimes present in M. pilo-
sula where the edges of the indusium attached to the leaf
tissue may be attenuated toward the margin of the segment.
In Microlepia the sorus is abaxial, borne well back of, or
near, the margin of the segment, the indusium is fully at-
tached (rarely only partially) to the leaf tissue along its
sides, these ti pp hing and rarely reaching the
margin; it is usually half cup-shaped, with the sides and
apex extended rather equally, or rarely the sides are at-
tenuated well beyond the apex toward or to the margin.
I would place the following critical species in Microlepia:
Dennstaedtia concinna Rosenst. (illegit.), Microlepia denn-
taedtioid: opel., Microlepia melanorhachis Rosenst., and
Dennstaedtia resinifera (Bl.) Mett. In Dennstaedtia con-
cinna the sorus is rather close to the margin and only a
moderate degree of modification of it and of the indusium
would make it, by definition, a taedtia. In Microlep
jamaicensis and a species from New Caledonia (Rosenst.
Fil. Nov. Caled. exsicc. 131) the sides of the indusium are
only partially attached to the leaf tissue. If some of the
species listed above are placed in the genus I suggest, no-
menclatu: changes are necessary. I :
make these, without a greater familiarity with the species,
except in the case of the single American one.
SPECIES IN DENNSTAEDTIA
The species of Dennstaedtia have been difficult to define.
This difficulty, I believe, is due more to the fragmentary
lack of defining characters. Most, if not all, of the remain-
ing problems exist where there is a lack of sufficient inform-
ation. The leaves of Dennstaedtia are usually large and the
specimens prepared from them commonly consist of such a
part as may readily be ac dated an
sheet and they are not often accompanied by notes on the
portion of the lamina represented or its shape. There is
iderabl tainty involved in relating these pieces to
a whole lamina so that one may compare characters of com-
26 ROLLA TRYON
parable portions of the leaf. There are so few collections
that provide data on the shape of the lamina that, although
evidently an important character, it has not been possible
to emphasize it. When a sufficient number of adequate speci-
mens exists it will also be possible to know if specific charac-
ters are to be found in the petiole and rhizome. There may
be differences between juvenile and adult leaves of the same
species and there is a possibility that the juvenile type may
persist in large leaves.
The key presents the species as less well defined than they
actually are. A key that utilized lamina shape and charac-
ters of the basal pinnae more prominently would reflect the
difference between the species to better advantage but would
be impractical to use with most specimens. A major difficul-
ty in the construction of the key, and one that will hardly
fail to pass unnoticed by its users, is that of the characters
available for a practical key some are variable and others, al-
though constant, are not unique. The identification of a
single fragment then often depends on a combination of its
characters and will not always be either easy or perhaps
possible.
GEOGRAPHY AND RELATIONSHIP OF THE SPECIES
The American species of Dennstaedtia have, with the ex-
ception of D. punctilobula and D. Wercklei, large and com-
plex leaves. The lamina is frequently 1-1.5 (-2) m. long and
the petiole is of similar length. The leaves are borne singly
on an extensively creeping rhizome and the species often
form large and conspicuous colonies, They grow predom-
inantly in moist and usually shaded places. The tropical
species are most frequent in forests but also grow along
forest margins, or in clearings and other open habitats.
The most pronounced group within the American repre-
sentatives of the genus is formed by species 7-11 which
share the characters of enlarged sterile vein tips and a close-
ly crispate, persistent pubescence on the under surface of
the axes. Of these five species, Dennstaedtia Sprucei and
D. Wercklei each have one or more unique characters. The
problems of their definition are discussed under their treat-
ments. All three are known to bear proliferous buds in the
axils of the pinnae and are the only American species that
have such buds. D. arb is distributed from Mexico
DENNSTAEDTIA IN AMERICA 27
to Panama, the Greater Antilles, and in the Andes south to
Bolivia. D. dissecta has a similar range but is more widely
distributed in northern South America and extends, in the
south, to southern Brazil. D. obtusifolia is the most widely
distributed of the three; its range is similar to that of D.
dissecta except that it also occurs throughout the Lesser
Antilles and is the only species of the genus in that area.
D taedtia globulif and D. bipinnata are evidently
related, sharing a similar type of pubescence and decurrent
herbaceous wings on the minor axes. Both species are dis-
tributed widely in tropical America, from Mexico to Pan-
ama, the Greater Antilles, northern South America to Boli-
via. The former, D. globulifera, has a wider range in the
south, extending to Argentina, Uruguay, and Braz‘1; while
the latter, D. bipinnata, extends north to Florida.
e three species, Dennstaedtia cicutaria, D. distenta and
D. glauca are not clearly closely related but do share charac-
ters of pubescence and of slender sterile vein tips. D. ci-
cutaria is the most widely distributed of the three, growing
from Mexico to Panama, the Greater Antilles, and in South
America southward to Bolivia and southern Brazil. D. dis-
tenta has a relatively restricted range from Mexico to Pan-
ama; Jamaica and Hispaniola. D. glauca, the only species
of Chile, also occurs from northwestern Argentina to south-
ern Peru.
punctilobula, the only species of temperate
to boreal regions, has no close relations with the other
American species. Rather, it is related to certain species of
eastern and southeastern Asia. It occurs in eastern ani
midwestern United States, north to Newfoundland. The
gland-tipped trichomes on the lamina are distinctive among
the American species. Dae
The occurrence of six of the American species In Central
America and Mexico and also in the Greater Antilles, but
not the Lesser Antilles, lends emphasis to the well known
floristic affinity between those two regions.
28 ROLLA TRYON
KEY TO THE SPECIES
a. Axis of the penultimate segments, as in figs. 8, 9, lacking perpen-
dicular herbaceous wings on the upper surface or perpendicular
b. Sterile vein tips, on the upper surface, slender, as in fig. 17, not
enlarged, ending well back of the glabrous margin; tertiary axes
glabrous to sparsely or densely pubescent beneath with straight to
tortuous, more or less spreading trichomes. c.
¢. Trichomes on the under surface of the pinnules gland-tipped,
or many of them so, fig. 2. 1. D. ilobule
¢c. Trichomes on the under surface of the pinnules not gland-
tipped, or absent.2 d.
d. Pinnae alternate, the lower ones stalked, with the basal pin-
nules not or scarcely reduced, as in fig. 4. e.
e. Many or most of the sori borne in a sinus, fig. 5; pinnules
more or less pubescent beneath, fig. 5; lamina deltoid.
(Trichomes usually abundant on the under surface of the
pinnules, some of them short, acicular, rigid, or rarely these
absent and only larger, sometimes lax and subtortuous ones
present; cells of the trichomes, except sometimes the lower
ones, mostly several times longer than broad). . <=
2. D. cicutaria.
e. All or most of the sori term ‘al on lobes, fig. 13; lamina
ovate- or deltoid-lanceolate. lai
d. Pinnae opposite or rarely subopposi
each with a pair of usually much reduced basal pinnules, fig.
6 (a detached pinna). (Major axes straw colored on the
b. Sterile vein tips, on the upper ‘ace, in fig. 30,
vat punctate, or rarely (in D. obtusifolia) slender; tertiary
axes glabrate to usually subappre: ascending pul it
neath with the trichomes more or less curled. (Pinnae subopposite
to alternate, the lower ones subsessile or less often short stalked).
£:
f. Lamina pinnate-pinnatifid or more complex. (Basal pinnules
of the lower pinnae more or less reduced, often strongly so, as
in fig. 6, a detached pinna). g.
g. Sterile veins ending well back of the glabrous margin, as in
ss
Se SES ae
* Rarely, specimens of D. obtusifolia have slender vein tips and are also
sterile
glabrate: they may be separated from D. cieutaria by the subopposite, nearly sessile
lower with reduced basal pinnules, from D. glauca by the sori borne predomin-
antly in a sincs, from D. distenta by the major axes that are brownish beneath.
DENNSTAEDTIA IN AMERICA 29
h. Pinnules 1-pinnate to pinnate-pinnatifid, figs. 23, 25-28,
apical segments of the pinna separate or nearly so to the
prolonged apex, the separate ones closest to the tip obtuse;
obtuse pinnules on the apical pinnae deeply pinnatifid;
lamina deltoid. i.
i. Mature sori, at least the basal acroscopic ones on an ulti-
segment, mostly 1. 5 mm. broad and about half
as thick, fig. 24. ...... obits ... 7. D. dissecta.
i. Mature sori mostly 0.5-1.0 mm. broad and about as thick,
fig. 29. 8. D. obtusifolia,
h. Pinnules entire to deeply pinnatifid, figs. 31, 33-35, apical
segments of the pinna confluent back of the prolonged apex,
the separate ones closest to the tip acute; obtuse pinnules
on the apical pinnae entire to lobed, lamina evidently ovate.
9. D. arborescens.
g. Sterile veins nearly reaching the persistently pubescent
margin, 37 10. D. S; i.
f. Lamina 1-pinnate, the pinnae entire, fig. 39. (Sterile veins
ending well back of the glabrous margin) ...... 11. D. Wercklet
a. Axis of the penultimate segments bordered on each side, on the
upper surface, by a pronounced herbaceous wing perpendicular to
the plane of the segment, fig. 21, the wing on the basiscopic side de-
current onto the axis of the next order either as an her'
wing, fig. 22, or as a pronounced ridge; trichomes on the under
or many of their cells several times longer than broad; pinnae alter-
nate, rarely subopposite, the lower stalked with the basal pinnules
not or scarcely reduced, as in fig. 4; sterile vein tips, on the upper
surface, slender, fig. 17, to rarely clavate, ending well back of the
glabrous margin). j.
j. Basal segments of the pinnules of the central pinnae usually sub-
opposite to nearly opposite, rather or quite equal in size, the in-
ferior not or slightly ascending, fig. 14, (apical pinnae with the
i imi! the basal segments of the pinnules of the
times shining above, dull beneath, the surface of the upper epider-
mis (dry) minutely pebbled by the individual epidermal cells; sori
globular, fig. 16, to less often subglobular, rarely cylindrical; la-
mina deltoid. 5. D. globulifera.
j. Basal segments of the pinnules of the central pinnae definitely
alternate, quite unequal in size, the inferior ascending to strongly
ascending, fig. 18, (apical pinnae with the basal pinnules similar,
the basal segments of the pinnules of the basal pinnae may be less
unequal and the inferior less ascending) ; leaf tissue usuall; i
dermis (dry) minutely striate to sometimes nearly smooth; sori
cylindrical to subcylindrical, fig. 20, rarely globular; lamina ovate
to lanceolate-ovate. 6. D. bipinnata.
30 ROLLA TRYON
ybe-°°* 1, Dennstaedtia punctilobula (Michx.) Moore, Ind. Fil.
; xevii. 1857. Fics. 1-2.
ol ° Nephrodium punctilobulum Michx. Fl. Bor.- Am. 2: 268.
1803. Holotype: Canada, “Polypodium punctilobulum”, Hb.
Michaux, Pp. (D. C. Eaton, Can. Nat. 13: 28. 1870, comments
on the identity of the type).
The gland-tipped trichomes on the lamina (Fig. 2) dis-
tinguish this species from all other American ones. It is
related, rather, to certain species of eastern and southeast-
ern Asia, especially to D. appendiculata, and D. scabra but
also to D. pilosella and Microlepia stenoloba (this last spec-
ies has not been transferred to Dennstaedtia) .
The named forms, f. cristata, f. nana, f. Poyseri and f.
schizophylla, are fully treated in Gray’s Manual of Botany,
ed. 8.
be
In the eastern portion of its range this species grows in a
variety of habitats, especially in open pastures, in meadows,
in woods, on rocky hillsides, streams banks and moist road
banks ; from Indiana westward it becomes restricted to sand-
stone cliffs.
Newfoundland southwestward through the southern parts
of Ontario, Indiana and Illinois, to lowa (US), Missouri and
Arkansas, and southward to Georgia and Alabama (US).
I do not believe it is necessary to cite specimens of this
well known and distinctive species.
‘wO4° 2 Dennstaedtia cieutaria (Sw.) Moore, Ind, Fil. xevii.
1857. FIGs. 3-5,
ecole? Dicksonia cieutaria Sw. Schrad. Jour. 1800: 91. 1801.
Holotype: Jamaica, Swartz, s-PA, fragment and photographs
us!.
20°49? Dicksonia rubiginosa Kaulf., Enum. Fil, 226, 1824. Holo-
type: Rio de Janeiro, Brazil, Hb. Mertens.
2 Dicksonia apiifolia var. dissecta Desv. Mém. Soc. Linn.
Paris 6: 318. 1827. Holotype: none cited, presumably Bra-
zil, Raddi.
cos “5 Dicksonia angustidens Presl, Tent. Pterid. 136. 1836.
Based, indirectly, on Dicksonia apiifolia var. dissecta Desv.
dee-2to2 Dicksonia umbrosa Liebm. Vid. Selsk. Skr. V, 1: 262 (re-
print 101). 1849. Holotype: Colipa to Misantla, Mexico,
March, Liebmann, c; isotype, us!.
>” 5, Dennstaedtia globulifera (Poir.) ca Bot. Jahrb.
4-16.
34: 455. 1904. FIGs.
*°! Polypodium globuliferum “ste -Encrel 5: 554. 1804.
_ Holotype: Santo Domingo, Plumier, Fil. t.
‘2 Dicksonia altissima Sm. Rees Encycl. 11. 1808. Based on
psa | sespisse globuliferum Poir. 2
cksonia punctulata Poir. ‘cong afar de Suppl. 2: 475.
1811. Holotype: Hb. Desfonta:
2b6-
° Dicksonia Pawan (Poir.) O. Ktze. Rev.
2 Dieksonia tenera Presl, Del. sma 1: 189. 1822. Holotype:
Rio de Janeiro, Brazil.
icksonia exaltata Kze. Bot. Zeit. 8: 59. 1850. Holotype:
Plumier, Fil. t.
30.
. Dennstaedtia tenera (Presl) Mett. Ann. Sci. Nat. V,:2:
"261. 1864.
014-1534 Dieksonia Lagerheimii Sod. Crypt. Vasc. Quit. 50. 1893.
Holotype: Voleain a Corazén, 2000-2800 m., Ecuador,
eclegtiad isotype or authentic specimen, ny! US eg e !
37 8.
1898.
(253 Dennstaedtia exaltata (Kze.) Hieron. Bot. Jahrb. 34: 454,
1904.
4,535 Dennstaedtia Lagerheimii (Sod.) C. Chr. Ind. Fil. 217.
1905.
38 ROLLA TRYON
~°Dennstaedtia Tamandarei Rosenst. Hedwigia 56: 359.
"4915. Syntypes: Brazil, Legru in 1913; Brazil, Tamandaré
5421. Rosenst. Fil. Bras. —— marked “n. sp. original”,
“Tamand” & Brade 5421, NY
* Dennstaedtia Bradeorum elas: Repert. Sp. Nov. 22:
3. ees vee vl = Rica, Brade 634; isotype, NY!, US!.
lif is closely related to the next spe-
cies, D. bipinnata, i in the characters mentioned in the key.
Of these, perhaps the most important is the character of the
perpendicular herbaceous wings that border the axes of the
penultimate segments and are decurrent onto the next axis.
The differences between the two are discussed under D. bi-
pinnata.
The distinctive characters of the basal segments of the
pinnules (nearly opposite, rather equal is size and the in-
ferior not or slightly ascending) may be developed only in
certain portions of the lamina. They are present in the cen-
tral portion of basal pinnae and progessively more basal
portions of the pinnae above. In the apical pinnae, where
the pinnules are not large enough to have basal segments,
the same characters are developed in the basal pinnules.
The apical portions of the pinnae and the basal portions of
the lower pinnae may have the basal segments of the pin-
nules similar to those in D. bipinnai
Wet forests, moist shaded hillsides, forest borders, stream
banks, canyons, and ravines, 200-2300 m.
Eastern and central Mexico to Panama; Greater Antilles;
Venezuela and Colombia south to Bolivia, Argentina, Uru-
guay and southern Brazil.
Representative specimens: Mexico. TAMAULIPAS: Sharp 5077 (US).
SAN LUIS POTOSI: Pringle 3824 (GH, US). VERACRUZ: Bourgeau 1941,
2342 (cu, US); Conzatti & Gonzales 613 (GH, US); Copeland 72 (GH,
4 x Noy. 27, 1937, nonsatd (GH): Kenoyer & Crum 4108 (GH).
78350, 81027 (US); Tiirckheim IT 1328 (us). Hecdicaa: Williams €
t
Haupt 42 (GH, us); Scamman & Holdridge 7396 (GH) ; Standley 43336
(vs). Panama: Killip 5252, 5487 (us); Maxon 4969, 5725 (GH, US).
Cuba: Clément 1185 (us); Ekman 14359 (us); Howard 5198 (GH,
Us) ; Jack 7037 (us), 7272, 7987 (GH, US). Jamaica: Webster & Gold-
berg 20 (US), 22 (GH, US), 92 (us). Hispaniola. Harti: Ekman H3117,
H5598 (us); Holdridge 1981 (us); Leonard 8768, 4026 (GH, US).
DOMINICAN REPUBLIC: Ekman H11634, H11778 (us) ; Miller 1222 (us);
Tiirckheim 3075 (GH, Us). Porto Rico: Hioram 256 (us). Venezuela:
DENNSTAEDTIA IN AMERICA 39
Dennsta
TE 4.
4, Colombia, H. H.
40 ROLLA TRYON
Linden 140 (GH). Colombia: Holton 70 (GH); Killip & Smith 19111
GH, US); H. H. Smith 1118 (GH, US); Triana 48 (us). Ecuador:
Camp E-3462 (GH, US); GALAPAGOS ISLANDS: Stewart 882, 883, 959
(GH, US). Peru: Killip & Smith 24648 (us), 25473 (GH, Us). Bolivia:
Buchtien 3382 (GH, in part) ; Cdrdenas 2853 (us); Herzog 2232 (us);
Steinbach 8939 (GH, US). Argentina. sUsUY: Lillo 3123 (GH). SALTA:
Rodriguez 1033 (GH). TUCUMAN: Lillo 2875 (GH) ; Venturi 255, 2516,
6136 (GH, US). CHACO: Schulz 714 (GH). MISIONES: Gozalbo 38 (GH) ;
Hindobro 5435 (GH); Rodriguez 664 (GH). Paraguay: Hassler 6842
(GH), 12244 (US); Morong 571 (us). Uruguay: Osten 6541 (us).
Brazil. RI0 GRANDE DO SUL: Leite 154 (GH). SANTA CATHARINA: Dusén
L. B. Smith 2246 (GH, us); L. B. Smith & Brade 2278 (GH, US).
MINAS GERAES: Regnell II 322c (us).
~22"°5 6. Dennstaedtia bipinnata (Cay.) Maxon, Proc. Biol.
oe Soc. Wash. 61: 39. 1938. Fics. 17-22.
~ Dicksonia bipinnata Cav. Deser. Pl. 174. 1802. Holotype:
Porto Rico, Ventenat, MA; isotype, Hb. Willd. B, fragment
us!. (Maxon, loc. cit., discusses the Hb. Willdenow specimen
accepted as an isotype).
26-2/°56 Polypodium bacciferum Poir. Lam. Eneyel. 5: 554. 1804.
Holotype: Port-de-Paix, Santo Domingo, Plumier, Fil. t. 31
and the identical Plumier, Amer. t. 45.
Dicksonit iantoides Humb. & Bonpl. ex Willd. Sp. Pl.
5: 488. 1810. Holotype: Caripe, Venezuela, Humboldt &
Bonpland, Hb. Willd., B, fragment us!.
246-2/5°* Dicksonia globuligera Desv. Mém. Soc. Linn. Paris 6: 317.
1827. Based on Polypodium bacciferum Poir.
266-0406? PD taedtia adiantoides (Humb. & Bonpl. ex Willd.)
Moore, Ind. Fil. xevii. 1857.
aedtia bipinnata is most closely related to the pre-
2g -oFe73
cur in the axils of the tertiary segments and of the pinnules.
The South ican specimens (Fig. 18) are usually of
a firmer texture and more deeply laciniate than those from
DENNSTAEDTIA IN AMERICA 41
42 ROLLA TRYON
Central America and the West Indies (Fig. 19). This var-
iant is not sufficiently distinctive. however, to deserve recog-
nition.
Wet forests or forest borders, road banks, rocky slopes
and open habitats, sea level to 2000 m.
uthern Florida; central Mexico to Panama; Greater
Antilles; Trinidad to Colombia, south to Peru and Bolivia.
Representative specimens: United States. FLORIDA: Palm Beach Co.,
May 14, 1926, Small et al. (GH, US). Mexico. HIDALGO: Kenoyer 710
(GH, US); H. E. Moore 3402 (GH). CHIAPAS: Ghiesbreght 356 (GH);
Miinch 155 (us). Guatemala: Heyde & Lux 4667 (GH, Us). Honduras:
Standley 18621, 55545 (Us) ; Thieme 5639b (GH, US). Salvador: Stand-
ley 19760, 20207 (GH, US). Costa Rica: Pittier 6925, 7494, 10347 (US);
5892, 7018, 7019, 7608 (GH); Scamman & Holdridge 7895
(GH) ; Skutch 3031 (GH, us); Standley & Valerio 45186, 45895 (US).
Panama: Killip 4538 (US), 5003 (GH, US) ; von Wedel 718 (Us). Cuba:
Britton & Shafer 2096 (us); Clément 1419, 5224 (us); Eggers 4938
(us); Hioram 6299, 6404 (GH); Linden 1747 (GH); Maxon 3960,
4144, 4400 (GH, US); Morton & Acufa 3613 (GH, US); Pollard & Pal-
mer 174 (us); Shafer 7868, 8499 (GH, US); Wright 895 (GH), 962
(GH, US). Jamaica: Maxon 10330 (US), 10368, 10371 10385, (GH, US);
Oreutt 5776 (GH). Hispaniola. Harti: Ekman H8199 (Gu, us) ; Leon-
ard 7919 (GH, US); Leonard & Leonard 12149, 14324 (GH, US). DOM-
INICAN REPUBLIC: Abbott 1592 (GH, US) ; Ekman H11466 (us) ; Fuer-
tes 1554 (GH, US); Valeur 739 (GH, US). Porto Rico: Britton 5214
(GH, US); Abbe & Marble Oe (us) ; Britton et al. 6189 (US); Eg-
‘rinidad: Protas 6020, 7265 ase Pendior 157 (cH). Neraca!
H. Smith 2431 (eH, us). Ecuador: Haught 3368 (GH, US). Peru
Killip & Smith 22651 (GH, pene 22875, 26799, 27796 (US); Mexia
6127 (GH, US); Spruce 4890 (NY). Bolivia: Buchtien 313, 3378 (US);
Herzog 321 (us); Krukoff pce (GH, US); Steinbach 3036 (GH); R.
S. Williams 1259, 1267 (GH, US).
os -7, Dennstaedtia —= oe ei Ind. Fil. 305.
Mee oth 64 oo 2 vag 0g 134. 1788, not Forst.
2¢¢-0+2te Dieksonia dissecta Sw. Schrad. Jour. 18002: 1801.
Pitota pe: Jamaica, Swartz, s-Pa, photograph and hare
2b- hb kein cornuta Kaulf. Enum. Fil. 227. 1824. Holotype:
Brazil, “ex Spreng.
500-5635? Dicksonia Millefolium Desv. Mém. Soc. Linn. Paris 6:
_,318. 1827. Holotype: Hb. Desv., P, photograph GH!.
266- otbse oo cornuta (Kaulf.) Mett. Ann. Sci Nat. V, 2:
DENNSTAEDTIA IN AMERICA 43
1
ig. 25, upper pin
ral portion of
PLATE 6. - 23, central portion of a pina,
_ onard Fig.
Fi cent
co, Broadway in 1911; Fig. 27, etre portion of a pinna,
nna,
Peidias: xitep 12141.
44 ROLLA TRYON
e shape and size of the mature sori distinguish D. dis-
secta (Fig. 24) from D. obtusifolia (Fig. 29). In the key
have expressed this difference in terms of the breadth and
the thickness. There is a simular relation of breadth and
length, although this is not as constant. However, it may be
useful for identification of immature specimens. In D. dis-
secta the sorus is about twice as broad as long, while in D.
obtusifolia it is about as broad as long. The basal acroscopic
sori of a segment are especially distinctive in D. dissecta; the
apical ones may resemble those of D. obtusifolia. I have not
been able to discover any means of identifying sterile speci-
mens of these species, but fortunately they are few.
The close relation of D. dissecta and of D. obusifolia, their
similar ranges and the occurrence of at least a few mixed
collections* suggest that perhaps they are variants of a
single species. However, D. dissecta is not known from
Cuba, Porto Rico or the Lesser Antilles, while D. obtusifolia
eccurs in all of those areas.
Scamman 7609 and 7610 from Costa Rica have prolifer-
ous buds in the axils of the pinnae.
Wet forests, shady woods and forest openings, 30-2600 m.
Southern Mexico to Panama; Jamaica and Hispaniola;
Trinidad to Colombia, south to Bolivia and southern Brazil.
ote atine specimens: Mexico: VERACRUZ: Copeland 71 (GH, US).
CHIAPAS, ht 368 (GH). la: Heyde & Lux 3212 (GH,
US); ‘Minch 151 (US); Tiirckheim 8629 (GH, US), II 2367 (US).
British Honduras: Schipp S-921 hae Nicaragua: Standley 10919
(Us). Costa Rica: Maxon 305 (us); Maxon & Harvey 8053 (US);
Pittier 63 (GH); Scamman. 7021, 7609, 7610 (GH); Standley 37173,
pound 51258 (us). Panama: Cornman 1195 (Us); Killip 5175 (GH).
jamaica:
alti: Ekman, H7527, H7528, H10185 (us); Leonard 4279,
4740 ste ae Trinidad: Lockhart (GH). Venezuela: Fendler 59 (GH);
Moritz 283 (GH, US). Colombia: Killip & Smith 16057 (GH, US) ; Lin-
den 1043 (US); Pennell 4424, 7602 (GH, US); Stiibel 438 (Us). Ecua-
dor: Eggers 15313 (US); Rimbach 105 (GH, US). Peru: Killip & Smith
24546 (US); Maebride 4844 (Us) ; Mexia 8248 (GH, US). Bolivia: Bang
2423 (GH, Us) ; Herzog 2260 (US); Rusby 127 (US). Brazil: RI0 GRANDE
DO SUL: Leite 3048 (US). SANTA CATHARINA: Luederwaldt 1821 (US).
PARANA: Dusén 14116 (US). SAO PAULO: Brade 8351 (US). RIO DE
JANEIRO: Glaziou 1232 (us).
%E-02722 8. ee (Willd.) Moore, Ind. Fil.
1861. FIGs. 26-
266-0 $270 Dicksonia cituijots Willd. Sp. Pl. 5: 483. 1810. Holo-
* Maxon 9375 and 9901, Mazon & Killip piss Jamaica, and Leite 3048, Brazil,
are all D. dissecta at us and D. obtusif<
DENNSTAEDTIA IN AMERICA 45
type: Ceara: Venezuela, Bredemeyer, Hb. Willd., B, photo-
graph US
Beececer eel ordinata Kaulf. Enum. Fil. 226. 1824. Holo-
type: Porto Rico, Ventenat. (Kze. Farnkr. t. 106b is evi-
dently drawn from the holotype).
2-222” Dennstaedtia ordinata (Kaulf.) Moore, Ind. Fil. 306.
1861.
soo © °°" Dennstaedtia producta Mett. Ann. Sci. Nat. V, 2: 260.
1864. Syntypes: Colombia, Lindig 329, B, photographs and
fragment _ P, photograph GH!; Colombia, Lindig 333, B,
GH!, NY!, US
$00 - Dicksonia incisa Fée, Mém. Fam. Foug. 11:94, t. 25, fig 1.
1866. Holotype: Guadeloupe, L’Herminier in 1864; authen-
tic tins L’Herminier 172 in 1862, P, photograph GH!.
266-2? Dicksonia cicutarioides Fée, Mém. Fam. Foug. 11: 95, t.
25, fig. 2. 1866. Holotype: Guadeloupe, L’Herminier in 1864.
Dennstaedtia incisa (Fée) Kuhn, Linnaea 36: 146. 1869.
« Dicksonia scandens Baker, Jour. Bot. 15: 162. 1877, not
Bl. 1828; illustrated in Ie. Pl. t. 1605. Holotype: Andes of
Quito, Ecuador, Sodiro, K, photograph and fragment us!
“ Dicksonia pubescens Baker, Jour. Bot. 19: 203. 1881, not
Schkuhr, 1809. Holotype: Antioquia, Colombia, Kalbreyer
1859, K, photograph and fragment us!.
»--056¢> Dennstaedtia Sodiroi Diels, Nat. Pflanz. 1‘: 218. 1899.
Eeeed on Dicksonia scandens Baker.
Goo" 365%, var. protrusa Christ, Prim. Flor.
Costar. 3: 38. 1901. Holotype: Costa Rica, Tonduz 117835, P,
isotype, GH!, us!.
Zé /_Dewstacdtia cicutarioides (Fée) Hieron. Bot. Jahrb. 34:
a54. 1
pope cicutaria var. deparioides s Rosenst. Hedwigia
432214. 1904. Holotype: Toledo, Sao Paulo, Brazil, Ulbricht
73, Pa!
vo" 5" Dennstaedtia pubescens (Baker) C. Chr. Ind. Fil. 218.
1905.
bec-% 2+ Dennstaedtia deparioides (Rosenst.) Rosenst. Hedwigia
46: 71. 1906.
dee 26 Dennstaedtia Kalbreyeri Maxon, Proc. Biol. Soc. Wash.
51: 3 40. 1938. Based on Dicksonia pubescens Baker.
lia is most closely related to D. dis-
secta and their distinguishing characters and problems of
separation are discussed under the latter treatment. i
This species and D. ordinata have commonly been main-
characters by which to separate Three other species
have been less often recognized as distinct from D. obtusi-
46 ROLLA TRYON
folia. Dennstaedtia producta Mett. is an unusually lacini-
ate form. Dennstaedtia incisa (Fée) Kuhn is a form with
especially small and narrow lobes. Dennstaedtia Kalbreyeri
Maxon I can interpret only as an unusual variation of D.
obtusifolia with a fine, stiff and abundant pubescence, and
the pinnules nearly at right angles to the pinna-rachis and
with very uniform tertiary segments. The type of D. Kal-
breyeri has some trichomes on the under surface of the
costa that resemble (although they are longer) those usually
occurring in D. obtusifolia. Another specimen (Cuatrecasas
9382, us) has the Nes of D. Kalbreyeri and the typical
of D.
er
Fée (loc. cit., t. 25. ge: 2) illustrates a specimen of this
species with proliferous buds in the axils of the pinnae.
Wet forests, forest clearings, wet banks and hillsides, sea
level to 2100 m.
Guatemala to Panama; Greater and Lesser Antilles;
Trinidad to Colombia, south to Peru, Paraguay and south-
ern Brazil.
Representative specimens: Guatemala: Tiirckheim 1053 (GH), 8629
(Us). Honduras: Ames 105 (Us). Nicaragua: C. F. Baker 2468 (US);
Levy 453 (GH). Costa Riea: Scamman & Holdridge 7899, 7900 (GH);
Skutch 5357 (us); Tonduz 11785 (GH, US); COCOS ISLAND: Pittier
16230 (GH) ; Snodgrass & Heller 952 (GH). Panama: Killip 2832 (GH),
12141 (GH, Us). Cuba: Clément 1178 (GH, US), 1497, 1557, 1609 (GH);
Ekman 3937 (us); Pollard & Palmer 165 (GH, US); Wright 895 (GH,
Us). Jamaica: Clute 292 (us); Maxon 8836 (GH, US); Wilson & Mur-
ray 581 (GH). Hispaniola. Harti: Leonard & Leonard 12280 (GH, US).
DOMINICAN REPUBLIC: Abbott 320 (us), 486 (GH, Us); Ekman H11473
(GH, US); Howard & Howard 9394 (GH, US). Porto Rico: Britton &
Cowell 1023, 2205 (us); Scamman 6524 (GH); Sintenis 1789 (US),
5443 (GH, US), 6536 (US). Guadeloupe: Duss 4184, 4341 (Us); L’Her-
minier 172 (GH); Stehlé 1464 (us). Dominica: Lloyd 900 (us). Mar-
+ Duss 1685, 4164 (GH, US), 4686 (US). St. Lucia: Box 469, 470
(us) ; Proctor 17905 (GH). St. Vincent: H. H. Smith 1723 (Gu, Us).
pecoogson Broadway 1862 (GH). Tebago: Broadway 4219 (GH, US),
223 (GH). Trinidad: Britton et al. 1353 (GH, US); Broadway 5354
ons 5914 (US), 9961 (GH,US); Fendler 84 (GH, US). Venezuela:
‘endler 374 (GH); MARGARITA ISLAND: Johnston 185 (GH, Us), Miller
& Johnston 160 (GH, Us). Colombia: poe 333 (GH, Ny, US); Pennell
4472 (US), 8918 (GH, US); Pennell et al. 8665 (GH, Us). Ecuador:
Hitchcock 21764 (GH, Us); Stiibel 947 (us). Peru: Kanehira 160 (us);
Killip & Smith 22647 (eu, us), 23915, 24916 gen Schunke 154 (us).
Paraguay: Hassler 6840, 12244 Sy Brazil. RIO GRANDE DO SUL:
Leite 3048 (GH). SANTA CATHARINA: Spannagel (Ros. exsicc. 241)
(Us). PARANA: Dusén 14678 (GH, us). SAO PAULO: Luederwaldt 21351
(GH). RIO DE JANEIRO: Glaziou 5256 (US).
DENNSTAEDTIA IN AMERICA 47
48 ROLLA TRYON
aeth? 9. D PES 52
i: ‘b (Willd.) Ekman ex Maxon,
Proc. Biol. Soc. Wash. 43: 88. 1930. FIGS. 30-35.
Davallia arborescens Willd. Sp. Pl. 5: 470. 1810. Holotype:
_,Santo Domingo, Plumier Fil. t. 6.
266-2%2"9 Nayallia concinna Pres], Rel. Haenk. 1: 66. 1825, not
Schrad. 1818. Syntypes: “Chile, 1790, Haenke’”, 2 sheets
2257 ig
Dicksonia concinna (Presl) Hook. Sp. Fil. 1: 74. 1844.
2") Deparia Mathewsii Hook. Sp. Fil. 1: 85, t. 30B. 1844.
Holotype: Peru, Mathews 1 782, K, photograph and frag-
ment US!; isotype, us!
2 Dicksonia consanguinea Kl. Linnaea 20: 445. 1847. Holo-
type: Puerto Cabello, Columbia, Karsten.
2% Dicksonia recognita Kze. Bot. Zeit. 8: 57. 1850. Holotype:
Plumier, Fil. t. 6.
*) Patania triangularis Presl, Epim. Bot. (reprint) 261.
1852. Holotype: Colombia, Linden 1042, PR; isotype, GH!.
2s~el2£? Dennstaedtia concinna (Presl) Moore, Ind. Fil. xevii.
1857.
Sco- 1? Sob Dennstaedtia macrophylla (Desv.) Moore, Ind. Fil. xevii,
1857.
2h6- 06 25) Dennstaedtia Pavonii (Hook.) Moore, Ind. Fil. 307. 1861.
26e- oF FSD, taedti i (Kl.) Moore, Ind. Fil. 305.
1861.
See- 26% Dicksonia adiantoides var. coronata Sod. Rec. Crypt. Vasc.
Quit. 23. 1883. Holotype: Ecuador, Sodiro; Corazén, Sodiro,
US!, K, photograph Us! and Canzacoto, Sodiro, Ny! are pre-
sumably authentic.
266-22 322 a coronata (Sod.) Sod. Crypt. Vase. Quit. 48.
1893.
Seo~ ne Dennstaedtia coronata (Sod.) C. Chr. Ind. Fil. 216. 1905.
**3 Dennstaedtia Mathewsii (Hook.) C. Chr. Ind. Fil. 218.
dee-08?
1905.
246-3777" Dennstaedtia grossa Christ, Bull. Herb. Boiss. II, 6: 192.
1906. Holotype: Navarro, Costa Rica, Wercklé 9, P; authen-
tie specimens collected by Wercklé, ny!. us!.
Maxon (loc. cit.) has discussed the application of Willde-
new’s name and the confusion that led to the inappropriate
epithet. The Plumier plate is not an entirely satisfactory
DENNSTAEDTIA IN AMERICA 49
match for material recently collected in Hispaniola, or else-
where, but unless it can be shown that it represents a differ-
ent species it seems best to accept its identity as Maxon de-
termined it.
Some forms of D. arborescens with deeply pinnatifid pin-
nules and some of the previous species, D. obtusifolia, with
1-pinnate pinnules are rather similar and fragmentary
specimens may be difficult to identify.
In the shape and size of the ultimate segments, D. arbore-
scens is the most variable of the American species. The three
principal variations are illustrated in Figs. 31, 33 and 34.
The form with broadly obtuse segments illustrated in Fig.
33 may occur in both fertile and sterile pinnae, or the sterile
ones may be truncate, as in Fig. 35. It is with some uncer-
tainty that I treat these all as one species and do not give
them some intraspecific status.
My conclusion that D. arborescens should be considered a
polymorphic species is based on the following considerations.
Dennstaedtia Sprucei, a distinctive species, exhibits some of
the variability in shape and size of the ultimate segments
that D. arborescens does. Some specimens, Fig. 36, are simi-
lar to those of D. arborescens in Fig. 33; others are substan-
tially the same as those of D. arborescens in Fig. 31. None
of the forms has a distribution that would favor its recogni-
tion. There are no other characters (or at least I have found
none) to support those of size and shape of the ultimate seg-
ments. Finally, complete intergradation exists between the
forms, although these int diate speci are not as
numerous as the others. The lack of a distinctive geographic
range and the parallel variation in D. Sprucei indicate, ci
believe, that no geographic subspecies or ti involv-
ed in the variation of D. arborescens. However, adequate
field studies may bring out facts of correlation with habitat
or local geography that may result in a reevaluation of this
conclusion.
leave no observable scar; thus it is not possible to determine
if specimens formerly bore them.
Wet forests, ravine banks, forest borders; 100-1500 m. in
Central America, 1200-2200 m. in the Greater Antilles and
400-2800 m. in South America. :
Southern Mexico to Panama; Cuba, Jamaica and Hispan-
iola; Venezuela to Colombia, south to Bolivia.
ore
2¢e
~ 06296
50 ROLLA TRYON
Representative specimens: Mexico. CHIAPAS: Miinch 60 (us).
Guatemala: Hatch & Wilson 114 (Us); Salvin (GH). Costa Rica:
Mazon & Harvey 8006 (GH, Us); Scamman 7020, 7897, 7898 (GH);
Standley 44096 (GH, US). Panama: Cornman 1226 (GH, US); Maxon
4968 (GH, US) ; Pittier 5338 (GH, US). Cuba: Ekman 5499, 7145, 7146
(us). Jamaica: Chrysler 2055 (GH, US). Hispaniola. HaITI: Ekman
H1682, H3122, H5446 (us), H7633 (GH). Venezuela: Fendler 60
(GH, Us) ; Moritz 386 (US). Colombia: Killip & Hazen 8987 (GH, US);
Killip & Smith 19970, 20414, 20462 (Gu, us) ; Linden 1042 (GH);
H. ro 1072, 2218 (GH, Us). Eeuador: Mexia 7208 (GH, US) ; Rene
bach 66 (US). Peru: Macbride 4176, 4842 (F, US) ; Mathews 1782 (us);
Killip & Smith 24642 (¥, GH, us) ; Spruce 4346 (GH). Bolivia: Buchtien
314, 3593 (GH, US) ; R. S. Williams 1243 (GH, US).
10. Dennstaedtia Sprucei Moore, Ind. Fil. 308. 1861.
Holotype: Ecuador, Spruce 5350; isotype, GH!, Cc,
photograph and fragment vs!. FIGS. 36-37.
~~ Dicksonia Sprucei (Moore) Baker, Jour. Bot. 15: 162.
1877.
This is one of the most distinctive species; the sterile
veins that nearly reach the margin of the segment and the
persistently pubescent margin (Fig. 37) are unique charac-
ters among the American species. Specimens have some-
times been identified as D. Pavonii, but that name is based
on Davallia arborescens ( =Dennstacdtia arborescens)
rather than the specimen of D. Sprucei (Peru, Ruiz &
a cited and illustrated by Hooker (Sper tl: 14 t
6A).
Wet forests and moist open places, 1500-2000 m.
Ecuador and Peru
= Kenadoc: Mille 165 (US) ; 1873, Sodiro (Ny) ; Spruce
5350 (GH). Peru: Killip & Smith 25848 (GH, US); Soukup 1826 (GH,
us).
11. Dennstaedtia hpi, A aaa Tryon, comb. nov.
Wercklei ae Bull. Heh Boiss, II, 4: 1100.
Saccoloma
1904. Holotype: Costa Rica, Wercklé 320, P, photograph and
figures in Am. Fern. Jour. 48: pl. 13 and 14. 1958.
‘<-082% Dennstaedtia arcuata Maxon, Am. Fern Jour. 35: zB
yy
+ ot ~
£199
1945. Holotype: sb esigean Killip 5565, US!; isotype:
C a, Dryander 2446, GH!, us!, dispéeak
6570, us!; Peru, Tan 674, US!).
rena | Wercklei (Christ) Kramer, Am. Fern Jour.
The 1-pinnate lamina with entire, slightly auriculate pin-
nae is sufficient to distinguish D. Wercklei from all other
DENNSTAEDTIA IN AMERICA 51
De: sae arborescens: Fig. 33
7 ng ERED
Dennstaedtia
Fig. portion of a sterile pinnul
lip & Smith 25848. De —- Wercklei:
; Fig. entral pinnae, 14, Colombia, Deyander 2 2456.
52 ROLLA TRYON
face of the costa and the enlarged sterile vein tips relate this
species ve Dz dissecta, D. obtusifolia, D. arborescens and
D. Spruce
Fy sei iMloe: cit.) has discussed and illustrated the pre-
viously obscure Saccoloma Wercklei and has (in herb.) iden-
tified D. arcwata as conspecific with it.
Forests, about 2000 m.
Costa Rica, Colombia and Peru.
Specimens seen: Colombia: Dryander 2446 (GH, ch Juzepezuk
6570 (Us); Killip 5565 (Gu, US). Peru: Bryan 674 (US
DUBIOUS AND EXCLUDED NAMES
Some of the following names are undoubtedly based on specimens
of Dennstaedtia but, not having seen the types, I have not been able
to place them in synonymy. From some portion of the protologue, it
is reasonably certain that none of them could be the correct name for
any of the species recognized in this treatment. Other names are
included that have been placed in Dennstaedtia but are referable to
another genus.
Dennstaedtia apiifolia (Sw.) Moore, Ind, Fil. xevii. 1857. Dicksonia
apiifolia Sw. Schrad. Jour. 18002: 91. 1801. =MAXONIA APIIFOLIA.
itaedtia divaricata (Sod.) C. Chr. Ind. Fil. 217. 1905. Dicksonia
divaricata Sod. Crypt. Vase. Quit. 48. 1893. Holotype: Volean Pulula-
hua, 2000 m., Ecuador, Sodiro,
aedtia erosa (Kze.) Moore, Ind. Fil. 306. 1861. Dicksonia
scribed as scaly and this would exclude it from Dennstaedtia; how-
ores the epg Linas may be erroneous for the name has been applied
pe coicram fluminensis (Fée) C. aS Ind. Fil. 217. 1905. Microle-
pia fluminensis Fée, Crypt. Vasc. Brésil 1: a, t. 51, fig. 1. 1869. Holo-
type: Brazil, Glaziou 2378. = MICROLEPIA
Dennstaedtia grandifrons Christ, iy “Flor. Costar. 3: 38. 1901.
Holotype: Costa, Rica, Tonduz 11931,
iformis (Fée) C. Chr. Ind. Fil. 217. 1905. Micro-
lepia estan Fée, Crypt. Vase. Brésil 1: 152, t. 51, fig. 2. 1869.
Syntypes: Brazil, Glaziou 2379, 3332. =MICROLEPIA SPELUNCAE?
Dennstaedtia i Christ, Bull. = Boiss. II, 5: 732. 1905.
Holotype: Chiapas, Mexico, Miinch 137,
Orbignyana Kuhn, Linnae ht 146. 1869; Chaetopt.
348. 1882. Holotype: Bolivia, DOrb
(Baker) C. chr sity Fil 218. 1905. Dicksonia
Pearcei Baker, Ann. Bot. 5: 197. 1891. =LoxsoMOPSIS PEARCEI.
aedti: Christ, Bull. Herb. Boiss. II, 5: 258, 732.
1905. =HYPOLEPTS sp.
Dennstaedtia v. vagans (Baker) Diels, Nat. Pflanz. 14: 218. 1899.
Dicksonia vagans Baker, Jour. Bot. 15: 162. 1877. Holotype: Andes of
bacco Ecuador, Sodiro, k.— GRAY HERBARIUM, HARVARD
THE LEPTOSPORANGIUM OF THE NEW ZEALAND
FERN ANARTHROPTERIS DICTYOPTERIS
KENNETH A. WILSON
Anarthropteris was described as a genus by Copeland
(1947) and includes only A. Dictyopteris (Mett.) Copel. a
fairly common epiphytic or epipetric New Zealand fern
which has also been reported from the New Hebrides (Dob-
bie, 1951). The genus was considered by Copeland to be most
closely related to Loxogramme and the two genera were
grouped by him with the grammitid ferns. A survey of the
mature sporangia of polypodioid and grammitid genera dis-
closed that the sporangial structure of Anarthropteris did
not conform with that of either the polypodioid or the gram-
mitid ferns, and it was, in fact, considerably different from
those of Loxogramme (Wilson 1959). Of considerable mor-
phological interest, however, was the structure of the spor-
angial stalk which was very irregular in its cellular
arrangement and suggested that longitudinal intercalary
divisions had taken place during the ontogeny of the stalk.
Since no vertical divisions had been observed in the sporang-
ial stalk cells of the polypodioid genera Phlebodium (Wilson,
1958a), and Pyrrosia, or the grammitid genus Xiphopteris
(Wilson, 1958b), it was felt that a study of the ontogeny of
hydroxide (5%) and stained in tannic acid and iron chloride
following the methods described in earlier papers on spor-
angial ontogeny. An herbarium specimen of the plant mater-
ial was prepared and has been deposited in the Gray
Herbarium of Harvard i ity. All illustrati
made with the aid of a camera lucida.
were
MORPHOLOGICAL OBSERVATIONS
The sporangium of Anarthropteris Dictyopteris develops
from a single superficial cell of the receptacle which becomes
54 KENNETH A. WILSON
swollen and then is divided by a more or less horizontal wall
on a level well above that of the surface of the adjacent re-
ceptacular cells (fig. 1). The second wall is horizontal and
is intercalated in the lower cell of the sporangial initial and
is produced on a level with the surface of the neighboring
receptacular cells. This second division produces Segment
O and separates a basal cell from the sporangial primordium.
The sporangial primordium is therefore two-celled at this
stage, and consists of the proximal Segment O and a distal
cell, the “mother initial” (fig. 2).
The mother initial divides by the formation of three obli-
que walls which produce Segments I, II and III, that contri-
bute to the jacket layer of the capsule and to the three-rowed
portion of the stalk subtending the capsule (figs. 3, 4).
Segment O may become divided by the intercalation of a hor-
izontal wall before the formation of Segment I (fig. 10), or
preg remain undivided until after Segment I is produced
(fig. 3).
Soon after the formation of Segment I, II and III, the
mother initial, which now has the form of an inverted three-
sided pyramid (fig. 4), divides and produces a horizontal
wall which cuts off the cap cell, Segment IV, of the sporang-
ium (fig. 5). This division results in the complete enclosure
of the mother initial by its daughter cells. Following this, the
enclosed tetrahedral mother initial divides in the same order
and in the same manner as it did in producing Segments I,
Il, IL, and IV, so that it becomes enclosed by still another
layer of cells, the tapetal initials (figs. 6, 7).
The division of the tapetal initials to produce the two lay-
ered tapetum is accompanied by divisions in the central cell
— eventually give rise to the spore mother cells (figs.
-9).
Even before the formation of the tapetal initials, inter-
divisions have begun to take place within the various
segments of the sporangial initial. In fact, intercalary divi-
sions have begun to take place before the formation of
segment IV (fig. 4).
The divisions that take place in Segments I, U1, III follow
the general sequence described for Xiphopteris serrulata and
for Phlebodium aureum (Wilson 1958a, 1958b). Seg-
ment I, then, when mature is composed of a tetrad of
cells in its distal portion which forms part of one of the
lateral f; of the capsule. Beneath these four cells is a row
of three or four cells that form part of the stalk (figs. 18a,
25). Segment II is the segment in which the stomium forms,
LEPTOSPORANGIUM OF ANARTHROPTERIS 55
i is. 1-9, Internal seg-
ntation. (Both sides of each sporangial primordi-
FIGS. 1-20. Sporangial ontogeny in Anarthropteris
mentation; 10-20, Superficial segmer
um are illustrated and designated by the letters “a” and “b.”) Figs. 1-24 drawn to
seale “A,” figs. 25-29 drawn to seale “B”; both seales in microns.
56 KENNETH A. WILSON
and in addition to contributing cells to the stomial region of
the annulus, this segment also contributes cells to each face
of the capsule as well as to the sporangial stalk (figs. 19b, 25,
26). Segment II contributes cells to the formation of the
bow, to each of the sporangial faces, and to the sporangial
stalk (figs. 25, 26). The cap cell, Segment IV, becomes sub-
divided and differentiated so as to form part of the bow, a
portion of the epistomium, and also a portion of both spor-
angial faces (figs. 25, 26).
There seems to be no definite regularity in the divisions
that take place in Segment O. Although the first two or three
FIGS. 21-29. Sporangial ontogeny in Anarthropteris Dictyopteris. 21-24, Various
i 3] ial stalk: ii fi
face; 26, Mature sporangium, distal face; 27, 29, Aborted sporangia; 28, Mature
divisions in this stalk-forming segment are as a general rule
ransverse, bsequent divisions may result in the inter-
calation of tr: walls, longitudinal walls, or at times
earl
in the ontogeny of the sporangium. Longitudinal walls have
been seen in the lower cell of Segment O during the period of
the formation of the tapetal initials (fig. 6). There is no
precise sequence or pattern in the intercalation of the verti-
cal walls. Most frequently the basal cell becomes divided by
a longitudinal wall, or at other times two or more divisions
LEPTOSPORANGIUM OF ANARTHROPTERiS 57
may take place in this cell (figs. 8, 9, 15-26). Similar divi-
sions may occur in other cells of Segment O, so that one or
more cells of this sporangial segment may be subdivided by
longitudinal walls. Occasionally the divisions that take place
are oblique rather than vertical (figs. 8, 18). In some in-
stances, after the vertical or oblique divisions have taken
place, additional horizontal walls are formed which subdi-
vide the cells (figs. 23b, 26).
Although most of the vertical or oblique divisions occur in
the cells of Segment O, the lowermost cells of Segment I
also rarely divide in a similar fashion (figs. 22a, 23b).
Mixed among normal young and mature sporangia are
found both paraphyses and elongated, aborted, tannin-filled
sporangia in various stages of development ( figs. 27, 29).
Even these aborted sporangia show the longitudinal walls
in the cell of the stalk.
The paraphyses of Anarthropteris develop well before the
sporangia are initiated. Unfortunately none of the material
available was young enough to permit a study of their onto-
geny. From an examination of the mature paraphyses it is
not possible for me either to support or reject the interpre-
tation that they represent transformed sporangia.
DISCUSSION
sporangia of Phlebodium aureum (Wilson, 1958a) Xiphop-
teris serrulata, and Pyrrosia nuda (Wilson, 1958b), that
only slightly inclined and the wall is produced above the
level of the surface of the adjacent receptacular cells. This
58 KENNETH A. WILSON
same type of division occurs in the sporangial initial of
Xiphopteris serrulata, and as a result of the position of the
first-formed wall the ial stalk of Xiphopteris is one-
some number of n=37 (Brownlie, 1958) does not help very
greatly in indicating any relationship. The t: ic posi
tion of Anarthropteris cannot be establist 1 with any degree
of satisfaction without additi studies.
Criteria of Comparison. ix + 359 pp. University Press, Cam-
i d.
BROWNLIE, G. 1958. Chromosome numbers in New Zealand ferns.
Trans. Roy. Soc. New Zealand 85: 213-216, pl. 17. 1958.
LEPTOSPORANGIUM OF ANARTHROPTERIS 59
CorpeLAND, E. B. 1947. Genera Filicum, the genera of ferns. xvi +
247 pp. Chronica Botanica, Waltham, sz
cone H. B. 1951. New Zealand Ferns. Fourth ed. Revised and
ited by M. Crookes. 406 pp. Whitcombe and Tombs, Auckland,
New Zealand.
Foster, A. F., AND E. M. GIFForb, JR. 1959. — Morphology
of eaaclae _* xi + 555 pp. Freeman, San ncisco.
Witson, K. A. 1958a. Ontogeny of the sporangium xy Phlebodium
(Polypodium) aureum. Am. Jour. Bot. 45: 483-491.
1958b. Ontogeny of the sporangia in Xiphopteris ser-
rulata and Pyrrosia nuda. Jour. Arnold Arb. 39: 478-493.
——___—.. 1959. Sporangia of the fern genera allied with Poly-
podium and Vittaria. Contrib. Gray Herb. 185: 97-127.
OBSERVATIONS ON THE JUVENILE LEAVES OF
PELLAEA ANDROMEDIFOLIA
ALICE F, TRYON?
The apogamous species of Pellaea, section Pellaea (Tryon
1957) are usually wide ranging and are morphologically dis-
tinct from the sexual types. A species in California, Pellaea
andromedifolia, has apogamous members which are scarcely
distinguishable from the sexual ones and have a limited and
disjunct range. The sexual plants are diploid (n=29) and
the apogamous are probably polyploid although the count is
not yet confirmed. Other apogamous races or varieties of
this section are all polyploid (Tryon and Britton 1958). The
most reliable differences that have been found in herbarium
material of the two types are the number of spores in the
sporangium and the ponding diff in spore size.
The apogamous plants occur within the range of the more
widely distributed sexual phase. They have been identified
from Baja California and in southern California in San
Diego and Riverside counties and from Butte, Lake and
Humboldt counties in northern California. The material
with which this paper is concerned is the apogamous race
collected among serpentine rocks on a grassy hillside with
scattered oak trees. The plants grew in both sunny situa-
tions and in the shade, from crevices of large rocks (west
of Redway, Humboldt county, California, Tryon & Tryon
5556, GH). :
pores from this collection and also from a collection
of the sexual race were sown on soil in terra cotta pots
which were placed in glass covered dishes and watered
from below. Spores from apog: S$ germ ted ii
ten days, while those from sexual plants germinated in
eighteen days. The first sporelings were observed on the
neglected, and although the apogamous plants remained
vigorous, the sexual ones were lost. Es
The lamina of the adult leaf of Pellaea andromedifolia is
elongate-triangular, 6-40 cm. long, imparipinnate, usually
tripinnate (less often bipinnate or quadripinnate). There
are about ten or twelve “pairs” of pinnae, usually alternate,
“7 For the use of specimens i ium and for facilities in the green-
he eo oot ine Biclngeal Laboratory of Harvard Universicy, I am
62 A. F. TRYON
up to 14 em. long with 8-50 segments. The sterile ultimate
segments are entire or ternate and usually ellipsoidal or
ovate in form. The open venation of the sterile segment is
unequally dichotomous with a central vein and nearly equally
dichotomous ultimate veins (Fig. A). The margins of the
sterile segments are slightly crenulate and the small crena-
tions roughly correspond to the vein ends. The petioles of
the adult leaves are straw colored to ruddy brown and dark
brown only at the base or in spots or streaks above the base.
The juvenile leaves were examined on some 150 plants of
Pellaea andromedifolia of different ages, varying from those
with a single leaf on the prothallus to those with twelve
leaves. Each series (Figs. D, E, F, G, I) illustrates the
smallest to the largest leaves on a single plant. The inter-
mediate stages are arranged from the sequences observed on
several plants and may not be in strict order, but this will
not affect the general conclusions. The illustrations were
prepared from tracings of the leaves.
The initial leaf is small, usually slightly longer than the
lobes of the prothallus (Fig. B 1,2). The petiole arises from
the cushion of the prothallus and is slightly longer than to
twice as long as the lamina. The petioles of the first leaves
are dark brown or blackish near the base. This color extends
progressively farther up the petiole and in later juvenile
leaves the petioles are wholly dark, the color obscuring the
first dichotomy of the vascular system of the leaf. The lam-
ina is broader than long and cordate. It has two large lobes,
each with a pair of less developed lobes, or four nearly equal
lobes which are shallowly bifid.
The vascular strand from the petiole divides into two
equal branches near the apex of the petiole, usually in the
expanded base of the lamina but sometimes at the apex of
the petiole (Fig. C 1,2). The two principal veins each divide
equally in the base of the lamina. The subsequent branches
again divide in the mid-portion of the lamina, resulting in
eight, nearly equal, ultimate veins which end short of the
which gives the impression that they are parts of a repeated
or duplicated system. Up to this point the leaves are quite
JUVENILE LEAVES OF PELLAEA 63
unlike the adult leaf. An abrupt change of form occurs us-
ually between the sixth to the ninth leaf when a terminal
lobe, demarked by strong lateral sinuses, is developed, hav-
ing an unequally dichotomous venation with a pronounced
central vein (Fig. D 9, 10; E 9; G5, 6). This stage seems
to be the forerunner of the adult condition, and the series of
leaves which follow appear more like the adult sterile leaf.
The first leaf of the adult type is compound with three dis-
tinct pinnae borne on short, dark colored stalks, and each of
these pinnae has a central vein and ultimate dichotomous
veins (Fig. G 7). In later leaves the stalk of the terminal
pinna elongates and rachis develops (Fig. G 8). Subsequent
leaves bear additional pinnae along the rachis and these in
turn become compounded (Fig. H). As the pinnae increase
in size, the marginal lobes become less prominent crenations.
a eS
andromedifolia
Fig. B. Separate plants with initial leaves attached to prothallia, X 3-—
Juvenile leaves showing position of first vascular dichotomy, X 3.— Figs. D and E-
Sequences of ji il from indi Jants, X 3.
64 A. F. TRYON
Variations from the above sequence were noted. In some
plants the initial leaf was only bilobed, or the four-lobed type
had one or more lobes of larger proportions; or more than
four lobes were formed (Fig. E 1). It was observed that
occasionally among the first formed leaves there is one of a
more advanced form which is often attached at the base of
the prothallus near the soil. These are noticeably larger
(Fig. B 3) than the usual type; the petiole may be short and
rigid, and the lamina has a terminal lobe with a prominent
central vein. The initial leaves of this kind are usually suc-
ceeded by those with two or four lobes and equally dichoto-
mous venation but some plants were noted with more than
one leaf of this advanced form (Fig. I 2, 3).
Pena
Peer } Sf ~
aw aye VAM ILI
QU AG) ZZ,
co. : VA a
(SS AOR
/ [ace
_ Leaf-tracings of Pellaea andromedifolia — Figs. F, G, I. Sequences of leaves from
individual plants. x 3. Fig. H. A twelfth leaf with the adult form, X 114.
In Pellaea andromedifolia the sequence of juvenile leaves
ne an equally di plan of venation and
lobing through an abrupt transition to one of pinnate ar-
ement with distinct pinnae and unequal dichotomy of
JUVENILE LEAVES OF PELLAEA 65
the principal veins. The recognizable, repeated form and
sequence in these leaves is evidence that there is an inherent
pattern in the juvenile leaves. The variations from the nor-
mal sequence which have been observed may be a response
to certain nutritional states.
Cther investigations which have been made on juvenile
leaves of ferns are mainly concerned with phylogenetic re-
lationships or physiological processes. Some of these that
are pertinent to the observations on Pellaea are reviewed.
The work of R. Orth (1938), on the morphology of juvenile
leaves of 17 species of ferns native to Germany, shows that
in each species there exist certain patterns and that these
can be classified in four groups. On the basis of the four-
lobed initial leaves, Pellaea andromedifolia belongs to the
same group as Pteridium aquilinum and is similar to it in
general form. Orth illustrated the sequence of leaves on a
single plant for a few species but most of his examples are
drawn from selected stages. Comparisons are drawn with
the adult form in each species and the variations in different
parts of the same adult leaf are considered.
In his studies on Diellia, W. H. Wagner (1952a) draws
comparisons with the aspleniaceous genera illustrated b:
Orth and remarks on the limitations of the system of classi-
fication based on the initial leaves. The three species of
Diellia, illustrated by Wagner, have the first leaf entire with
a single unbranched vein, or two-lobed with equal dichotom-
ous venation and these would be placed in two different
groups in Orth’s classification. The form of the sporeling
leaves in Diellia is different from that in Pellaea andromedi-
folia in having a simple lamina with few irregular lobes.
The compound condition is reached after the ninth or tenth
leaf and the venation progresses from a simple or equally
dichotomous condition to a reticulate one. Wagner (1952b),
using a technique of mass sampling, illustrates comparative
series of juvenile leaves in Pt and P.
aureum. The series of the former resembles that illustrated
by Orth for the closely related P. vulgare more than it does
the series in P. awreum — thus, characters of the juvenile
ais support those of other parts of the plant in showing
the relationships of these three species.
Phylogenetic conclusions have been drawn on the basis
of the form and vein pattern in juvenile leaves and it is of
interest to examine these since there are two essentially op-
posed points of view. It was proposed by F. O. Bower (1923)
that an understanding of the evolution - the fern leaf and
its architecture will be found, in part, in the comparison of
66 A. F. TRYON
the juvenile and adult leaves of the same plant. He draws
such comparisons and illustrates the sequence of leaves in
Anemia adiantifolia, Osmunda regalis, and Cyathea insignis.
There are marked differences in form between the series
shown for each species ; however, the initial leaf in each case
has equally dichotomous veins and shows a progressive
change to unequal dichotomy with the development of the
central vein in the major segments. Bower (1923, p. 88)
ludes : “Compari of the juvenile leaves of other Ferns
supports the view that equal dichotomy was the prior, and
probably the original state in the construction of the leaf,
and that some form or another of dichopodium of the main
veins is a state derivative from it”.
e most recent work in which phylogeneti 1]
are drawn, in part, from juvenile leaf form is that of W.
H. Wagner (1952c), and his conclusions are quite opposed
to those of Bower. Foliar dichotomy, including vascular
dichot sis idered to a dification of ancestral
pinnately-organized leaves. Wagner (1952c, p. 591) states,
“Foliar dichotomy in living ferns appears to represent de
novo dichotomy rather than retained psilophytalean dichoto-
my”. One of the sources of difficulty in a comparison of the
theories of Bower and Wagner is a matter of terminology
and this is apparent in reference to the juvenile leaves. The
leaves which Wagner describes as midribless and midribbed
are referred to by Bower as equally dichotomous and sym-
podially dichot pectively e terms pinnate and
dichotomous have been applied as opposing terms ; however,
the former refers to a kind of arrangement and the latter
to a type of branching.
Wagner uses the juvenile leaves in Osmunda cinnamomea
dition is preceded by leaves of the two- or four-lobed form
with equally dichotomous venation.
JUVENILE LEAVES OF PELLAEA 67
Among the physiological investigations, morphogenetic
studies by R. H. Wetmore (1953, 1954) have been made on
juvenile leaves of Todea barbara, Osmunda cinnamomea,
Cyrtomium falcatum and Pteridium aquilinum var. latius-
culum. In these the juvenile leaf form was changed through
variation of the sugar concentration in the media upon
which the plants were grown. On media containing a low
a higher sucrose concentration (0.5%) the initial leaf and
those following, prior to the production of a more complex
type, were nearly all three-lobed, and the two-lobed type
was rare. Wetmore (1954, p. 34) reports on this experi-
ment: “Never was a two-lobed leaf evident on media con-
taining 1 percent and 2 percent sucrose. Here the normal
sequence was quickly dispensed with and pinnate leaves of
the adult nature were the regular type’. Wetmore examined
the meristematic centers of juvenile leaves and indicated
that the shape of the leaf is determined by meristematic ac-
tivity along the margins and at the apex of the leaves. The
two-lobed leaves result largely from marginal extension of
the meristem, and when this is supplemented by apical cell
division, the three-lobed or pinnate leaf is formed. He con-
cludes that the sequence of development as observed in
sporeling leaves is infl d by the bohydrate concen-
tration and that it is difficult to see a recapitulatory se-
uence.
Regenerated tissue on excised sporeling leaves was re-
ported by E. W. Brown (1918) in Phegopteris polypodioides
(Thelypteris Phegopteris). The new, induced tissue formed
a prothallial-like structure from which four leaves were
produced. The first two were most complex, having three
lobes each with a strong central vein, and these were fol-
lowed by a leaf with two lobes and equally dichotomous veins
and a final leaf which was entire with a single vein. The
nutritional state was considered as a possible factor in this
sequence. Whatever the cause, the example does illustrate
a sequence of leaves in reverse order of that found in the
juvenile leaves of normal plants and that the potential for
each of these forms also exists in regenerated tissue. :
The prothallus and sporeling leaves of Pteris longifolia
were studied by H. G. Albaum (1938) in an investigation
related to growth subst and metaboli: Albaum indi-
cated that upon the initiation of the first leaf the center of
growth is transfered from the prothallus to the meristematic
areas of the leaf. A successful test was made for growth
68 A. F. TRYON
substance and ted, using 3-Indole
acetic acid, which eonirdlled “the ne of sporeling
leaves.
In conclusion, it has been shown that the sequence of ju-
venile leaves in Pellaea andromedifolia proceeds from a two-
or four-lobed lamina with equally dichotomous veins to a
more complex type with the initiation of a terminal segment
anda prominent central vein formed by unequal dichotomy.
In this series as well as in those illustrated by Orth, Wagner
and Bower, certain characteristics are apparent in the ju-
venile leaves which may supplement those of the adult
leaves in understanding the relationships of ferns. The ju-
venile leaves have been examined in relatively few gion
of ferns. It i is. evident from such studies as have been mad
pertaining to the evolution of the fern leaf. The studies by
Wetmore show that, within the juvenile plant, the potential
for the production of a leaf of either simple or more complex
form may be influenced by variations in the nutritional state.
LITERATURE CITED
ALBAUM, H. G. 1938. Inhibitions due to growth hormones in fern
prothallia and sporophytes. Amer. Journ. Bot. 25 :124-132.
Bower, F. 0. 1923. The Ferns. Vol. I. Analytical Examination of the
Criteria of Comparison. 359 pp. University Press. Cambridge,
d.
Englan
Brown, E. W. 1918. in Phegopteris Polypodioides. Bull.
Torr. Bot. Club, 45:391-397.
OrrH, R. 1938. Zur logie der Primarbla inheimiscl
Farne. Flora 133:1-55.
Tryon, rg F. 1957. A revision of the fern genus Pellaea section Pel-
laea. Mo. Bot. Gard. 44:125-193.
Tryon, A. F. anp D. M. Britton. 1958. Cytotaxonomic studies on the
fern genus Pellaea. Evolution 12:137-145.
Waener, W. H. 1952a. The fern genus Diellia; its structure, affinities
and taxonomy. Univ. Cal. Publ. Bot. 26:1-212.
1952b. Juvenile leaves of two Polypodies. Amer. Fern
Jour. 42:81-85.
1952e. Types of foliar dichotomy in living ferns. Amer.
Jour. Bot. 39:578-592.
WETMORE, R. H. 1953. Carbohydrate supply and leaf development in
ferns. Science 118:578.
1954. The use of “in Vitro” cultures in the investiga-
tion of growth and differenti jation in vascular plants. Brookhaven
CONTRIBUTIONS FROM THE GRAY HERBARIUM
OF HARVARD UNIVERSITY
Epirep BY
Regp C. Rouuins AND Rosert C. Fostex
NO. CLXXXVIII
CYTOTAXONOMIC AND EVOLUTIONARY STUDIES
IN THE NORTH AMERICAN SPECIES OF
GUTIERREZIA (COMPOSITAE)
By
Orro T. Sousric
THE SOUTH AMERICAN SECTIONS OF ERIGERON
AND THEIR RELATION TO CELMISIA :
By
Orro T. SOLBRIG
PUBLISHED BY ;
THE GRAY HERBARIUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASS., U.S. A.
1960
CONTRIBUTIONS FROM THE GRAY HERBARIUM
OF HARVARD UNIVERSITY
EDITED BY
Reep C. Rotins anp Rosert C. Foster
NO. CLXXXVIII
CYTOTAXONOMIC AND EVOLUTIONARY STUDIES
IN THE NORTH AMERICAN SPECIES OF
GUTIERREZIA (COMPOSITAE)
By
Orto T. SOLBRIG
THE SOUTH AMERICAN SECTIONS OF ERIGERON
AND THEIR RELATION TO CELMISIA
By
Otro T. SOLBRIG
PUBLISHED BY
THE GRAY HERBARIUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASS., U.S. A.
Issued December 13, 1960
CYTOTAXONOMIC AND EVOLUTIONARY STUDIES
IN THE NORTH AMERICAN SPECIES OF
GUTIERREZIA (COMPOSITAE)*
Gutierrezia is an American genus of Compositae-Astereae
found in western America, Mexico, and southern and west-
ern South America. The genus is poorly known botanically.
Several species have been described, but no intensive taxo-
nomic study of them has ever been made. Morphological and
cytological studies are also lacking.
The principal objective has been to study the genus from
various approaches, such as cytology, morphology, distri-
bution, ecology, including detailed investigations of popula-
tions in nature. Unfortunately the problem of transportation
to the different localities where Gutierrezia grows, as well
as lack of time, has prevented the author from studying all
species with the same degree of intensity. This applies par-
ticularly to the South American representatives, which had
to be excluded from this study for these reasons.
This work is therefore not primarily a systematic study,
although a taxonomic revision of the North American spe-
cies is attempted. Nevertheless it is hoped that this study,
incomplete as it is, will help to clarify the taxonomy of the
genus and furnish some information about the life-history
of its species.
ACKNOWLEDGMENTS
Professor Lincoln Constance gave invaluable help during the course
of this i igation and the preparation of the manuscript. Hours of
conversation relating to subjects botanical and otherwise gave me a
better and wiser understanding of science. Professor Herbert G.
Baker’s advice was very helpful in providing other dimensions and
outlooks to the problems encountered and so was the help received from
Professor G. Ledyard Stebbins, Jr. To all three of them I express my
ks.
W. A. Weber, Messrs. J. Swanson and a 5
Francia Chisaki obtained seeds or living material of Gutierrezia
standing of the problems. I am very grateful to all of them. :
A “James Gowey Fellowship in Botany” from the University of
T Revised from a thesis submitted, in partial fulfillment of the requirements for the
degree of Doctor of Philosophy, to the Graduate Division of the University of Cali-
fornia, Berkeley.
4 OTTO T. SOLBRIG
California, Berkeley, permitted me to devote full time to research dur-
ing the last year of graduate work. Grants in aid from the “Associ-
ates in Tropical Biogeography” and the “Patent Fund” of the Uni-
versity of California made field ‘work in Mexico, South America, and
the United States possible.
I have studied material from the following herbaria, to whose cur-
ators I wish to express my appreciation for making this possible:
Academy of Natural Sciences of Philadelphia; California Academy of
Sciences; Chicago Natural History Museum; Gray Herbarium and
Arnold Arboretum of Harvard University; Jepson Herbarium, Uni-
versity of California; Missouri Botanical Garden; New York Botani-
cal Garden; Pomona College; Rancho Santa Ana Botanical Garden;
Rocky Mountain Herbarium, University of Wyoming; Southern Metho-
dist University; United States National Herbarium; University of
California, Berkeley; University of California, Los Angeles; Univer-
sity of New Mexico; University of Washington; and Washington State
University.
HISTORY OF THE GENUS
The genus Gutierrezia, with the one species, G. linearifo-
lia, was described in 1806 by Lagasca. It is dedicated to a
member of the Spanish noble family Gutierrez. The type-
material is unknown, since no specimens named by Lagasca
exist and the original description is so general that it fits
several modern species. Lagasca indicated the type locality
only as “Hab. in N.H.” (Habitat in “Nova Hispania” ?).
The only other use of the name G. linearifolia Lag. known
to me is that by Hooker and Arnott (1835) for plants col-
lected by Darwin in South America. The description and
the localities given suggest that they referred to what we
know today as G. Gilliesii. Asa Gray (1884) noted that, “G.
linearifolia Lag., the original species (of which no speci-
men named by Lagasca is extant) . . . may with the high-
est probability be referred to a Chilean species, the Brach-
yris [Gutierrezia] paniculata D.C.” This species is closely
allied to G. Gilliesii and its characters correspond to Lagas-
ca’s description.
Pursh (1814) described specimens collected by Lewis “on
the plains of the Missouri” as Solidago sarothrae. Nuttall
(1818), unaware of Lagasca’s paper, described the new
genus Brachyris and the species B. euthamiae based on ma-
terial collected by himself and on Solidago sarothrae. A
second species of Brachyris, B. divaricata, was described by
the same author in 1841. Sprengel ( 1825) changed Brach-
yris into Brachyachyris; he stated no reason for this change,
NORTH AMERICAN GUTIERREZIA 5
but it is supposed that it was purely etymological’.
De Candolle (1836) was the first to study the totality of
the material then known. He accepted Nuttall’s Brachyris,
dividing it into two sections, Eubrachyris an hiach
ris. Under Eubrachyris he placed B. euthamiae Nutt. and
five species newly described by himself; in Amphiachyris
he placed another new species, B. dracunculoides DC. In ad-
dition he established the genus Hemiachyris, with one spe-
cies, H. texana DC., and the genus Odontocarpha for a new
species from Chile, O. Poeppigii, and placed this last genus
in the tribe Vernoniae, its clearly asteroid characters not-
withstanding.
Hooker and Arnott (1841) recognized that Gutierrezia
and Brachyris were congeneric and further remarked, “We
are far from certain if the B. paniculata, euthamiae, cali-
fornica and texana are not all forms of the G. linearifolia
Lag.” This seems to indicate that Lagasca’s species was not
included among those specifically named. If this is so, the
original material must come from South America, since,
aside from the species indicated above, no other North
American species fit the original description.
Torrey and Gray, in the “Flora of North America” (1841-
43), did not include in their treatment of Gutierrezia, Am-
phiachyris dracunculoides, which had been elevated to
generic rank by Nuttall (1841). However, they did include
Gutierrezia terana (DC.) T. & G. In their study they recog-
nized fewer species than did De Candolle; Asa Gray’s treat-
ment in the “Synoptical Flora” (1884) does not differ
appreciably from that of 1841.
Hoffmann, in his h of Compositae (1897), for
Engler’s “Pflanzenfamilien”, divided Gutierrezia into three
sections, Brachyris, Amphiachyris, He h
and i is. His
characterization of the sections is similar to that of De Can-
dolle, who had described Hemiachyris as a genus.
No monographic study of Gutierrezia has been attempted
since De Candolle, and the North American species were
treated for the last time by Gray in 1884. Nevertheless,
more than 50 species have been described from North and
South America since then by various authors, particularly
E. L. Greene, R. Philippi, J. Lunell, P. A. Rydberg, and more
Tin the original description Nuttall derived the name Brachyris from Brachys: short,
and Achyris: chaffy seale.
6 OTTO T. SOLBRIG
recently S. F. Blake, A. Nelson, and A. L. Cabrera. The
genus has been studied on several occasions for different
local floras; the various authors differ in their treatments,
but they coincide in placing in synonymy a large number of
the described taxa.
CYTOLOGY AND GENETICS
The i igation of ck number and morphology
as an aid to taxonomy and evolutionary understanding is a
well known technique. The analysis of artificially obtained
interspecific hybrids has also been used widely enough to
itate no introductory emphasis. Both of these ap-
proaches have been used in the present investigation.
CROSSES AND RESULTS
Plants were grown in the greenhouse of the Department
of Botany of the University of California at Berkeley in the
spring and fall of 1956, from seed obtained partly by the
author and partly by various correspondents. The seeds
were germinated in two-inch pots and the plants were later
transferred to four-inch pots. A second transplantation was
performed, usually a year later, to eight-inch pots. Several
plants were grown from each seed collection. In the fall of
1957 a large number of the plants were transplanted to the
experimental area of the University of California Botanical
Garden in Strawberry Canyon.
To perform crosses, the maternal plants were decapitated
with a razor blade just before the tubular flowers opened,
and were then washed for several minutes with a thin jet
of luke-warm water from a plastic squeezing-bottle. The
washing was repeated twice a day until the stigmas came
out. To avoid accidental pollination by insects, the heads
were covered with a net of cheesecloth. The heads were pol-
linated by hand and pollination was also effected by tying
together heads from the parent plants under the cheesecloth
net.
All the crosses were done reciprocally between both par-
ents and were repeated several times in each case. Control
tests were run with unpollinated decapitated heads which
did not set seed in any case, and with non-decapitated heads
covered with the cheesecloth net, which did set some seed.
This was taken as an indication of self-compatibility.
A few words about seed setting in Gutierrezia species
NORTH AMERICAN GUTIERREZIA 7
might be appropriate here. It has been observed that a large
amount of Gutierrezia seed collected in the field does not
germinate. This might be due to a lack of optimal conditions
in the laboratory, but since morphological examinations in-
dicated a large amount of shrunken and apparently inviable
seed, it seems likely that the species of Gutierrezia are poor
seed producers. It is assumed that poor pollination is ac-
countable for this, but it is likely also that the physiological
bracteata
Gutierrezia. A thick full
full line indicates a sterile hybrid;
Fic. 1. Attempted artificial crosses between species of
line represents a presumably fertile cross; a thin
a dotted line indicates that no hybrid was pbtained.
balance ne2ded for seed production is very delicate and easily
upset. The possibility therefore exists that the failure of
the crosses attempted is attributable to physiological and not
genetical causes.
In figure 1 are depicted the crosses attempted and the
results obtained. As can be seen, in only one case was a cross
8 OTTO T. SOLBRIG
between two different levels of ploidy successful. The hy-
brid plant obtained from several attempted crosses between
G. californica (n = 12) and G. bracteata (n = 8), was very
slow growing and abnormal in appearance. The plant was
sterile and the meiotic analysis revealed that division of
PMC’s was arrested at metaphase I and consequently no
pollen was formed.
The only other successful cross was one between members
of two populations of G. sarothrae from Nevada and Colo-
rado, The lation from Nevada belongs to
the more robust and erect form, while that from Colorado
is a rather dwarfed form of slow growth. Several plants
were veces from this cross and the hybrids were vigorous
and of n Unfortunately, the plants were
paar ey sat before meiosis could be studied.
The planning of hybridization experiments conducted dur-
ing 1956 and 1957 was hindered at the time by a poor knowl-
edge of relationships in the genus. It is now realized that
the significance of these crosses is less than might have been
anticipated because of two main reasons: (1) the crosses
attempted were not always those which might have shed the
most light on an understanding of the evolution of the group;
and (2) the work is incomplete and more experiments are
needed. Technical difficulties also taxed heavily the success
of this approach. Nevertheless, one conclusion of taxonomic
and can perhaps be drawn. That
is the experimental confirmation of the theoretical assump-
tion that in Gutierrezia, polyploidy presents an effective
barrier to genetic interchange. Occasional hybrids between
plants with different chromosome numbers can be obtained
in the experimental garden and are found in nature (see
next tion), but if an ional exchange of genes be-
tween different polyploid levels occurs, it must be rather ex-
ceptional and of little evolutionary significance. A mechan-
ism for an interchange of genes between different levels of
ploidy, as discovered by Zohary and Nur (1959) in Dactylis,
is not likely.
CYTOLOGY
TECHNIQUE. Studies were made of mitosis in root tips
and of meiosis in dividing pollen mother cells.
Root tips were obtained both from germinating seeds and
NORTH AMERICAN GUTIERREZIA 9
from growing seedlings and fixed in Carnoy’s fixative, or
in any one of several modifications. The root tips were then
macerated in the usual way with 1 N HCl or a mixture of
concentrated HCl and 95% ethyl alcohol and then placed in
a drop of aceto-carmine or aceto-orcein on a slide and
squashed under a cover glass. Some root tips were processed
according to the Feulgen technique.
Most floral buds used for the study of meiosis were fixed
in the field; others were obtained from plants grown in the
greenhouse or the Botanical Garden. Buds were fixed in 3
parts absolute alcohol and 1 part glacial acetic acid for 24
hours and then washed twice and stored in 70% ethyl alcohol
in the refrigerator at near 0° C. until they were processed.
Propionic acid was used occasionally in place of glacial acetic
acid. Drops of a saturated solution of Fe(OH), in absolute
alcohol, which acts as a mordant, were usually added to im-
prove staining. Meiotic slides were prepared by teasing the
flowers apart in a drop of aceto-carmine, heating slightly,
and squashing under a cover slip. Permanent slides were
made whenever possible.
RESULTS. Because of the small size of the chromosomes,
studies of chromosomal morphology did not reveal any struc-
tural details. Pretreatment with paradichlorobenzene was
equally unsuccessful for the same reason. The cytological
investigation was therefore restricted to the determination
of chromosome number and to the discovery of possible
meiotic irregularities.
Gutierrezia texana and G. glutinosa, the only annual spe-
cies of the genus, revealed a haploid chromosome number of
four in all the plants investigated (Table 1). No irregu-
larities in meiosis were noted.
Gutierrezia sarothrae and G. serotina (Table 1) both have
four pairs of chromosomes. Meiosis was regular in all plants
investigated. Four of the populations of G. sarothrae studied
have eight pairs of chromosomes. Meiosis in these plants, as
far as it has been possible to detect, is perfectly normal and
no multivalents have been found.
Eight or twelve pairs were found in populations belonging
to the Gutierrezia bracteata complex. Meiosis was usually
normal, but some irregularities were di d in several
instances. Anaphase bridges were common, but no frag-
10 OTTO T. SOLBRIG
ments could be detected, — indicates that the bridges
were of the “sticky” type. Two plants belonging to different
populations are remarkable pir to merit more detailed
description.
‘TABLE 1. CHROMOSOME NUMBERS IN SPECIES OF GUTIERREZIA
Species Locality Collector n 2n
G. sarothrae Hee mi, E. of Ash Fork, Arizona Solbrig 23801 4
8 mi. E. of Hyde Park, Sa ” 2805 4
es 3 mi, W. of Aguanga, = 2760 4
9.9 mi. S. of Santa Ysabel. Calter 2 27634
2 mi. W. of Temecula, C: ” 2758 4
9.8 mi. S. of Santa Ysabel, California” 2765 (4
1.6 mi ta.
C: i o 2769 4
8.5 mi. E. of Chula Vista, California Ws 2766 4
6.2 mi. E. of Chula Vista, Cali fornia” 2768 4
11.1 mi. E. of Idyllwild, California ” 27784
Fort Collins, Colorado A, Weber 4
Virginia City, N R. H. Miller e528
Mt. N - 4
Field Creek, Orego: R. 4
‘asatch Mts., Utah R. K. Vickery Jr. 4
Wasatch Mts., Ui 2 4
5 miles N. of Payson, Arizona Solbrig 2794 8
8 mi. W. of Seli zona * 2802 8
19.8 mi. =o: of Dam, Arizo x 2792 8
Jct. Payson-Phoenix and Hwy. 488,
ike: = 2793 «8
G. ser 17 mi. E. of Tucson, amit 4
G. microcephala Silver Canyon, ee Mts., w Oalifornts Rancho peed Ana
9389 8
Morongo, San Bernardino Co.,
California 3382 8
mi. E. of Roswell, New soe G. W. Thomas 8
5 mi. W. of Bronco, Texa: » 8
cal Mexico » 8
Douglas, Arizona Solbrie 2789 16
G. bracteata Cuyamz Valley, California “ia 2166 12
Padres Nat. , Californi ” 2167 12
22.5 mi. E. of Idyllwild, California 2774 12
14 mi. N. Jet. Hidden Valley &
roads, California 2830 12 24
ape &S. So Obispo Co. line,
“ 2753 12
Taba Canyon, California ” 2775 12
» California * 159 8 16
a Creek, Yolo Co., California Rancho Santa Ana
Bot. Gard. 8 16
La Panza, California Solbrig 2751 8
7.2 mi. W. of Patterson, California a 2748 «8
G. californica © Oakland : ‘ornia ” 2154 12
G. texana Red River St., Austin, Texas B. C. Tharp 4
Austin Chalk, Austin, Texas B. C. Tharp 4
e Dallas, L. Shinners 4
slutinosa San Luis Potosi, Mexico Solbrig &
San Luis Potosi, San Luis Potosi, Ornduft 4590
Cs
NORTH AMERICAN GUTIERREZIA 11
The first is from a population in San Luis Obispo County,
California. Most of the cells have eight pairs. Nevertheless
we find a small number of multivalents (2% ) and also univa-
lents (8%). Supernumeraries, easily distinguishable by
their size and disposition at meiosis, were also present. We
are possibly in the presence of a small translocation here,
but there may be also some homologies between non-sister
chromosomes.
The other plant is from near Coalinga, Fresno County,
California. In this area both eight- and twelve-chromosome
populati eccur, ti in the same locality. In this
plant, most cells showed nine pairs of chromosomes, although
it varied from nine to eleven. Univalents and multivalents
were also present. The most plausible explanation is that
this plant might be a hybrid between eight- and twelve-
chromosome Gutierrezia bracteata, as the presence of 10
pairs in some cells would indicate, the tenth chromosome
having been lost in a pre-meiotic division. The existence of
cells with eight pairs and even eleven indicates the possibil-
ity of irregular division prior to meiosis. This does not rule
out the origin of this plant through a back-cross, but this
possibility seems to me to be remote.
Gutierrezia californica has 12 pairs of chromosomes in all
the plants studied, and meiosis was regular.
The picture presented by the Gutierrezia microcephala
complex is a very different one. The first plants investigated
came from seed received from the Rancho Santa Ana Botan-
ical Garden, and proved to have eight pairs of chromosomes
at metaphase. It was observed at the time that there were
very few PMC’s undergoing meiosis in each preparation.
This was attributed at the time to the fact that there are
only one or two flowers per head with pollen. In the fall of
1958 buds were gathered from several populations in Cali-
fornia and Arizona. These plants showed very irregular
meiosis with little or no fertile pollen. Most cells which could
be counted showed eight pairs of chromosomes, but some
had 16 or even 32 chromosomes. One population of Gutier-
rezia microcephala showed 16 pairs of chromosomes and a
regular meiosis.
As a result of these studies we may conclude that: (1)
Gutierrezia is a polyploid complex with a base number of
12 OTTO T. SOLBRIG
x = 4; and (2) different levels of ploidy are genetically iso-
lated.
DISTRIBUTION AND ECOLOGY
Gutierrezia sarothrae is the most widely distributed of
all North American species of the genus. It is found through-
out the region extending from northern Mexico to southern
British Columbia and Alberta in Canada, between the Sierra
Madre Occidental and Sierra Nevada-Cascades in the west
and the Great Plains on the east. It grows also in southern
California and in Baja California, Mexico (fig. 2). Gutier-
rezia californica is restricted to the area of San Francisco
Bay, California, while G. bracteata grows in the central and
-
i
Fic. 2. Distribution of Gutierrezia sarothrae (dotted area).
NORTH AMERICAN GUTIERREZIA 13
south Coast Ranges and from Inyo County to the Tehachapi
Range and occasionally southward to about Baja California,
Mexico (fig. 3). Gutierrezia serotina grows in an area
a Ale
Fic. 3. Distribution of Gutierrezia californica (half-full dots) and G. bracteata
(full dots). Physiographic map of California used by apecial permission of Ginn and
Company. Copyright by Ginn and Co.
14 OTTO T. SOLBRIG
around Tucson, Arizona, and occasionally southward into
Mexico. Gutierrezia microcephala, on the contrary, is more
Texas, and northern Mexico( fig. 4). Finally, Gutierrezia
texana ranges from northern Mexico to southern Oklahoma,
and G. glutinosa from central Mexico to Texas, through
northeastern Mexico and eastern New Mexico (fig. 5).
In order to discover whether or not environmental con-
ditions determine this distributional pattern, it was neces-
sary to investigate the ecology and distribution of the species
of Gutierrezia in more detail. These studies consisted of
careful observations of the type-of distribution and growth,
ecological preferences, and plant associates. Mechanical an-
alyses were performed on 19 soil samples associated with
six of the eight species, and several pH readings and some
nitrate determinations of soils were made in the field
SOIL STUDIES
Table 2 shows the results of mechanical analyses made on
19 soil samples from California, Arizona, Oregon and Idaho.
TABLE 2. MECHANICAL ANALYSIS OF 19 SOIL SAMPLES
Moisture? Organie Matter Coarse Sand
Species Coll. No, A B x A B = A x
G. sarothrae 2756 = 060 052.056 395 401.398 4.184 4,221 4,202
2763224 183.203 455 «ABZ «443 4.19 — 4.196
2766 «= 242 694.468 515.394.4540 4.721 «4.578 4.
2765 . 337 .331 917 806.861 1.146 2.481 1.813
2301 9.556.451.5038 155 .10T = .781 +390 37:
29055 430 647 649 1.664 2.257 3.417
2912 128 118 .123 139 7202.57 2.579
2792 094 092 312 584.448 «2.970 2.750 2.360
2794 = 033.051 .042 1.522 1.363 1.442 1.48 485
G. serotina ee 063 .066 if 291 287 5.452 5.394 5.423
G. microcephala 2307 024 016 123.147 185 © 6.793 6.981 6.887
27894122 107 «114 657 652 654 2.773 2.839 2.806
G. bracteata 2722034 034 450° 427 «488 3.475 3.552 3.518
2604 074 .05T 304 361.332 2.487 2.940 2.713
2755211147 .179 575 472523 1.638 1.942 1.
2723207 173.190 51l 476 = 493 6 4.799
2748 = 036. 010.023 66 382 178 5.786 6.124
ves 2743 162.142 1152 646 63 1.986 —
G. californica 2428 234 234 1.605 — 1.605 3.420 3.297 3.355
NORTH AMERICAN GUTIERREZIA 15
TABLE 2 (cont.)
Fine Sand Clay Silt Total
y. Saas 5 aes A pane Ae By cea A B =
4.132 4.766 4.449 998 .506 .752 -061 .080 9.869 10.007 9.937
4.319 — 4319 +805 _ 805 _ 6 10.055 _— 10.022
3.089 3.564 3.326 909 863 .886 091.097 9.579 12.114 10.846
1 52 3.305 2.952 3.128 146.148 9.386 10.050 9.718
4.537 4.500 4.518
1.938 3.280 2.609 158 4 8.296 9.931 9.113
94 — +294 _ 152 9.743 9.719
1.668 1.447 1.557 089 .071 © 9.299 9.627 9
1.159.988 1.073 095.075 «= 9.179 — — 8,852.
-T1T 691.704 055 .056 = 9.601 9.727 9.664
109.076 .092 100.100 9.8549, 3
1.696 1.509 1.552 O71 = .087 9.446 9.371 9.35:
774 857.815 054.058 007 7.379 8.191
685.873.1779 070 .076 = 8.241 9. 621
1.990 1.886 1.913 +134 .195 10.381 9.793 10.061
1.413 1.520 1.466 060 .055 9.407 9.662 9.533
8 380.376 099 .099 9.843 9.903 9.872
5.483 — 5.483 1,213 — 12138 3 9.582 =— 582
3.085 2.881 2.983 1.832 1.940 1.886 082.088 10.271 _ 10.154
TABLE 2 (cont.)
2 - 2 2
z Be 3 3 4
eee eee ik es eee pee 2
Roi eps Mee eat 2A Bite ea a
Species Coll. no. SS x BS x xs a
G. sarothrae 2756 0.6 4.0 42.0 44.5 TS 1.0 Sand
27680 2.0 4 42.0 48.2 8.0 6 Sand
2766 14.7 4.5 46.5 3 8.7 1.0 Sand
2769 s 8.6 18.1 34.3 31.3 15 Sandy-Clay-Loam
2301 5.0 s 6 4 Sandy:
G. serotina 2777 0.7 2.9 54.2 31.3 7.0 0.6 Sand
G. microcephala 2807 0.2 14 68.9 26.3 0.9 1.0 Sand
G. bracteata 2722 0.3 44 35.1 33.3 5 6 Sand
2604 0.6 3.3 27.1 7 73 os Sent
2755 1.8 5.2 17.9 54.6 19.1 2.0 Sandy-Loam
2723 1.9 4.9 47.2 1 4.7 0.6
2748 0.2 17 61.2 30.8 1.0 Sand
27:
G. californica 2428 2.3 16.1 33.6 29.8 18.7 0.8
The samples were taken in localities supporting a good
growth of Gutierrezia. The soil surface was scraped clean
of debris and vegetation, and the sample was taken from
the upper layers where the bulk of the roots of the Gutier-
16 OTTO T. SOLBRIG
rezia plants were found. In every case only one horizon was
involved.
The technique in the analysis of particle-size followed is
the so called “Beaker method” as described by Piper (1942).
The analyses were run on 10 grams of the soil sample after
it had been passed through a 5-mm. screen and thoroughly
mixed. Each analysis was repeated twice unless otherwise
indicated.
The analysis was intended as a rough guide to detect
possible gross requirements of the different species in re-
lation to soil texture. As can be seen, the results are rather
uniform, the variation in soil type is not great from sample
to sample, and the differences cannot be correlated with the
occurrence of any one of the species. This does not rule out
variations in tolerance to different soil types by the species,
especially to heavy soils, but this is still to be shown experi-
mentally.
Table 3 indicates the results of pH readings performed in
the field. A Beckmann portable pH meter was used in most
cases, the readings being made directly from a soil solution
extracted with distilled water. A few of the readings were
obtained by a colorimetric method. The reason for making
TABLE 3. PH DETERMINATIONS MADE ON SOILS SUPPORTING GUTIERREZIA
Species Locality pH
G. bracteata Corral Hollow, Alameda County, California 7.5
Patterson, Santa Clara County, California 7.6
La Panza, San Luis Obispo County, California 8.5
G. sarothrae Idyllwild, Riverside County, California 6.4
‘emecula, San Diego County, California 6.2
John Day, John Day County, Oregon 71
‘ansen, Twin Falls County, 0 2
Ash Fork, Coconino C rizona 8.0
G. serotina Tueson, Pima County, Arizona 8.0
these readings was the same as that which led to the under-
taking of soil.analyses, and the results were similarly uni-
form. Table 4 shows the results of a few determinations of
the content of nitrates in the soil. The nitrate was deter-
mined colorimetrically by the diphenylamine method from
a soil solution extracted with a weak acid, with the aid of a
commercial field soil kit known as the “La Motte Field Soil
Kit”. Since no significant differences were obtained, it was
deemed unnecessary to perform more of these determina-
NORTH AMERICAN GUTIERREZIA 17
Even though the analytical soil data failed to reveal any
ecological differences between species, they provide some
indication of the over-all requirements of the genus as a
whole. The North American species of Gutierrezia seem to
grow on loose, sandy, alkaline or neutral soils, and apparent-
ly those also with a low content of organic matter and
nitrates.
A careful chemical analysis of the principal elements of
the soil does not fall within the scope of this work. A chemi-
cal analysis per se, in the present state of our knowledge
about the complex soil-plant relations, would not have been
very significant in any case if it had not been complemented
by experiments to determine the importance of trace ele-
ments, differential absorption of different soil constituents,
etc. Nevertheless, it would be helpful if such data were to
be available some day, especially in relation to Gutierrezia
californica sensu stricto (see page 50). This species is of
interest in this connection because of its restriction to the
San Francisco Bay area and because it grows on serpentine
GUTIERREZIA
TABLE 4, NITRATE NITROGEN oN
Species Locality Nitrate p./million
G. bracteata Patterson, Santa Clara County, California 5
La Panza, San Luis Obispo County, California 15
G. sarothrae Temecula, San Diego County, California 15
It is known that many species of plants cannot grow on
serpentine, while others are restricted to it. Finally, a third
group can grow either on or off serpentine (Kruckeberg,
1951, 1954, 1957). The reasons for this behaviour are not
altogether clear; several authors have investigated this fas-
cinating problem and various explanations have been sug-
gested (Walker, 1954).
None of the species of Gutierrezia can be classed as a so-
called “serpentine species” with the possible exception of
G. californica. In northern California, where serpentine out-
crops are rather common, the author knows of only four
populations of Gutierrezia which grow unmistakably on
serpentine: the Oakland Hills, Point Bonita, and Angel
Island populations of Gutierrezia californica and the large
serpentine outcrops of New Idria in the central Coast
Ranges, where both eight- and twelve-chromosome popula-
tions of G. bracteata are found.
18 OTTO T. SOLBRIG
A few preliminary experiments were performed in order
to determine tolerance to serpentine in different populations
of Gutierrezia californica and G. bracteata. Seeds of the
Oakland Hills population of G. californica and of the Corral
Hollow and Cache Creek populations of G. bracteata (both
of these last with eight pairs of chromosomes) were sown
glutinosa (dotted area), G. serotina (half-
istribution of Gutierrezia
cireles), and G. grandis (full full shaded circles). Goode Base Map used by
ger permission, Dept. of Geography, Univ. of Chicago. Copyright by the University
in two different pots, one with pure serpentine soil, and the
other with a mixture of equal parts of sand and greenhouse
top-soil. In each case both sets of seeds germinated normally
and no difference could be detected relating to speed of ger-
NORTH AMERICAN GUTIERREZIA 19
mination or number of seedlings. The seedlings were later
transplanted to individual pots and grown to maturity with
the same type of soil as that in which they had been germi-
nated. In all three cases the plants on soil grew slightly
better than those on serpentine, notwithstanding the fact
Fic. 5. Distribution of Gutierrezia microcephala (crosshatched area) and G. tezana
Apes dots). Goode Base Map used by special permission, Dept. of Geography, Univ.
of Chi Copyright by the University of Chicago.
that one set of seeds (Oakland Hills) came from plants
growing on serpentine in nature.
This experiment, aside from showing that Gutierrezia
californica can grow off serpentine under experimental con-
ditions, also demonstrates that there is no element in serpen-
tine deleterious by itself to the growth of those plants of G.
bracteata which normally do not occur on serpentine.
In addition to the experiments just reported, plants from
the Oakland Hills and Angel Island populations of Gutier-
rezia californica and from the New Idria population of G.
bracteata (12-chromosome type) were grown in pots in the
greenhouse and also in the Botanical Garden in common
agricultural soil without any visible effect on them, aside
from the production of larger and more vigorous specimens
20 OTTO T. SOLBRIG
because of the greater care (mainly freedom from compe-
tition) and watering they enjoyed.
While performing these rough experiments the author ob-
served that the pots which contained serpentine required
more frequent watering in order to maintain in them the
same moisture content as in pots containing non-serpentine
soil. To verify this observation and its effect on the plants,
a group of pots with plants of Gutierrezia californica (Oak-
land Hills population) and G. bracteata (Cache Creek popu-
lation) grown in serpentine were watered with exactly the
same frequency as plants of the same populations growing
in non-serpentine soil. The plants in pots of serpentine soil
died after some two months of this treatment, due to lack
of a sufficient supply of water. This low water-retention
capacity of serpentine soils is a known characteristic (Walk-
er, 1954). In the author’s opinion, serpentine soils are to be
put in the same class with loose sand soils, which are also
poor water retainers.
Since the plants of Gutierrezia californica have a shallow
root system and grow in association with mesic grasses it
may be, therefore, that their presence in the San Francisco
Bay Area on serpentine is d ined by this “loose” charac-
teristic which gives the plants an advantage over other com-
ponents of the vegetation, especially grasses, while on the
heavier soils this situation would be reversed. In this case
it may be that the low water-retention capacity of serpen-
tine, rather than its mineral content, accounts for the growth
of Gutierrezia on it.
FIELD OBSERVATIONS
The various species of Gutierrezia usually grow in com-
pany with perennial grasses and shrubs in the open broad-
leaf shrub communities. Munz and Keck ( 1949), in their
study of California plant communities, characterize Gutier-
rezia sarothrae as a member of the Shad-Scale Scrub. Never-
theless, after visiting and studying populations of all the spe-
cies from central Mexico to Idaho, it has been observed that
the species of Gutierrezia are not restricted to defined com-
munities. This is not to say that they are not characteristic
Im a certain area of a certain type of vegetation, or that their
tolerances are so broad that they grow under almost any
conditions. This is certainly not the case: limitations of soil
and climate are as important to them as to any other plant
+,
NORTH AMERICAN GUTIERREZIA 21
species. But aside from certain broad physiographic and
ecological characteristics, in the many localities visited,
Gutierrezia was the only conspicuous pratinen present in all.
Whenever possible, a list was made of the species growing
in association with Gutierrezia (table 5). Unfortunately,
when the genus blooms in late summer or fall, most of the
herbaceous vegetation is dried up and not easily identifiable.
The list of table 5 is therefore tentative and incomplete, and
is presented only as a guide to the type of a Aire sh
may be expected to occur in association with Gutierr
‘TABLE 5. PLANTS GROWING IN ASSOCIATION WITH VARIOUS SPECIES OF GUTIERREZIA
A. Gutierrezia —Pinus ponderosa Dougl. P. cembroides Zucc., Juntperas
sarothrae.
communis L., Avena fatua L., Bromus tectorum L., B. brizaeformis L., B. japonicus
L., Poa sp., Allium sp., Atriplex canescens (Pursh) Nuttall, enone SP
Salsola is
Castilleja
rhinoides Benth., Helianthus annus L., Solidago sp., Cirsium waar ane risen:
C. undulatum (Nutt.) Spreng., gra ea canadense L., Grindelia sp., Lactuca scariola
L., Chrysothamnus nauseosus (Pall.) piss Sicdcurans ‘tridentate Note, Senecio canus
ita Achillea lanulosa Nutt., ease rage us seaber He
B. Gutierrezia californica. isa eacrvomiscfotia es Polypodium scoulert
G., Koeleria cristata (L.) es calgon pomeridianum (Ker) Kunth, Quercus
ic californica Nui
agrifolia Née, Eriogonum nudum Dousgl., ve
utt., Rhi T. & G., Silene gallica L., Esel ifornica
Cham., Arabis glabra (L.) Bernh., Dudleya sp., Lath: vestitus Nutt., ae albi-
frons Benth., Sic Gray, Sanicula bipinnatifida Dougl.,
villosa Benth., Achillea lanulosa Nutt., Artemisia californica
Cc. itierrezia — Pinus sabiniana Dougl., iperus communis L., Avena
fatua L., A. sativa L., Festuca sp., Quercus douglasii H. & A., Q. lobata Né ‘ucea
whipplei Torr., Eriogonum sp. ‘rayia spinosa (Hook.) Mogq., Adenostoma z
tum H. Euphorbia peplus ifornicum (H. &
A.) Greene, Arctostaphylos sp., Monardelia villosa Benth., Seutellaria
sp., Madia sp., Trichostemma sp., Artemisia californica Less., Lepidospartum squama-
ex Gray, ee, sp.
‘riogonum sp., Cercidium microphyllum (Torr.) _Rose
& a Totton evicoee ta juliflora (Swartz) De. Carnegia gigantea (Engelm.) Britt. &
OTHER OBSERVATIONS
Gutierrezia has a shallow root system which puts it in
direct competition with shallow-rooted herbaceous plants,
especially grasses. Since it blooms in the fall, it is in a vege-
tative state all through the summer, a season of very little
or no rainfall throughout its range. Water relations there-
fore become critical and no doubt play an important role in
the distribution of the species, both in determining the range
and the localities within the range.
The author has observed that wherever one finds a popula-
tion there is some kind of physical accident in the environ-
ment which presupposes a higher moisture content in the
22 OTTO T. SOLBRIG
soil than in the surrounding area, such as a depression, a
gulch, sometimes even a creek bed, or running water. Ex-
posure also plays an important role and it is not uncommon
to find plants growing only on north or northeast slopes.
This is probably due to the north-facing slopes having less
sun exposure, with less evaporation and less transpiration
by the plants.
It has already been mentioned that Gutierrezia must com-
pete with grasses, especially perennial ones. The competi-
tion here is believed to be mainly for the available water
supply in the ground. Xerophytic grasses are better adapted
morphologically and anatomically than Gutierrezia and ap-
parently eventually crowd it out of better soils. In poorer
soils, on the other hand, the situation appears to be reversed,
and a possible explanation may be the requirement by grasses
of more fertile soils (Tisdale, personal communication, and
my own field observations).
This dynamic balance between grasses and Gutierrezia
is expressed most clearly when the natural equilibrium is
disturbed. One frequent cause of disturbance is over-
grazing. Grasses are eaten avidly by both cattle and sheep
while Gutierrezia is eaten very little. In such cases the
latter plants spread rapidly, to the point that throughout
their range both Gutierrezia sarothrae and G. microcephala
are considered, wherever they grow abundantly in range-
lands, to be indicators of over-grazing. A second frequent
ease of unbalance is produced when the soil is disturbed by
road construction or some similar accident. Gutierrezia
plants may frequently be seen growing in the barren slopes
where the subsoil has been exposed, while the grasses rapidly
take over the better spots.
The distribution of rainfall also plays an important role
in determining the area of distribution. The concentration
of species and abund: of ind in the southwest
United States and northern Mexico seems to be directly cor-
related with late summer and fall rains. Gutierrezia bracte-
ata is, of course, an exception to this, since, after the spring
rains have ceased, it receives very little or no rain before
r-October. It is interesting, therefore, to note that
Gutierrezia bracteata is the latest bloomer, by about a month,
of all the species of the genus.
Cee
NORTH AMERICAN GUTIERREZIA 23
These field observations are substantiated by those made
on the cultures in the Botanical Garden at Berkeley. Plants
of Gutierrezia californica (Oakland Hills and Angel Island,
California, populations), G. bracteata (Yolo County, Corral
Hollow, Temblor Range, and Cuyama Valley, California,
populations), G. sarothrae (Fort Collins, and Grand Junc-
tion, Colorado; Wasatch, Nevada; and Lake Elsinore, Cali-
fornia, populations), G. microcephala (Clark Mts., Nevada,
population), G. glutinosa (San Luis Potosi, Mexico, popula-
tion) and G. texana (Austin, Texas, population), were grown
together. Water was provided in amounts sufficient so that
it never became a limiting factor. All the plants attained
much greater stature than they normally do in nature, but
their flowering time was similar to that which had been
observed in the field. The first to bloom were the Colorado
populations of G. sarothrae, about mid-June. A month later
all populations of G. sarothrae were in full bloom. In Au-
gust, G. microcephala and G. glutinosa began blooming ;
toward the end of the month the Texas populations of G. tex-
ana and the populations of G. bracteata and G. californica
were in anthesis. By October all but these last two species
had passed their blooming period. Finally, in December G.
bracteata and G. californica, especially the latter, were still
blooming, while the Grand Junction population of G. saro-
thrae was flowering for a second time.
If we correlate these observations with rainfall, we see
that the Colorado plants come from a region where the sum-
mer rains start first, and the California plants from the
region which gets its rain latest. Undoubtedl i
tal conditions can hasten or retard flowering time but, as
the experiments of Clausen, Keck, and Hiesey (1940, 1945,
1948) and Clausen and Hiesey (1958) have shown, flower-
ing time seems to be determined genetically.
VARIATION STUDIES
Prelimi barium studies revealed among the per-
ennial North American species the existence of an over-all
morphological similarity ; no clear-cut qualitative differences
could be detected. In addition all those characters, which
classically had been used to separate the species of Gutier-
rezia, showed some degree of variation. Field studies con-
firmed these observations and also showed a higher degree
1,
24 OTTO T. SOLBRIG
of inter-than intra-population variation. These preliminary
observations were restricted mainly to such characters as
size of heads and number of flowers, which have usually
been employed to characterize the various species.
In order to evaluate the amount of variation and to find
correlations, if any, with specific taxonomic units, it was
decided to study and measure the largest possible number of
characters. Another object of this investigation was to de-
termine the validity of the morphological characterization of
the species.
An approach different from the mere study of herbarium
specimens was thought to be important since studies of this
kind in the past had failed to produce satisfactory results in
Gutierrezia. It was decided therefore to study the variation
of populations in nature with the aid of simple standard
statistical methods. This, it was hoped, would produce a
better understanding of the population dynamics.
MATERIAL AND METHODS
Eleven characters were selected after some preliminary
inquiry. These are the following: height of plant; height
and width of involucre; length and width of achenes of both
ligulate and tubular flowers; length of pappus of both ligu-
late and tubular flowers ; and number of ligulate and tubular
flowers per capitulum.
A total of 17 populations, belonging to five species, from
the states of Oregon, Idaho, California, and Arizona was
studied. These populations were chosen at random. Lack
of time and transportation facilities precluded the study of
populations in other states. A sample of 50 individuals was
selected from each population. In order to randomize, each
population was measured as to length and width and a rough
outline of it was drawn on paper. A rectangle was then cir-
cumscribed around it and each side was divided into six
equal parts, and a grid consisting of 36 equal divisions was
constructed. The vertical columns and horizontal bars were
then numbered from one to six, so that each square of the
grid was identified by two numbers. Finally with the aid
of two different colored dice, ten squares in the grid were
selected. Taking a corner of the rectangle circumscribing
the population as an origin, five plants of each of the selected
Squares nearest to the origin were then chosen. After the
NORTH AMERICAN GUTIERREZIA 25
height of the plant had been measured, one head of each
plant was collected and deposited in an envelope, for sub-
sequent measurement and study in the laboratory. It is not
therefore possible to establish any correlation between a
single measurement of height of plant and any of the other
characters, although it is possible to do so for any of the
other characters.
This laborious sampling method has been followed in
order to try to eliminate any kind of voluntary or involun-
tary bias, so as to obtain a true random sample from which
meaningful statistical conclusions can be drawn. It is also
hoped that randomization has largely changed systematic
errors such as possible differences in soil, moisture, expo-
sure, etc. within the population, into fluctuating errors, and
that these have been subsequently made negligible by the
use of the common standard statistical methods used in this
study. If the sources of bias and systematic errors have been
taken care of, and we may rather safely assume so, the
samples are a true representation of the phenotypes of the
populations studied.
Gutierrezia californica was not included in this study ini-
tially. By the time it had become apparent that data from
this species would be of interest, its flowering season had
passed. Consequently, the largest number of herbarium spec-
imens of two populations well known to the author were
gathered and one head from each available plant was re-
moved and studied. This sample is less reliable than those
obtained directly in the field, since it is very possible the
collectors selected specimens for their vigour, size, or some
other special characteristic. Another source of error is
introduced by the fact that they were collected in different
years and hence under different seasonal climatic conditions.
Nevertheless, since both populations are small, the data may
have comparative value and certain ingful preli
conclusions may perhaps be drawn from this material.
STATISTICAL METHOD. Hubbs and Perlmutter ( 1942) have
provided a graphic method for the rapid comparison of rep-
resentative samples of several populations. This method is
a modification of that of Dice and Lleraas (1936) and con-
sists in plotting in a graph the range and the mean of the
sample and in addition one standard deviation and two
26 OTTO T. SOLBRIG
standard errors on each side of the mean. If the ranges of
the four-standard errors so plotted do not overlap or just
barely touch, and provided that the ratio between them is
no greater than two, the mean of the two samples can be
assumed to be significant (P=0.01 to 0.005). If there is
overlap, when it is no more than 33% of the length of the
shorter of the two four-standard errors rectangles com-
Fic. 6. Maximum (left bar), minimum (right bar), and mean (central bar) plant
measurements of fifteen populations of Gutierrezia.
NORTH AMERICAN GUTIERREZIA 27
pared, Hubbs and Perlmutter present tables (t values) which
show that the mean still can be significant (P = 0.05). In
cases where the larger four-standard errors rectangle is two
to four times longer than the shorter, an overlap of 50% or
even 75% still can be somewhat significant (P = 0.072 for
50% overlap and a ratio of 2/1; P = 0.089 for 75% overlap
anda ratio of 4/1).
This method has been used in comparing the means of
six characters: length and width of achene of tubular and
ligulate flowers and length of pappus of tubular and ligulate
flowers (fig. 9-14).
On the other hand, a frequency distribution was thought
to be more meaningful for comparing the number of both
ligulate and tubular flowers, since in this case actual number
rather than mathematical mean is the important feature.
Measurements of the involucre were combined in a pictorial
diagram so that width, length and shape can be compared.
Finally, size of plant was plotted on a map in relation to the
geographical location of the population.
RESULTS
PLANT HEIGHT. Nine populations of Gutierrezia sarothrae,
three of G. bracteata, two of G. microcephala, and one of G.
serotina were investigated as to this character. Fifty plants
were measured in every case with exception of a population
of G. sarothrae from San Diego County, California (Solbrig
2769) which consisted of only 35 plants.
Measurements were taken in the field with the aid of a
steel measuring tape. The height recorded is that between
the base of the plant and the tip of the longest branch. All
measurements of plant height are expressed in inches, since
no tape marked in meters could be found, and converting
inches to centimeters would have introduced a source of
inaccuracy.
There was an appreciable difference in the size of the
plants both between and within populations. The mean
height of the populations also varied widely, as did the range
and the standard deviation. No ingful lati
could be detected between the variation of these parameters
and any definite geographic or specific character.
The mean heights of the populations of G. bracteata and
G. microcephala are situated in the upper range of the dis-
28 OTTO T. SOLBRIG
tribution; the only population of G. serotina investigated
has the smallest mean height, while the populations of G.
sarothrae range from values not significantly larger than
those of G. serotina to some which fall well within the range
of G. bracteata.
INVOLUCRE. Size and shape of involucre is an important
character in the separation of genera in Astereae, and it
also has been employed in the delimitation of species of
Gutierrezia. It is important, therefore, to know to what
extent this structure may provide reliable taxonomic charac-
ters.
TABLE 6. MEAN, MODE, RANGE, STANDARD DEVIATION AND STANDARD ERROR
F PLANT HEIGHT
ne G
z = 4 3 ecw e 8
g Fy Se le ge g aie:
a _ a
G. sarothrae Temecula, Calif. 2758 50 22.4 22 10 38 28 13.90 1.966
Chula Vista, Calif. 2766 50 17.3 16 10 27 17 9.70 1.371
Rancho Santa Fe, Calif. 2769 35 20.3 20 13 29 12 7.48 1.263
Payson, Arizona 794 50 17.38 17 10 28 18 5.74 .811
Hyde Park, Arizona 2805 50 7. 018: 1% B18: 0
John Day, Orego: 2905 50 9.3 9 4 14 10 4.84 .684
Hansen, Idaho 2011 50 9.1 11 4 16 12 4. 5
G. bracteata Pat nm, Calif. 2743 50 15.0 15 7 18 6.62 .936
Pond Ranch, Calif. 748 50 23.7 23 14 31 17 9.88 1.397
emblor Range, Calif. 2755 50 18.5 19 12 27 15 6.90 .975
G. microcephala Saint David, Ari: 50 17.8 17 10 26 16 6.74 .953
Park, Arizona 2804 50 126 10 7 20 18 6.00 .848
G. serotina Tueson, Arizona 271T 60 7.0 7 £18 9 4.24 599
Figure 7 shows the shape and the mean dimensions of the
involucre in the populations of Gutierrezia studied for this
structure. The range of di i is illustrated by means
of two superimposed crosses, the larger being proportional
to the maximal sizes and the smaller to the minimal ones;
the range is the difference between the largest and the small-
est cross. In both crosses the vertical bar represents height
and the cross bar indicates width.
Two populations may be singled out easily from the rest:
population No. 2777, representing G. serotina, and popula-
tion No. 2804, which belongs to G. microcephala. The shape
and also the proportion between length and width (almost
as high as broad in G. serotina and several times longer than
broad in G. microcephala) are so characteristic that the two
NORTH AMERICAN GUTIERREZIA
species can be identified readily by these characters.
The involucre of G. californica is of approximately the
same shape as that of G. bracteata and G. sarothrae. The
mean dimensions of the former are larger than those of the
latter species. In spite of a slight overlap in range with G.
bracteata it is believed that this character is of some taxono-
mic value.
TABLE 7. MEAN, MODE, RANGE, STAND,
OF HEIGHT OF INVOL
JARD DEVIATION AND STANDARD ERROR
29
¢ 3 S et ee |
2 = NBs: et *
z Fa Piet es z ss 3 ipa d
& 4 ©
G. sarothrae epee mee 2758 50 43.6 45 49 35 14 5.64 .797
Chi 2766 50 35.1 31 45 25 20 2.35 .332
Rancho Sania F aFe, Calif. 2769 50 34.8 35 39 27 12 6.00 848
Payson, A 2794 50 56.5 55 65 45 20 8.52 1.205
‘Aah Fork, Aris na 2801 50 37.3 39 43 27 16 1.61
Park, Arizona 2805 50 2 40 49 35 14 2.58 364
John Day, Or 2905 9 39.3 * 45 33 12 8.58 3.031
flansen, Idaho AML gm cA ee So sea ae . .
G. bracteata Patterson, Calif. 2743 50 51.2 51 59 40 19 3.40 .480
ee Calif. 48 50 51.6 51 58 46 12 2.25 318
lor Range, Calif. 2755 50 55.7 59 63 49 14 8.60 1.216
G. microcephala ae Park, Arian 2804 50 35.9 35 41 17 24 8.38 1.185
G. serotina Tucson, 2777 +50 2: 29 «45 19 26 8.78 1.241
G. californica Angel Island, Calif, 2428 10 57.0 * © © & . .
Oakland, Cali! 2154 19 52.1 * Ge ae = -
* Data insufficient
All measurements expressed in mm. X 10.
‘TABLE 8. MEAN, MODE, RANGE,
OF WIDTH OF INVOLUCRE
$
= 2 4 7 ss 8 & ™
i : ee ee eee
4 CA
G. sarothrae Temecula, Cali 2758 50 24.8 25 19 29 10 2.68 .379
Chula Vista, 3 2766 50 18.5 19 18 27 14 2.69 .380
Santa Fe, Calif. 2769 50 21.5 19 15 25 10 4.12 .582
n 2794 50 21.2 20 18 28 10 2.48 | .350
Ash Fork, 2801 50 19.6 19 13 25 12 2.34 .330
Hyde Park, Arizona 2805 50 16.7 17 11 27 16 2.52 .356
John Day, Orego 2905 9 30.1 * 25 35 10 7.16 2.530
Hansen, Idaho Sil: 8 SLE. Fae bs +
G. bracteata Pa . Calif. 2743 50 33.9 32 29 42 18 3.56 503
Pond Ranch, Calif. 48 50 33.0 32 26 48 22 3.25 459
Temblor Range, Calif. 2755 50 26.1 29 19 35 16 7.30 1.032
G. microcephala Hyde Park, Arizona 2804 50 10.0 10 5 16 11 2.49 .352
G. serotina Tueson, Arizona 27 26.5 25 19 35 16 7.46 1.055
G. californica Angel Island, Calif. 2 Ste, Sou ee Le . td
66 18 STO FSS = .
30 OTTO T. SOLBRIG
Gutierrezia sarothrae and G. bracteata cannot be sepa-
rated on the basis of involucral characters alone, even though
in the populations studied the involucres of G. bracteata
VY 9
ica serotina
2804
microcephala
ae eace
2758 2769 2794
tie 8 é ————
a 28 ye 2gil
Fic. 7. st width of the invol Each
ao t and e involuere.
NORTH AMERICAN GUTIERREZIA 31
were slightly larger than those of G. sarothrae. The overlap
is so large and consequently the chances of error so great
that, even if we were to assume that size were a valid dif-
ference, the use of this character would lead to confusion
rather than to clarification.
Nevertheless, certain trends can be pointed out: (1) the
trend toward larger involucres in G. bracteata (see above) ;
(2) the relatively narrower involucre of G. sarothrae; (3)
the funnelform involucres of G. bracteata in contrast to the
more turbinate ones of G. sarothrae. These slight differences
in shape can be correlated with a tendency in the involucral
bracts of G. bracteata to reflex slightly and consequently to
loosen the involucres somewhat, while in G. sarothrae the
involucral bracts are closely appressed at anthesis.
Aside from the differences noted above between popula-
tions belonging to different species, figure 7 shows also that
populations of the same species differ from one another not
only in the mean of the characters considered but also in the
range. Population No. 2794 of Gutierrezia sarothrae is of
interest in this respect. As may be seen in figure 7, the in-
volueres are larger, although not broader, than in all the
other populations studied of the same species. It is thought
to be significant that this population is a polyploid with n =
8, while all the others are diploid with n= 4. It is also of
interest that the mean length of the invol of populati
No. 2794 approaches that of the 8-cl populati
of G. bracteata.
NUMBER OF FLOWERS. This character is very important
because it has been used extensively to separate species. In
this case it is believed that frequency distribution and range
are more meaningful than mean and standard deviation, and
therefore the latter will not be considered (although it has
been calculated and the interested reader is referred to tables
9and 10).
Figure 8 shows the distribution of the frequencies of ligu-
late and tubular flowers of 14 populations of Gutierrezia.
In most cases the mode class has a frequency of about 50%.
An exception to this is the only populati of G. mi phale
studied, which shows a greater frequency for the modal class
and far less variation than all the other populations investi-
gated. Herbarium studies appear to confirm this observa-
tion.
32 OTTO T. SOLBRIG
A second feature which may be observed is that the range
between the lowest and highest values observed is not
great. The Oakland Hills population of G. californica might
be an exception to this, but since the representativeness of
the sample is questionable (see page 25) this cannot be con-
firmed.
Another characteristic is that different populations of the
n 50
microcephaic
Fic. 8. Frequency
(broken line) in one head.
of number of ligulate (full line) and tubular flowers
NORTH AMERICAN GUTIERREZIA 33
same species have different modal classes and also vary in
the range. This finding is believed to be important because
all the species considered have been characterized at one time
or another by number of ligulate or tubular flowers or both.
TABLE 9. MEAN, MODE, RANGE,
OF NUMBER OF TUBULAR FLOWERS IN ONE HEAD
3 2 % aes I,
g 3 2) ee 3 6 g me RS
a 3 é za 2228
G. sarothrae Temecula, Calif. 2758 50 53 5 4 7 8 .7T 108
Chula Vist: if. 2766 50 46 5 0 6 6 1.09 .154
Rancho Santa Fe, Calif. 2769 50 5.9 6 4 7 8 .70 09
Payson, izona 2794 50 36 4 0 4 4 1.30 .183
Ash Fork, Arizona™ 2801 50 3.8 4 ek eae | 127
Hyde Park, Arizona 2805 50 3.5 4 | es Sire 50.070
Hansen, 2911 5.6 6 C9 em .
G. bracteata Patterson, Calif. 2743 50 9.0 9 6 11 5 1.09 £154
Pond Ranch, Calif. 2748 50 89 9 8s 11 3 75 = 106
‘emblor Range, Calif. 2755 50 43 4 | Pare, Soe | 86 121
G. microcephala Hyde Park, Arizona 2804 50 — 1 Coes 42 «059
G. serotina Tueson, ‘izona 2777 50 5 ae 5 10 «5 1.25 176
G. californica Angel Island, Calif. 2428 16 10.2 10 8 12 4 s °
2154 25 11.1 10 8 15 7 6.72 1.148
* Data insufficient.
* Sample taken trom two plants.
TABLE 10. MEAN, MODE, RANGE, STANDARD DEVIATION aur STANDARD ERROR
OF NUMBER OF LIGULATE FLOWERS IN 0)
g 2 z Ss 6 ¥ & x
g z - eile z g2gcgocs =
a it &
G. sarothrae Temecula, Calif. 2758 50 5.5 5 4 T 8 65 081
Chula Vista, Calif. 2766 50 5.5 5 4 T 8 147 207
Rancho Santa Fe, Calif. 2769 50 5.8 6 4 8 4 84 118
Payson, Arizo 2794 50 46 5 4 5 1 .50 070
Ash Fork, Arizona? 2801 50 44 4 3 5 2 60 .084
Park, Arizona 2305 50 46 5 4 5 1 50 .070
Hansen, I Pa eae aan ee ee, Se .
G. bracteata Ps , Calif. 2743 50 7.6 8 5 9 4 98 .181
Pond Ranch, Calif. 2748 50 65 6 5 8 8, 78 .103
Temblo: , Calif. 5 50 49 5 8 6 3 Th .104
G. hala Hyde , Aris 2304 50 —- sats OO 1 dD
G. serotina Tueson, Arizona 2777 60 6S oF S$ S$ 5 123. .178
G. californica Angel Island, Calif. MAA ES EE SOR NS *
Oakland, Calif. 2154 25 72 7 5 10 5 6.84 1.268
insufficient.
* Sample taken from two plants.
Neither in range nor in mean is there any absolute differ-
ence between species. A general trend is nevertheless pres-
ent. Gutierrezia californica has from 8 to 15 tubular flowers,
34
OTTO T. SOLBRIG
with a mode of 10, which is more than in any other species.
Ligulate flowers vary from 5 to 8 with modes of 7 and 8.
The populations of G. bracteata have a range of from 6 to
11 tubular flowers, and 5 to 9 ligulate flowers in two popula-
BEPGGga
eae
33
5 15 25
ioe os eee eee
pee ee
i
se es |
oe
——
ae
See eee
——=.
oO 8 ite 16
(2758 ch
‘2766 rm
2769 cb
2794 patente
2801 ——
(2805 ——
za re
277 —
2743 raat
2748 ==)
2785 ==
Fic. 9. Mean (vertical line), range (middle horizontal line), standard deviation
(dark lower bar) and stan
tubular
dard error (light upper bar) of length of pappus 0!
flowers. Fig. 10. Id. of pappus of ligulate flowers. For further explanations
see text.
NORTH AMERICAN GUTIERREZIA 35
tions, with modal classes of 9-6 and 8, respectively. How-
ever, a third population (Solbrig 2755) has only 2 to 6 tubu-
lar and 3 to 6 ligulate flowers and modes of 4 and 5. This
last population thus follows the pattern of G. sarothrae
rather than that of G. bracteata. This feature shows up in
other characteristics too, as will be seen. At this stage of
our investigations, however, there is insufficient informa-
tion to give an explanation for this behavior.
Gutierrezia serotina has 5 to 10 tubular and 3 to 8 ligu-
late flowers per capitulum in the population studied. The
modal classes are 8 and 7, respectively.
‘TABLE 11, MEAN, MODE, RANGE, STANDARD DEVIATION AND STANDARD ERROR,
OF LENGTH OF PAPPUS OF TUBULAR FLOWERS
- > S 2
2 3 4 2 Pie:
3 3 Pole ate tet ese crs
é 3 é = aa gs
G. sarothrae Temecula, Calif. 2758 50 146 15 12 19 7 1.60 220
a Vista, Calif. 2766 50 114 12 8 15 7 1.56 220
Rancho Santa Fe, Calif. 2769 50 9.3 10 5 1 6 106-17
, Arizona 2794 50 16.6 16 12 20 8 157 .222
Ash Fork, Arizona 2801 50 10.6 10 8 14 6 1.88 .195
Hyde Park, Arizona 2305 50 11.8 12 10 15 5 1.05 .148
sen, 2911 28 136 15 6 20 14 8.30 .634
G. bracteata Patterson, Cali 2743 50 17.9 19 15 22 7 1.77 .250
Pond Ranch, Calif. 48 50 20.2 20 16 Cie 219
Temblo: , Calif. 2755 50 17.5 18 12 26 12 298 .421
G. serotina Tucson, Arizona 50 138 12 10 20 10 3.22 .455
All measurements expressed in mm. X 10.
‘TABLE 12, MEAN, MODE, RANGE, STANDARD DEVIATION AND STANDARD ERROR
OF LENGTH OF PAPPUS OF LIGULATE FLOWERS
; 2 ea
= 3 : 2 wae >
a a =
G. sarothrae Temecula, Cal ems 50 5.8 56 4 8 4 2101 .142
Chula Vista, Calif. Bo 5.9 5 2 9 7 L6T 236
Calif. 69 50 34 8 2 5 3: 16 .10T
Payson, Arizona 50 10.7°10 7:18 6 LIT .165
Ash Fork, Arizona 2801 50 36 3 Ee ee Be nts
Hyde Park, Arizona 2805 50 5.2 5 3 9 6 .95 «184
Hansen, Idaho 2911 81 81 67 4°15 11 2.48 1483
G. bracteata Patterson, Calif. 274g 50 91 9 612 6 .9 .140
Pond Ranch, Calif 2748 50 12.4 12 9 15 6 146 .206
emblor Range, Calif. 2755 50 98 10-11 6 15 9 1.73 244
G serotina 50 10 055 14 9 «(270 240
36 OTTO T. SOLBRIG
The number of tubular flowers varies from 3 to 7 in G.
sarothrae (one head had no tubular flowers) with modes of
4or 5. The polyploid population No. 2769 had a mode of 6,
4 11 18
T T T ere rt eee
2758 =
+
2766 ——
2760 Se Se
2798 __
+
201 ane — — aes
2805 oo
29 eae
;
2777 >
——
2143 pa aa
———
748 —
2755 — i
t
2 4 Z
oo apaerer ants Vara sires i 0
2758 ata
2766 ees
2769 — :
2704 : aes
2801 ——
2808 as :
1 ne aro
29. eae
?
2TiT a REI
2743 i
Boe: pa
2755 po ee :
Fie, 11, (vertical line), range (middle cabelas line), peers devia-
tion (dark lower bo and standard error (light upper bar) of length of a of
2
tubular 2. Id. of width of achene of tubylar flowers. Poe
explanations see ee
NORTH AMERICAN GUTIERREZIA
37
as did population No. 2911. Only five heads could be counted
in this last population so that these data cannot be taken
into consideration. The number of ligulate flowers varied
from 4 to 8, while the most frequent modal classes were 5
and 6 if we disregard population No. 2911.
Gutierrezia microcephala showed 0 to 1 tubular and ligu-
late flowers. One head had 2 tubular flowers, while 1 was
the mode for both ligulate and tubular flowers.
‘TABLE 13. MEAN, MODE, RANGE, STANDARD DEVIATION AND STANDARD ERROR
OF LENGTH OF ACHENE OF TUBULAR FLOWER
3
3 = 2 EEE OR, p
3 3 Z eden Bo oes. e
i i E olibtiela. a a8
G. sarothrae Temecula, Calif. 2758 50 10.0 10 8 4 6 1.37 193
Chula Vista, Calif. 2766 50 9.3 9 6 18 7 127 179
Rancho Santa Fe, Calif. 2769 «5 88 9 eet eae | 4 32
son, Arizona 2794 50 10.8 10 8.15 T 14 202
Ash Fork, Arizo1 2301 50 9.4 10 sl 3 97 137
Hyde Park, Arizona 2805 50 92 9 8 ll 3 65 .091
n, Idal 2911 28 85 9 411 6 188 360
G. bracteata Patterson, Calif. 2743 «50 12.0 11 8 18 8 198 272
md Ranch, i 274 0 117 UW 10 16 6 1.61 227
Temblor Range, Calif. 2755 50 10.7 10 8 15 T 1147 207
G. serotina Tucson, Arizona 27 50 10.5 10 8.14 6 197 278
All measurements expressed in mm. X 10.
TABLE 14. MEAN, MODE, RANGE, STANDARD DEVIATION AND STANDARD ERROR
OF WIDTH OF ACHENE OF TUBULAR FLOWER
3 2 2
g ¢ a = |8
a 3 &
G. sarothrae 3 2758 50 3.4
Chula Vista, Calif. 2766 50 4.0
Rancho Santa Fe, Calif. 2769 503.5
» Arizor 2794 50 4.3
Ash Fork, Arizona 2301 50 4.3
Hyde Park, Arizona 2305 50 3.4
nsen, 2911 28 4.4
G. bracteata Patterson, Calif. 2743 50 4.5
d Ranch, Cali: 2748 50 4.8
Temblor Range, Calif, 2755 50 4.6
G. serotina i: 45
Tueson,
All measurements expressed in mm. X 10.
Mode
Or ON ON OT OF oo Om Om Oo
Min.
co oe 2 oo oe Oe
Max.
Agatwaaaanan
Range
a
1.40
The information presented above may be summed up in
the following tabulation:
californica
- range mode range mode range
ligulate 58 T8 39 85 88 T 48 56
tubular 8-15 10 221 49 510 8 37
total 13-23 17-18 5-20 7-14 818 15 7-15 912
sarothrae microcephala
mode range mode range mode
01 1
02 1
0-3 2
38 OTTO T. SOLBRIG
2777 a.
shee oS See
mag ——
2755 ot
2777
——
2743 ——
——
248 =
Mean vertical line), range (middle horizontal line), standard deviation
(dark ipa bar) and standard error (light upper bar) of — of achene of ligulate
flowers. - 14. Id. of width of achene of ligulate flowers. w further explanations
see ae
NORTH AMERICAN GUTIERREZIA 39
Thus, G. microcephala, and to a certain extent G. califor-
nica, may be characterized by the number of flowers in a
head. The other species cannot be identified on this charac-
ter alone. However, if the flowers of a sufficiently large num-
ber of heads are counted, some clue may be obtained as to
their identity, and in some cases a preliminary determina-
tion can be made.
PAPPUS. The pappus of the ligulate flowers is shorter than
that of the tubular ones. Nevertheless, there is considerable
similarity in the relative distribution of the means. The
mean of pappus length of G. bracteata is significantly larger
than that of G. sarothrae, with the exception of the poly-
MEAN, dipiates ipl STANDARD DEVIATION we! STANDARD ERROR
Tase 15.
LENGTH OF ACHENE OF LIGULATE FLOW
Hy 2 z eit eee |
= E=4 | Coe ae =
oe ie et
a 3 a
G. sarothrae Temecula, Calif. 2758 50 11.5 11 9 18 9 LiL 157
ula Vista, Calif. 2766 50 10.5 10 8 16 8 1.46 .206
cho Santa Fe, Calif. 2769 50 10.1 10 9 18 4 67 094
Payson, Arizona 2794 50 11.7 11 8 16 8 1.59 .224
Ash Fork, Arizo1 2801 50 11.1 10 9 15 6 41.31 .185
Park, Arizona 2805 50 10.2 10 9 12 8 61 .086
insen, Idaho 2911 31 10.1 10 15 9 1.986 1.444
G. bracteata Patterson, Calif. 2743 50 12.5 12 10 22 12 2.38
Pon ch, Calif. 2758 50 11.3 11 10 17 7 1.52 .214
Temblor Range, Calif. 2755 50 11.4 9 15 6 1381 185
G. serotina Tucson, Arizona 2777 10.2 10 6 20 14 2.02
All measurements expressed in mm. X 10.
TABLE 16. MEAN, MODE, RANGE, ST/
"ANDARD DEVIATION AND STANDARD ERROR
OF WIDTH OF ACHENE OF LIGULATE FLOWER
« > CS 2
< = - 2 4s ow
8 $ go 28 28.5 S83 =
a fe 2
G. sarothrae * 2758 50 3.0 3 56 8 100 000
Chula Vista, Calif 66 87S 2°56. 8 980.189
ncho Fe, Calif. 2769 50 32 3 2 5 8 690. 087
Payson, Ari 279450 05 5 8 OT 4 L110 357
Ash Fork, Arizona 2801 50 3.6 3 2 6 4 1.310 .185
Hyde k, i: 2305 50 3.1 8 2 & SS 10 018
Hansen, Idaho 2911 50 45 & yee Ors Se 2
G. bracteata P: , Calif. 2743 50 4.7 4 S 8S 5 1.170 .165
, Calif. 2748 50 5.1 5 & 8.5... 4 131
Temblor Range, Calif. 255 50 44 4 8 8 5 1.050 .148
G. serotina Arizona 2717 50 39 4 2 10 8 1270 .179
All measurements expressed in mm. X 10.
40 OTTO T. SOLBRIG
ploid population of this last species (see page 31). The length
of Spat of G. serotina is somewhat intermediate.
ACHENE. No significant differences in length or width of
achene could be detected. Gutierrezia bracteata had the larg-
est achenes and some populations of G. sarothrae the small-
est; however, the differences are believed to have no taxono-
mie significance.
CONCLUSION. A certain amount of variability in the char-
acters of the populations of an outbreeding species can be
predicted on theoretical grounds. The variability encount-
ered in species of Gutierrezia is therefore not surprising.
Nevertheless, unbiased estimates of the amount of variabili-
ty in higher plants are few and usually deal with woody
plants (Critchfield, 1958). Statistical methods are of com-
mon use in genetics and related fields but are used only
occasionally in taxonomic studies. The novelty in this study
lies in the sampling method, which is a variation of the “com-
pletely randomized” method, applied to observational, rather
than experimental studies.
As for the results themselves, “key” characters and their
variability are expressed in terms of frequencies and ranges
and can be dealt with in a more objective fashion as a con-
sequence. The most important aspect is that in most cases
we have a continuous range of variability for most charac-
ters throughout the range of the genus. Species therefore
become meaningful only when they are expressed in terms
of series of populations with “frequency peaks” for more
than one character, rather than in terms of presence or ab-
sence of a fixed frequency of a character (such as an « num-
ber of flowers per capitulum).
EVOLUTION
GENERAL CONSIDERATIONS
Several groups of plants in which polyploidy has played
an important role in speciation are known today (Goodspeed,
1954; Clausen, Keck and Hiesey, 1945, ete. For a review see
Stebbins, 1950). In most cases polyploid species can be sep-
arated morphologically from their diploid ancestors and
relatives. Nevertheless, some instances are known in which
the polyploid plants cannot be separated from the diploids
in any other way than by counting their chromosomes (Bell,
1954; Thompson, 1951; Cave and Constance, 1950; Heckard,
NORTH AMERICAN GUTIERREZIA 41
1969). Some of the species of Gutierrezia have polyploids
which cannot be separated either morphologically or ecol-
ogically from plants with a lower level of ploidy. Tetraploids
of Gutierrezia sarothrae with n= 8 are apparently insuf-
ficiently distinct from diploids with n = 4 to permit their
identification on gross morphology. One tetraploid popula-
tion from Payson, Arizona (Solbrig 2794) was studied in
some detail: the mean of several but not all characters meas-
ured was slightly larger in the tetraploids than the mean of
the same characters in the diploids. There nevertheless was
an overlap in the range, which seems to justify not giving
a separate designation to the polyploid in spite of the prob-
able barrier that exists to crossing. Pollen grain and stomata
diameter measurements did not reveal any significant dif-
ferences. Neither could any substantial difference be de-
tected betwen the diploids and the known polyploid plants
of G. microcephala and G. bracteata.
Stebbins (1950), after surveying the corresponding litera-
ture, has indicated that the production of a “gigas” effect
as a result of autopolyploidy is correlated with the existence
of a strongly heterozygous diploid. Assuming that the poly-
ploids encountered are autopolyploids, this might indicate
that the diploids are rather I Nevertheless, it is
the author’s opinion that the polyploids are the result of
tal allopolyploidy (Stebbins, 1950). The author has
no direct or indirect evidence to support this assumption
other than the indirect evidence of work done with other
plants (Goodspeed and Clausen, 1928; Sears, 1948 ; Schnack
and Solbrig, 1952, etc.) and the more direct, but still insuf-
ficient, proof of the perfect or nearly perfect pairing at
meiosis observed in all instances in the polyploids.
Most species of Gutierrezia grow in small to moderately
large populations. Exceptions to this are, to a certain extent,
the annual species, G. texana and G. glutinosa, the perennial
mi phala and populati of G. sarothrae in the south-
western United States. All the other species observed grew
in an over-dispersed fashion, the populations containing ap-
proximately 20 to 500 plants. The area occupied varied from
a few hundred square meters to some hectares. Each popu-
lation studied showed greater uniformity within than be-
tween populations. This pointed out that there was some
kind of isolation mechanism which prevented the free flow
42 OTTO T. SOLBRIG
of genes from one population to the other. presto. an
investigation of the breeding I
which it was felt might shed some light on this problem.
As has been indicated before, the flowers are mechanic-
ally adapted for outbreeding, even though tests performed
showed that there apparently is no genetical mechanism to
prevent selfing. In outbreeding plants one usually expects
a greater amount of heterozygosity than in self-pollinating
plants; the Astereae, so far as known, are insect pollinated
and Gutierrezia is no exception.
All plants of Gutierrezia observed are visited by insects;
the amount of visitation varied from very heavy to light.
These variations are somewhat correlated with time of day
and the number of other plants in bloom in the surrounding
vegetation. As a rule, in the months of August to October,
when Gutierrezia blooms, there are few or no other plants
flowering in the surrounding area with the exception of
shrubby Compositae (usually Astereae!). The sporadic col-
lections of insects made were not very meaningful, since no
systematic studies and observations of pollinators were at-
tempted. Nevertheless, some observations may be recorded.
In several instances populations of Gutierrezia bracteata
and G. sarothrae in California were heavily infested with
beetles of the genus Epicauta. These beetles are gregarious,
feed on pollen, and crawl over the plants in large numbers.
Undoubtedly by doing so they achieve at least some pollina-
tion. But since, unlike bees, they tend to remain on one
plant, visiting one flower after another until the food source
is depleted and only then moving to another plant, they neces-
ores pollinate mostly flowers with pollen from the same
plant.
Beetles of the genus Crossidius also were collected on
Gutierrezia sarothrae. This genus of beetles is largely con-
fined to arid regions of western North America. Its larvae
are all root borers, chiefly in roots of Chrysothamnus, Hap-
lopappus, Artemisia and Gutierrezia (Linsley, 1957). The
adults feed on pollen of the same species and also use the
heads as mating places; in doing so they contribute to the
pollination of the flowers. They are better fliers than Epi-
cauta and it is believed that they contribute to outbreeding.
Several species have been recorded by Linsley (1957) on
Gutierrezia sarothrae, G. bracteata, and G. microcephala.
NORTH AMERICAN GUTIERREZIA 43
In some cases species of Crossidius seem to be specific on
Gutierrezia.
In addition to beetles, bees were also collected. Honeybees
were observed in several instances gathering nectar and
pollen. But since this species (Apis mellifera) is of relative-
ly recent introduction in the United States, it is not consid-
ered to be as important as native bees which also were ob-
served. It is of interest to note here that no large bees, aside
from the domestic bee, were found, while some small ones
were. Ina recent monograph of Perdita, a genus of oligolec-
tic bees (Timberlake, 1954, 1958), several species are indi-
cated as feeding on Gutierrezia bracteata, G. sarothae, and
G. microcephala. In some instances the Perdita species seem
to be monolectic in relation to species of Gutierrezia, but the
collections are far too few to allow an absolute statement.
According to MacSwain (personal communication), it is
most likely that other genera of oligolectic bees feed on
Gutierrezia. Nothing is known about oligotropic and mono-
tropic bees in relation to the genus.
The species of Perdita are small and their flights probably
do not cover a large area. They would therefore contribute
to cross pollination of plants within a single population, but
they would not be very effective in carrying pollen between
populations separated by as much as a few miles. It may be
said that, in general, bees contribute to reinforce the geo-
graphical isolation of populations (Grant, 1949).
The preliminary picture that emerges from these obser-
vations is as follows: The species of Gutierrezia are mech-
anically adapted to outbreeding, although there seem to be
no barriers to selfing (see section of Cytology and Genetics).
The flowers are pollinated by insects, mainly beetles and
small bees. These insects are generally poor fliers, which,
due to their habits or constitution, tend to remain within a
population and its surrounding area. They produce outbreed-
ing between plants of the same population, hence the intra-
population variability, but they contribute also to some
extent to the selfing of plants. On the other hand, their
limited range of the geograph
barriers between populations. ws
Several papers dealing with evoluti lat of
flowers and bees have appeared recently (Grant, 1950 b;
Leppik, 1957, etc.). Others have investigated the relation-
44 OTTO T. SOLBRIG
ships between oligolectic bees and flower constancy (Linsley,
1958; Linsley and MacSwain, 1957, 1958, ete.). Oligolectic
bees can be defined ‘“‘as those in which the individual mem-
bers of the population, through its range and in the presence
of other pollen sources, consistently and regularly collect
pollen from a single plant species or a group of related plant
species, turning to other sources, if at all, only in the face
of a local pollen shortage” (Linsley and MacSwain, 1957).
The ability to recognize the pollen source is apparently trans-
mitted to the larva when it is fed with pollen of the host plant
and is most likely related to olfactory — Lavon char-
acteristics in the pollen. Morphol p
adaptations exist in both the bee, to peinit a better ‘syn-
chronization with the host, and in the plant, to attract the
insect. In the presence of a shortage of the pollen source the
oligolectic bee can change to other sources (Linsley and Mac-
Swain, 1957, 1958). It is possible, therefore, that in each
population of Gutierrezia pollinated by oligolectic bees, as
for example Perdita, the insects are adapted to a certain
pollen or flower type and favor it over other types. This
could be in response to a mutation especially favorable to
the bee (Grant, 1950a) or as a result of chance, as will be
seen.
New populations of Gutierrezia are established when new
favorable sites are open. As has seen (under Ecology and
Distribution) this may be due to such causes as over grazing
or soil disturbance. These new populations are probably in-
itiated by a few seeds (and hence there is an advantage to
the species of being capable of self-pollination), since the
seeds are not _ adapted to dispersal. Under favor-
able iti lation might in time expand to a few
hundred Sree The ‘genotypes of the first members of the
colony will be determined strictly by chance between the
genotypes present in the neighboring populations, provided
not one genotype in the sum-total of those present in neigh-
boring populations has a comparatively very high frequency.
process is part of what is known as random genetic
drift (Wright, 1949; Dobzhansky, 1953). A combination of
random genetic drift and selection by oligolectic bees could
explain the uniformity within populations of Gutierrezia
and the differences between populations (which often are
statistically significant, see Variation Studies). This is to
NORTH AMERICAN GUTIERREZIA 45
be viewed as a possible although not necessarily a probable
explanation, and it does not necessarily rule out environ-
mental or edaphic selection even if no such differences could
be detected (see Ecology and Distribution).
SPECIATION
The basic chromosome number for Gutierrezia is x = 4.
This number was found in Gutierrezia texana, G. glutinosa,
G. serotina, and in the great majority of the populations of
G. sarothrae investigated. The tetraploid populations of G.
sarothrae and the diploid populations of G. bracteata and G.
microcephala have n = 8. Twelve is found in several poly-
ploid populations of G. bracteata and in G. californica, Fi-
nally, one population of G. microcephala had 16 pairs of
chromosomes.
Four is a rather uncommon number in Compositae-
Astereae, while nine seems to be the basic number of the
tribe (Raven, Solbrig, Kyhos and Snow, 1960). Apparently
a process of chromosomal reduction has taken place in those
genera which show less than nine chromosomes (cf. also
Stebbins, Jenkins and Walters, 1951). In Gutierrezia an
increase in chromosome number through polyploidy has
taken place secondarily.
Gutierrezia sarothrae is envisioned as the North Ameri-
can species most directly related to the basic stock. All other
perennial species in North America are considered to be
derived from it or from one of its direct ancestors, either
by adaptive radiation or by polyploidy or both. This assump-
tion is based on cytological, distributional and morphological
data.
As has been stated, Gutierrezia sarothrae has four pairs
of chromosomes in most of the populati investigated
which is considered to be the basic chromosome number in
the genus. It is the most widely distributed of all the species
and its range overlaps that of all the others. It also shows
the widest ecological as well as morphological diversity. In
short, it seems to have all the characteristics necessary to be
considered the most primitive living North American species
of the genus.
Polyploid populations of G. sarothrae show what might
possibly have been the first stages in the speciation and dif-
ferentiation of G. bracteata and G. microcephala. The poly-
ploid populations of G. sarothrae, in addition to having a
46 OTTO T. SOLBRIG
double number of chromosomes, show slight deviations in
the mean of certain characters. Some of these same devia-
tions are so pronounced in G. bracteata and G. microcephala
that they can be identified readily, which is not true of the
8-chromosome population of G. sarothrae. The inconclusive
evidence supporting the view that the polyploid populations
arose as a result of hybridization between two populations
of a lower chromosomal level has been stated above. The
same process occurring between populations of G. microce-
phala and G. bracteata could account for the polyploids of
those two species, or these might be the result of back-
crossing with polyploid or diploid G. sarothrae. An attempt
to i this possibility through the study of herbari-
um ppeetnens of 12- and 8-chromosome G. bracteata and
4-chromosome G. sarothrae failed to yield any conclusive
answer. Nevertheless, there were indications that the 12-
chromosome G. bracteata might be an allopolyploid between
8-chromosome G. bracteata and 4-chromosome G. sarothrae.
The morphological characters of the 12-chromosome G.
bracteata seem to be intermediate between the characters
of the two other groups. More —_ especially of an experi-
mental nature, is needed in this
Gutierrezia serotina seems to hea an 7 exireria morphological
variant of G. sarothrae. It would be interesting to know
whether it is also isolated from it genetically. Cytological
information is lacking in the case of G. grandis, and no ideas
as to it ion can
Gutierrezia californica seems to be related to G. bracteata
and might have been derived from it. On the other hand, G.
californica is very similar morphologically to the Chilean
G. paniculata. The fact that G. californica grows in such a
specialized habitat as Scented would seem to speak against
the idea that G. californica was derived directly from G.
paniculata through long-range dispersal. Also, the fact that
a hybrid between G. californica and G. bracteata was ci
tained seems to rule out this hypoth thel
information is needed, especially with regard to G. ae
No suggestions on their evolution can be advanced in re-
spect to the annual species. The pappus of G. glutinosa is
more like that of the other species of the genus, while G.
tezana, in spite of its reduced pappus, has an involucre more
NORTH AMERICAN GUTIERREZIA 47
in accordance with the generic type. Xerophytic annuals
seem to be specialized life forms within the tribe, which
consists mostly of shrubs (Raven, Solbrig, Kyhos, and Snow,
1960). If this is so, it would reaffirm the advanced nature
of the genus. It also suggests that section Hemiachyris has
been derived from section Gutierrezia.
In luding this di ion, it is appropriate to empha-
size the importance that the type of distribution has played
in the evolution of the genus. In a group like Gutierrezia,
where we are dealing with a series of populations more or
less isolated from each other, the role of the population as
the recombination unit is greatly increased. If we consider
also the fact that most species are distributed over a wide
territory, in which profound environmental differences un-
doubtedly occur, we can easily understand the range of mor-
ical diversity
In addition to the influential micro-environmental differ-
ences which might account for part of the differences be-
tween populations, random genetic drift is thought to be
largely responsible for the morphological diversity between
populations. This is inferred from the following circum-
stances: (1) We are dealing with a series of populations
separated by relatively large distances. (2) The populations
are generally small. (3) The seed-dispersal mechanism is
rather inefficient for the family. (4) Once the populations
are established, there seems to be little chance for inter-
change of genetic material. On the other hand, the popula-
tions are rather stable in the sense that being composed of
perennial plants they do not show the yearly fluctuations in
number characteristic of populations of annuals. Neverthe-
less, field observations indicate that they respond strongly
to yearly environmental fluctuations, as measured by the
number of seedlings found. Since the ability to produce off-
spring is in the last analysis the crucial point where selection
pressures are expressed, these annual fluctuations in pro-
duction of seedlings may be as vital to the species and per-
haps of the same or a greater degree of importance for the
operation of such a process as random genetic drift, as the
more impressive fluctuations of populations of spring an-
nuals in the California deserts (Epling, Lewis and Ball,
1960).
48 OTTO T. SOLBRIG
RELATIONSHIPS WITH OTHER GENERA
The phylogenetic relationship of Gutierrezia, with other
genera is still an open question. An exl
of most genera of Astereae is needed before any conclusive
answers can be given. However, it may be appropriate to
record some suggestions that have occurred to the author in
the course of the present investigation.
Gutierrezia was placed by Hoffmann (1897) in subtribe
Solidagineae because of the homochromous character of the
capitula, i.e., both tubular and ligulate flowers are of the
same color, usually yellow. In the light of work done in other
genera of Compositae (Babcock and Cave, 1938) it would
seem that such a characteristic may not be of much impor-
tance. Nevertheless, the Solidagineae have been accepted by
most authors, even though relationships with other sub-
tribes, particularly the Asterineae are obvious.
The chromosome numbers for at least some species are
known in about half of the genera of the subtribe. Nine is
the most common number encountered in some of the large
genera such as Solidago, and sections of Haplopappus, as
well as in A Chrysopsis, and
Heterotheca (Raven, Solbrig, Kyhos, “and Snow, 1960, and
literature in same ; Solbrig, 1960).
The pappus characteristic of the tribe is one comprising
well developed hairs, as in Haplopappus. Gutierrezia and a
few other genera differ in having a scaly or reduced pappus;
these genera are Grindelia, hc et aie a Xanthoce-
phalum. Amphipappus and A y hat in-
termediate, while Hysterionica has a papel Sth a row of
hairs and a row of minute scales. All but Amphipappus,
Hysterionica, and Grindelia are closely related (Shinners,
1950; Solbrig, 1960). While Amphipappus and Hysterionica
are not closely allied to the rest of the genera cited above,
there are certain characteristics in Grindelia (such as chro-
mosome number, type of pappus, and shape of involucre)
which seem to indicate affinity with these genera. However,
probably no linear relationship exists between any of them,
and the striking similarities between some species belonging
to different genera are probably due to convergent evolution.
Finally, the heterochromous genus Greenella should be
mentioned. The morphological similarity of one of its spe-
cies, Greenella arizonica A. Gray, to Gutierrezia glutinosa
NORTH AMERICAN GUTIERREZIA 49
is unquestionable and some authors have doubted the dis-
tinctness of the genus (A. Gray, 1884; Shinners, 1950). The
other two species of Greenella, G. discoidea and G. ramulosa,
are sufficiently distinct to establish the validity of the genus.
Until more information is available, it is believed that Green-
ella arizonica should be maintained in ts present status, since
it has enough characters in addition to its heterochromous
character to separate it from Gutierrezia.
TAXONOMIC TREATMENT
Gutierrezia Lag. Nov. Gen. et Spec. 30, 1816
Brachyris Nutt. Gen. 2: 163, 1818.
Brachyachyris Spreng. Syst. 3: 574, 1825.
Hemiachyris DC. Prodr. 5: 313, 1836.
Odontocarpha DC. Prodr. 5: 71, 1836.
Suffruticose, erect, glabrous to nent Letina te te ne ao
nuals. Leaves ee: alternate,
entire, narrow. Heads corymbose or paniculate oan in
clusters or solitary at i end of branchlets or solitary in the is of
leaves, on long to very short peduncles. Heads turbinate to campanul-
ate, the involucral bracts in two to many rows, imbricated, with green
and swollen tips. Flowers yellow, the ligules not more than 144 times
the involucre; tubular flowers campanulate with very short tube and a
narrow throat with 5 small triangular lobes; stamens 5; styles of
tubular flowers with long branches very much exserted at maturity and
long stigmatic hairs in the upper part and stigmatic papillae in the
lower part of the stigmatic branches; styles of the ligulate flowers only
papillate; pappus of short to maeane aoe squamellae, shorter or ab-
sent in the ligulate flowers; pase ee slightly flattened,
tomentose, with hairs arran;
Type-species, Gutierrezia iinearifotia sy (No specimens identified
by Lagasca have been found)
SPECIES
A. Plants perennial; involucre turbinate except in No. 1; flowers 20
or fewer per head eo eae ees Section Gutierrezia.
about 10 mm. long, solitary or in small clusters at ends of
branches; leaves broad, 5-6 mm. Wide ......-.+-+--+--++ 6. G.
B. Heads less than 10 mm. long, or if longer the leaves not more
than 3-4 mm. wide
C. Involucre campanulate; heads numerous, 3-6 mm. high,
3-5 mm. broad, solitary at the ends of branchlets; plants not
more than 50 em. high .. ww. 5. G. serotina.
Cc. Involucre turbinate; heads in clusters at ends of branchlets or,
if solitary, more than 6 mm. hi:
D. Heads with only 2 or 3 flowers; involucre very narrow;
achenes of disk fi borted 4. G. mic hale
50 OTTO T. SOLBRIG
D. Heads with more than 4 = involucre turbinate; ach-
enes of disk flowers fertil
E. Heads clustered at es of Pinger flowers 5 to 10;
involucre narrow, less than 5 mm. .... 3. G. sarothrae.
E. Heads mostly solitary at ends cr feokiate? Diet
usually more than 10; involucre more than 5 mm.
F. Inflorescence Toodely corymbose; heads 6-10 mm. We
mm. wide; open, little-branched shrub
1G. ecuoring
F. Inflorescence paniculate; heads 4-7 mm. high 2-5 mm.
wide; globose, mash nace shen. .. 2. G. bracteata.
A. Plants annual; involucre tv to 1
flowers more than a per nena. Section Hi iach
. Involucre turbin eads long: 1 along
length of ae pappus of di psec absent or reduced;
tubular flowers 10-20 7. G. texana.
2
Involucre campanulate; heads at ends of branches; pappus pre-
sent in both ligulate and tubular flowers; tubular flowers 20-40
8. G. glutinosa.
1. Gutierrezia californica (DC.) T. & G. Fl. N. Amer. 2: 193, 1842
Brachyris californica DC. Prodr. 5: 313, 1836.
Perennial shrubs or bushes. Stems semi-woody, glabrous or sparsely
tomentulose, rugose or irregularly striate, mostly simple or few-
branched, arising from a short, woody base. Leaves arranged singly
ion sake branches, linear, 5 to 35 mm. long, 1-3 mm. broad, the surface
punctate, hairy, entire ‘with ciliate mar: margins. Heads solitary or some-
times in groups of 2-3 at ends of branches, arranged in loose-co:
inflorescences. Peduncles 5 mm. or more long. The involucre 6 to
10 mm. high, 4-6 mm. broad, turbinate to campanulate, the bracts ca.
obtuse, some somewhat c: carinate, the outer shorter and narrower than
the inner, coriaceous or or sub-coriaceous, usually with a green, somewhat
swollen tip, the inner bracts with a m membranaceous border, glabrous.
Receptacle flat or somewhat convex, alveolate. Ligulate flowers about
9, the ligules narrowly lanceolat ate, 3-4 mm. long, ca. 1 mm. broad;
of stigmatic papillae at the base of the stigmatic branch on each
side; the ligulate flowers with only a border of stigmatic papillae along
branch. toment-
REPRESENTATIVE 7,
2428 page Oakland, Redwi ood. Park, 1 1956 seated 2154 (ue) ; Hebe
i 1
NORTH AMERICAN GUTIERREZIA 51
Alameda County, Oakland Hills, beyond Paroltas, 10-XI-1869, A.
Kellogg and W. G. W. Harford 387 (us, MO); Hills near Oakland,
1-VII-1889, E. R. Drew (uc) ; Alameda County, dry summit of Redwood
Ridge, 21-IX-1931, Constance 382 (uc) ; Oakland Hills, VII-1888, E. L.
Fis. 15. Sestererey cabiisenion i A. Involnere X 6; By Style of tubular fower X 29
Gutierrezia
paccmaaite da cat irene tubular flower X 20.
52 OTTO T. SOLBRIG
Greene (US); San Francisco, Wilkes (Us); Near Oakland, H. G.
Bloomer (NY).
utierrezia bracteata Abrams, Bull. Torr. Club 34: 265, 1907
Cuchi californica var. bracteata (Abrams) Hall, Univ. Calif.
Publ. Bot. 3: 36, 1907.
Small suffrutescent shrub up to 50 em. high, paniculately branched,
the branches slender, flexible, the bark light to dark gray, the branch-
lets greenish, a root-crown present or lacking. Leaves arranged singly
along the branches, guns clustered at the middle of the branches,
filiform, 30-50 mm. ig 1-2 mm. wide, the surface smooth, shor
ee to glabro = pace smaller leaves (“bracts” in Abrams
original description) in the axils of branches. Heads in open- paniculate
inflorescences, single or occasionally paired at ends of oe
Peduncles 5-50 mm. long. The involucre 5-6.5 mm. high, 2-3.5 mm
diameter, conical to turbinate, involucral bracts arr: arranged in agence
mately 3 rows, narrow and elongated, carinate or at least strongly con-
becoming loose at maturity, up to 4 mm. long and 2 mm. wide. Ligulate
flowers usually 5 (3-6), 6-8 mm. long; tubular flowers usually 4 (2-6),
4-6 mm. long. Pappus of 10 to 12 scales, 1-2 mm. long in tubular flowers
and 0.5-1.5 mm. in ligulate taicng Achenes cylindrical, hairy, 1-2 mm.
aaah 0.5-1.0 mm. wide at anthesi:
ALITY. California, ay eacig g County, between Banning
oC Seven Dales XI-1889, C. R. Orew
DISTRIBUTION. Inner coastal ranges e California from Yolo County
to Riverside Counts: occasional southward to about central Baja Cali-
fornia, Mexico. (fig. a
REPRESENTA’ MENS. California. Alameda County, Corral Hol-
low, 21-X-1956, Solbrig 2159 (UC); Cuyama Valley, 54 miles E. of
Hwy. 101, 22-X1-1956, Solbrig 2166 (uc) ; Cache Creek, 5.8 miles N. W.
of Rumsey, 19-X-1954, Everett & Balls 20366 (UC, WTU, NY, RM); Los
Angeles County, Azusa, 5-VI-1902, Abrams 1577 (F, RM, UC, MO, POM,
GH, NY); Waltham Creek, 5 miles W. of Alcalde, 14-XI- 1936, Belshaw
2735 (UC, WS, Mo, GH) ; Upper end of Cuyama Valley, 18-XI-1932, Wolf
4417 (A, Ws, WTU, UC); Corral Hollow, 18-X-1861, Brewer 848 (ws,
US, UC, Mo).
3. Gutierrezia sarothrae (Pursh) agen fe Rusby, Trans. N. Y.
Acad. Sci. 7: 10,
Solidago sarothrae Pursh, Fl. Amer. jad 2: 540, 1814; Xantho-
cephalum sarothrae (Pursh) Shinners, Field and Lab. 18: 29, 1950.
euthamiae Nutt. Gen. N. Amer. Pl. 2: 163, 1818; Brachy-
achyris euthamiae (Nutt.) Spreng. Syst. 3: 574, 1825; Gutierrezia
euthamiae (Nutt.) T. & G. Fl. N. Amer. 2: 193, 1841-43. Based upon
Nuttall, cor the arid plains of the Missouri from the Arikarees to the
Bracyris divariata Nutt. Trans. Amer. Phil. Soc. 7: 313, 1841;
Gutierrezia div aricata (Nutt.) T. & G. Fl. N. Amer. 2: 194, 1841-43.
Based upon Nuttall, “On the Platte near the Rocky Mountait 38
NORTH AMERICAN GUTIERREZIA 53
Gutierrezia Haenkei Sch. Bip. Flora 38: 115, 1855. Based upon
Haenke, “Mexico, Acapulco”.
Gutierrezia juncea Greene, Pittonia 4: 56, 1899. Based upon Skehan,
“near Gray, New Mexico”, VIII-1898.
Gutierrezia diversifolia Caos Pittonia 4: 53, 1899. Based upon
Watson 551, “Laramie, Wyoming”.
Gutierrezia divergens Greene, Pittonia 4: 58, 1899. Based upon
Parish, “San Bernardino Mesas” [California].
Gutierrezia lepidota Greene, Pittonia 4: 57, 1899. Based upon Greene,
“Plains about Grand Junction, Colorado”, 27-VIII-1899.
Gutierrezia fasciculata Greene, Pittonia 4: 56, 1899. Based upon
Greene, “Grand Junction, Colorado”, 26-VIII-1896.
Gutierrezia filifolia Greene, Pittonia 4: 55, 1899. Based upon E. O.
Wooton, “White Mts. of New Mexico”, 24-VIII-1897.
Gutierrezia longifolia Greene, Pittonia 4: se ee. Based upon E. O.
Wooton, “White Mts. of New Mexico”, VIII-18:
Gutierrezia tenuis Greene, Pittonia 4: 55, gn Xanthocephalum
tenuis Ce) Shinners, Field & Lab. 18: 29, 1950. Based upon
“Foothills of the mountains back of Silver City, New Mexico”,
30- 1X 1880.
Gutierrezia myriocephala A. Nels. Bot. Gaz. 37: 264, 1904. Based
upon A. Nelson 8645, “near Badger, Laramie County, Wyoming”, 3-IX-
1901.
Gutierrezia eas ia Rydb. Bull. Torr. Club 31: 647, 1904. Based
upon Clements 16, “Manitou, Colorado”, 1901.
Gutierrezia busaes Rydb. Bull. Torr. Club 31: 647, 1904. Based
upon Earle 474, “Gray, Lincoln County, New Mexico”, 1900.
Gutierrezia laricina Greene, Rep. Sp. Nov. 7: 195, 1909. Based upon
C. R. Orcutt, “Colorado Desert in California”, X-1889.
Gutierrezia Goldmanii Greene, Rep. Sp. Nov. 7: 195, 1909. Based
upon E. A. Goldman, “Florida Mts., in extreme southern New Mexico”,
8-IX-1908.
Gutierrezia furfuracea Greene, Rep. Sp. Nov. 7: 195, 1909. Based
upon E. A. Goldman, “Cactus flat, upper Rio Gila, southeastern New
Mexico”, 13-X-1908.
Gutierrezia fulva Lunell, Amer. Midl. Nat. 1: 235, 1910. Based upon
J. Lunell, “Des Lacs, Ward County” [North Dakota].
Gutierrezia Greenei Lunell, Amer. fone ls Nat. 1: 233, 1910. Based
upon J. Lunell, “Leeds, North Dakota’
rrezia ionensis Lunell, Amer. Midi. Nat. 2: 194, 1911. Based
upon J. Lunell, “east of Ione, Morrow County, Oregon”, 16-VII-1903.
Gutierrezia sarothrae var. pauciflora Eastw. Proc. Calif. Acad. Sci.
IV, 18: 480, 1929. Based upon Mason 1971, “Turtle Bay, Lower Cali-
fornia, Mexico”, 2-VI-1925.
Gutierrezia globosa A. Nels. Amer. Jour. Bot. 23: 265, 1936. Based
upon A. Nelson ek 739, “Between San Ysidro and Bernalillo, New
Mexico”, 29-VIII-1
Duele co anne nes ae. Jour. Bot. 23: 265, 1936. Based
54 OTTO T. SOLBRIG
upon A. Nelson, “Ten miles east of Santa Fe, New Mexico”, 30-VIII-
1931.
Shrubs, 15-90 em. high, profusely branched prevalently from the base,
the secondary branching variable, usually slight and concentrated in
the upper part, from shallow, fasciculate roots, or one central taproot
Root-crown woody, up to 7.5 em. thick. Leaves of two types, the primary
scattered along the stems, 2-7 cm. long, up to 3 mm. wide, the secondary
fascicled in axils of primary leaves or branchlets, up to 2 em. lon
1-2 mm. wide, linear-lanceolate, entire, glabrous to tomentulose, often
with regular amounts of resin on both surfaces. Heads in corymbose
inflorescences, usually in fascicles of 3-10 at the ends of branchlets,
sometimes single. Involucre 3-10 mm. long, 2-5 mm. wide, turbinate,
involucral bracts narrow, acute, with a green and swollen tip, imbric-
and often covered with resin. Ligulate flowers usually 4 or 5 (3-7),
2-5 mm. long; tubular flowers 3 or 4 (2-6), 2-3 mm. long; pappus about
as long as achene, of 8-10 scales, in ligulate flowers 1/2 -1/3 shorter.
Achenes terete, 1-2 mm. long, hairy.
TYPE-LOCALITY. “On the plains of the Missouri”, Lewis.
DISTRIBUTION. Between the Cascades-Sierra Nevada and the Rocky
Mountains from southern Canada to northern and central Mexico,
eastern foothills of the Rocky Mountains and infrequent on the Great
we Also in southern California, and Baja California, Mexico
ep
REPRESENTATIVE SPECIMENS. Canada. Along Highway from Calgary
to Waterton Lakes, near Pincho, 31-VIII-1938, Nelson & Nelson 3122
(RM, MO, UC). Mexico. Chihuahua. Rocky hills near Chihuahua, 26-IX-
1885, Pringle 337 (PH, GH, US, WTU). Baja California. Between Ojos
Negros and Neji Rancho, 16-IX-1929, Wiggins & Gillespie 4155 (F, MO,
NY, A, GH, US). United States. Arizona. 12.6 miles E. of Ash Fork,
26-VIIT-1958, Solbrig 2301 (uc). 5 miles N. of Payson, 26-VIII-1958,
Solbrig 2794 (uc). California. Dry Ridges, Bear Valley, 7-VIII-1902,
Abrams 2916 (La, UC, Ny, WTU, PH, US, F). Ramona, X-1903, Brandegee
(GH, US, Mo, RM, NY). 2 miles W. of Temecula, 18-VIII-1958, Solbrig
2758 (Uc). Colorado. Denver, 4-IX-1920, Clokey 2880 (US, GH, NY, UC,
WTU, MO, F,RM). Greenhorn, 2-IX-1921, Clokey 4320 (ws, Mo, WTU, RM,
Uc, Us). Buena Vista, 1-2-VIII-1919, Eggleston 15348 (GH, NY, MO, F).
Canyon, Charleston Mountains, 9-VIII-1937, Clokey 7737 (mo, SMU, UC,
NY, RM, WTU, Ws). Kyle Canyon Fan, Charleston Mountains, 9-VIII-
1938, Clokey 8155 (Mo, Ny, F, UC, RM, WTU, ws). New Mexico. Sierra
Blanca, 18-VIII-1904, Metcalfe 1230 (UNM, MO, Ny, UC). Pyramid
Peak, 16-VIII-1930, Fosberg S3775 (px, MO, UC, US). Oklahoma. Altus,
24-X-1936, Hopkins & Van Valkenburgh 695 (SMU, UC, US, RM, MO, WTU,
Ws). Texas. Randall County, Palo Duro State Park, 20-X-1945, Cory
50407 (UC, SMU, US). Culberson County, north of Van Horn, 10-X-1944,
Waterfall 5780 (cu, MO, SMU). Utah. Western Bear’s Ear, Elk Moun-
NORTH AMERICAN GUTIERREZIA 55
tains, 2-VIII-1911, Rydberg & Garrett 9370 (RM, US, UC, NY). Washing-
ton. Asotin County, north of Rogersburg, 7-IX-1936, Constance 1813
(GH, MO, US, WTU, WS). Wyoming. Laramie, 12-IX-1899, Nelson &
Nelson 6858 (RM, NY, GH, MO).
G. Stamens
flower X 20; I. Aborted pollen grains X 1000; 4. Siple of ligulate flower X 20: K.
voluere X 10; L. Ligulate flower X 10; M. Tubular flower
56 OTTO T. SOLBRIG
This species is the most abundant, the most widespread,
and the most variable. This variability, associated with a
disjunct type of distribution, has led those botanists prone
to hasize small diff to describe a large number
of taxa, as can be seen by the long list of synonyms. Never-
theless, when one considers the group in its totality, one
realizes that these “species” represent only special combina-
tions of characters, often not even representing one breeding
population, but just a few isolated individuals. Ni evertheless,
some of the described taxa might be deserving of varietal
rank. Although no subspecific taxa have been proposed since
it is felt that more work is needed, certain regional types
might be mentioned.
(1) Southern California and Baja California, Mexico.
Some specimens from this region have the heads solitary and
terminal, which causes them to resemble G. bracteata. In-
tensive study of some populations shows, nevertheless, that
they belong to G. sarothrae on the basis of a large series of
plants which show the typical characters. They also have
the chromosome number of G. sarothrae.
(2) North of Utah-Colorado line. These are small, very
globose plants, some 20 em. tall. Specimens of this type were
cultivated at the University of California Botanical Garden
Sait sana . 7
h
their mor gical characteristics. They
apparently hybridize freely with more typical forms and
probably represent an ecotypic form. The type of the spe-
cies comes from this area,
(3) Arizona, New Mexico, and Mexico. This common
form has few flowers, a very narrow involucre and is rem-
iniscent in aspect of G. microcephala. All kinds of in-
termediates to the typical form occur, however, and make
an interpretation difficult. These plants have fertile tubular
flowers and more involucral bracts than G. microcephala,
and hence can be easily separated from that species.
(4) Uintah Mountains in nortt Utah. A few
specimens with large heads, very similar to those of G.
bracteata, have been collected in this area. Not enough
information is available on this material, and since there
are also intermediates, a decision will have to be made when
more is known about the Gutierrezias in this area.
NORTH AMERICAN GUTIERREZIA 57
4. Gutierrezia microcephala (DC.) A. Gray, Pl. Fendl. 74, foot-
note 18,
Brachyris microcephala DC. Prodr. 5: 313, 1836; Gutierrezia eutha-
miae T. & G. var. microcephala (DC.) A. Gray, Syn. Fl. N. Amer. 1
(2) : 115, 1884; G. sarothrae var. microcephala (DC.) Benson, Amer.
Jour. Bot. 30: 631, 1943; Xanth hal i phal (DC.) Shin-
ners, Field & Lab. 18: 29, 1950.
Xanthocephalum lucidum Greene, Pittonia 2: 282, 1892; Gutierrezia
lucida (Greene) Greene, Fl. Francis. pt. 4: 361, 1897. Based upon a
imen from “the region of the Mohave Desert and southward”
[California], no collector indicated.
Gutierrezia linoides Greene, Leafl. Bot. Obs. 2: 22, 1909. Based upon
J.C. Blumer, “Limestone of the Chiricahui Mts.” [Arizona], 1907.
Gutierrezia glomerella Greene, Pittonia 4: 54, 1916. Based upon
Wooton 449 “Organ Mts., New Mexico”.
Gutierrezia digyna Blake, Contr. U.S. Nat. Herb. 22: 591, 1924;
Xanthocephalum digynum (Blake) Shinners, Fleld & Lab. 18: 29, 1950.
Based upon V. L. Cory 26,335, “12 miles east of Marfa, Presidio County,
Texas”, 19-X-1937.
Globose shrub, 30-100 em. high. Branches slender, grayish to
yellowish-green, about 1 mm. in diameter, glabrous, slightly fistulate,
arising from a well developed root-crown or short stem, up to 1-2 cm.
in diameter, bark rugose, gray. Leaves narrow-lanceolate of two types,
cauline 2-5 em. long and 2-4 mm. wide, axillary leaves shorter and much
narrower, often borne in fascicles, lanceolate, entire, often resinous, or
either type absent, the axillary in young plants and the cauline in older
specimens or under special environmental conditions. Heads in glome-
rules at the end of branches, sessile. Involucre 3-4 mm. long, 1-1.5 mm.
wide, narrowly turbinate, involucral bracts less than 10, lanceolate,
with a slightly thicker and greenish tip. Ligulate flowers usually 1 or
2 (up to 4), 3-4 mm. long; tubular flowers 1-3, 2-3 mm. long. Pappus
of few scales up to 2-3 mm. long, shorter in the ligulate flowers. Styles
developed hairs, that of ligulate flowers only with stigmatic papillae.
Achenes of tubular flowers aborted, those of the ligulate flowers fertile,
2-3 mm. long, hairy. i
TYPE-LOCALITY. “In Mexico prope locum dictum Saltillo”, Berlandier.
hwest United States from western Texas to
California, and northern and central Mexico (fig. 5).
REPRESENTATIVE SPECIMENS. Mexico. ila. Saltillo, 1898, Palmer
143 (uc, F, MO). United States. Arizona. Douglas, 23-VIII-1958, Sol-
brig 2789 (uc). 3.5 miles S. of St. David, 23-VIII-1958, Solbrig 2790
(uc). Ft. Lowell, 20-X-1903, Thornber 93 (Us, uc). California. San
ino and Riverside County line, road from Yucca Valley to San
Bernardino, 27-VIII-1958, Solirig 2807 (uc). Wyman Canyon, White
Mountains, 29-VI-1931, Duran 3151 (UC, F, MO, US, GH). Colorado.
Naturita, 11-VIII-1914, Payson 595 (GH, MO, WS, F)- Deer Ann, 25-
VIII-1901, Baker 915 (Mo, Us, UC, GH). Nevada. Kyle Canyon, Charles-
ton Mountains, 27-V1-1936, Clokey 7413 (GH, UC, PH, SMU, US, MO, WS,
58 OTTO T. SOLBRIG
WTU, F). Texas. Vicinity of El Paso, 8-X-1913, Rose & Fitch 17847 (Us,
MO). rmstrong and White Canyons, near the Natural Bridges,
4-6-VIII-1911, Rydberg & Garrett 9425 (US, UC, NY).
5. Gutierrezia serotina Greene, Pittonia 4: 57, 1899
Gutierrezia polyantha A. Nels. Amer. Jour. Bot. 25: 117, 1988. Based
upon Nelson & Nelson 1638, ‘a few miles north of Tucson, Arizona”,
26-IV-1935.
Small hemispheric shrub, 10-40 cm. high. Branches leafy to the top,
green or lightly gray, slightly striate, glabrous or nearly so, with very
few or no side branches, abundant from root-crown, 2-4 cm. in diameter.
Root a slender taproot with abundant secondary roots. Leaves single
along the branches or in fascicles at axils of single leaves in lower
third of branches, linear, 1-2 cm. long and up to 1 mm. wide, glabrous.
Heads solitary oF in loose glomerules at the tip of branches, arranged
in corymbose i % sessil le or short: Inv Neo
2-4 mm. high, 2-3 mm. wide, I to turbinat:
volucral bracts broad, almost as wide as long, obtuse, pooner screed
membranaceous, with a slightly thickened, greenish or brown tip, ar-
in 2 or 3 loose series. Ligulate flowers about 7 (4-8), 4-5 mm.
long; fabarae flowers about 8 (5-10), 3-4 mm. long, salverform. Pappus
of 10-12 lanceolate scales, 1.4 mm. (1-2) long in the tubular flowers, 1
mm. (0.5-1.5) long in the ligulate ones. Achenes of ligulate and tubu-
lar flowers approximately the same size, 1.5 mm. long; 0.5 mm. wide,
hairy, with the trichomes arranged in parallel rows.
TYPE-LOCALITY. “Plains about Tucson in southern Arizona”, 3-III-
1892, Toumey.
DISTRIBUTION. Plains about Tucson, occasional southward into Mex-
ico. (fig. 4)
REPRESENTATIVE SPECIMENS. Arizona. Tanque Verde Ranch, 17 miles
E. of Tucson, 22-VIII-1958, Solbrig 2777 (uc). 7.9 miles E. of Oracle
Junction, 23-VIII-1958, Solbrig 2791 (uc). Oracle, Spring 1941, H. S.
Gentry 6111 (uc). Martinez’s Ranch, 16 miles E. of Tucson, 8-IV-1940,
L. J. Brass 14305 (GH, Ny, Uc). Redington Pass, Pima County, 28-IV-
1938, R. C. Foster 617 (cu,uc). Near Willcox, 25-V-1928, R. H. Pee-
6. PPS grandis Blake, Contr. U. S. Nat. Herb. 22:591, 1924
Xanthocephalum grande (Blake) Shinners, Field & Lab. 18: 28, 1950.
Large shrub, 30-60 cm. high. Branches few, not more than 2-4 mm..
mm. le, margin y glab:
multinerved, the central nerve and to a certain extent the lateral ones
Prominent. Heads few, solitary at the ends of branchlets or in glo-
merules of 3-5, arranged in loose oose corymbose inflorescences, sessile or
pedunculate, peduncle 1-10 mm. long. Involucre 5-6 mm. high, 3-4 mm.
NORTH AMERICAN GUTIERREZIA 59
wide, turbinate, appressed, involucral bracts elliptical, obtuse, in sev-
wi
mm. high. Pappus of about 10 scales up to 1 mm. in length in the
tubular flowers, somewhat shorter in the ligulate ones. Achenes about
1-2 mm. long, hairy.
TYPE-LOCALTY. “Head of Cafion de las Baretas, in the Sierra Madre,
near Icamole, Nuevo Leén, Mexico”, 3-II-1907, W. E. Safford 1257.
DISTRIBUTION. Th’s species is known only from a few localities in
Sierra de Parras, X-1910, C. A. Purpus (MO, NY, F, US, UC, GH). San
Potosi. Sierra de Catorce, no date, F. Rodriguez (F). Monts du
Real de Catorce, 1827, Berlandier 1360 (GH). Potrero, 8-IX-1939, F.
Shreve (uc, GH).
7. Gutierrezia texana (DC.) T. & G. Fl. N. Amer. 2: 194, 1842
Hemiachyris texana DC. Prodr. 5: 314, 1836.
Robust annual, 20-80 em. high. Stems one to several, subdivided or
not, 2-4 mm. diameter, grayish-green, slightly ribbed, glabrous, branch-
lets grayish-green, fissured, glabrous or very slightly pubescent, not
more than 1 mm. in thickness, arising from base. Root a slender tap-
root not more than 10 em. long. Leaves lanceolate, up to 5 em. long
and 5 mm. wide, acute, glabrous, entire, su’ rugose or smooth, mul-
tinerved with the middle vein prominent on both surfaces, the lateral
longitudinal ones less so. Heads at the ends of branchlets and axils
of leaves and branchlets. Peduncles 2-3 mm. long. Involucres 3-4 mm.
high, 3-6 mm. broad, turbinate-campanulate, involucral bracts numer-
ous, about 3 mm. long and 1.5-2 mm. wide, in about three irregular
series, elliptical, margins membranaceous, tips green, midrib region
darker-colored. Ligulate flowers 10-15, tube 1-2 mm. long and ligule
2-3 mm. long and 2 mm. wide; tubular flowers about 12-18, corolla
campanulate, throat very short, about 0.5 mm. long, tube 1.5 mm. long.
Pappus of about 10-12 scales, in the tubular flowers not longer than
12 mm., in the ligulate ones reduced to a crown of minute setae about
stigma b with
‘lae only and the stigma branches shorter in the ligulate flowers.
Achenes of both ligulate and tubular flowers developed, 1 mm. long,
‘0.5 mm. wide at anthesis, growing to twice that size at ity in
some cases.
TYPE-LOCALITY. “Mexici prov. Texas”, Berlandier 1763, 1765.
southern Oklahoma to San Luis Potosi, Mexico,
REPRESENTATIVE SPECIMENS. Illinois. East St. Louis, 1897, Letter-
‘man (MO, PH, NY, US). Texas. Near Dallas, VIl-VIUIL.-, Reverchon
1297 (NY, MO, US, F, GH). Comanche Springs, 1849-1851, Lindheimer
921 (Mo, F, US, UC). Dallas, 27-X-1900, Bush 1624 (NY, GH). Fort
Worth, 30-IX-1902, Tracy 8159 (GH, F, NY, mo). Austin, 20-X-1937,
“Tharp 283 (PH, WS, UC).
60 OTTO T. SOLBRIG
8. Gutierrezia glutinosa (Schauer) Sch. Bip. Flora 38: 115, 1855
Hemiachyris glutinosa Schauer, Linnaea 19: 724, 1847.
Gutierrezia sphaerocephala A. Gray, Pl. Fendl. 73, 1849; Xanthoce-
phalum sphaerocephalum (A. Gray) Shinners, Field & Lab. 18: 29,
1950. Based upon Fendler 527, as prairie, from the upper to the
middle Spring of the Cimarron, VIII.
Gutierrezia eriocarpa A. Gray, Pl. Wright. 1: hee 1852; Xanthoce-
phalum sphaerocephalum var. eriocarpum (A. G Shinners, Field
and Lab. 18: 29, 1950: Based upon Wright 280, oP raition along the
Rio Grande, Tex:
Gutierrezia Coulteri Hemsl. sens Pl. Nov. 33, 1879. Based upon
Gutierrezia Berlandieri A. Gray, Proc. Amer. Acad. Arts & Sci. 15:
= 1880. Based upon Berlandier 1298, “Near San Luis Potosi” [Mexi-
antes annual 2-60 em. high. Stem branching from the base or at
about 1/3 its height from the ground, gray or greenish-gray, 2-3 mm.
in diameter, branchlets slender, light green, fistulose. Root a short
taproot. Leaves lanceolate, 2-3 cm. long, 2-3 mm. wide, entire, surface
smooth to rugose, glabrous. Heads at the end of branchlets, not nu-
merous, sometimes less than 10, pedicellate. Involucre campanulate,
3-5 mm. high, 4-8 mm. wide, involucral bracts elliptical, arranged in
2 poorly defined series, acute, with membranaceous margins, tip and
mid-rib section slightly darker and thicker. Ligulate flowers 10-20,
corollas 3-4 mm. long; tubular flowers 20-40, corollas slightly cam-
panulate 2-3 mm. long. Pappus variable in length, less than 1/2 mm.
to as long as the achene, shorter in the ligulate flowers. Achenes about
1.5 mm. at anthesis, up to 2.5 mm. at maturity, turbinate, hairy, with
the hairs in rows.
TYPE-LOCALTY. “Crescit in terris mexicanis”, Aschenbach 2:
DISTRIBUTION. Central and northwestern Mexico to Sina
Texas and southern New Mexico (fig. 4).
REPRESENTA’ SPECIMENS. Mexico. Coahuila. 2 miles NW of
Frontera, 24-26-VIII-1938, Johnston 7182 (GH, Us). Chihuahua. Plains
near Chihuahua, 11-X-1885, Pringle 622 (wTU, F, NY). United States.
New Mexico. 20 miles S. of Roswell, 8-IX-1900, Earle & Earle 533
(Ny). Near Las Cruces, 11-VI-1897, Wooton 130 (Ny, GH, Mo). Tex-
as. Ft. Quitman levee road, 18-VI-1943, Waterfall 4596 (Mo, GH).
Chisos Mountains, 15-18-VII-1921, Ferris & Duncan 2748 (NY).
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62 OTTO T. SOLBRIG
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THE SOUTH AMERICAN SECTIONS OF ERIGERON
AND THEIR RELATION TO CELMISIA*
In his studies on the family Compositae, M. Henri Cassini
(1816-1830) laid the foundation for the modern treatment
of this large and complex group. Fifty years after Cassini
had begun his studies, George Bentham, using to a great
extent the criteria elaborated by Cassini, attempted a world-
wide study of generic | Jaries within Compositae (Ben-
tham, 1873; Bentham & Hooker, 1873). His work was so
complete and of such fine quality that it has remained large-
ly unchallenged up to our day, Hoffmann’s (1893) later treat-
ment for Engler’s Pflanzenfamilien not differing appreci-
ably.
Nevertheless, due to the discovery of many new and some-
times critical species and genera in the last hundred years,
our understanding of Compositae has increased considerably.
Also, with the completion of the main phase of plant explor-
ation in many temperate areas, our knowledge of plant dis-
tribution has been greatly widened since the publication of
Bentham and Hooker’s Genera Plantarum. Furthermore,
cytology, genetics, anatomy and embryology have added new
dimensions to taxonomy and to our understanding of evolu-
tionary processes. Geologists, although still disagreeing, can
furnish more concrete ideas today than in Bentham’s time
about such topics as continental drift and land bridges; and,
finally, our knowledge about climatic conditions in the Terti-
ary era, probably when most Compositae genera were dif-
ferentiated, has also been augmented to a large extent. In
the light of all these advances, it is not surprising that many
of the generic boundaries established by Bentham are in
need of revision.
The tribe Astereae is one especially suited for a careful
analysis of generic boundaries. More or less detailed studies
covering most or at least part of the tribe, have been com-
pleted on chromosome morphology (Huziwara, 1959), chrom-
osome numbers (Raven, Solbrig, Kyhos & Snow, 1960,
together with citations), embryology (Harling, 1951), and
wood anatomy (Carlquist, in preparation). Several genera
have been monographed recently and papers dealing with the
redefinition of certain genera have been published. Also, in
TTam grateful to Dr. Rolla M. Tryon for reading the manuseript and making valu-
able suggestions.
66 OTTO T. SOLBRIG
a number of regional floras the problem of generic limits has
been analyzed on a local basis. If a study of generic relation-
ships is to reflect natural affinities and the correct evoiution-
ary history of the tribe, undoubtedly it has to be world-wide
in scope and has to involve as many different approaches as
possible.
Erigeron is one of the larger genera of the tribe Astereae,
both in number of species and in area of distribution. Al-
though it is truly a genus of the northern hemisphere, with
species native to both the Old and the New World, and with
a probable center of origin in western North America, it is
also found in South America, mainly along the Cordillera de
los Andes. Erigeron is closely related to two other large
genera, Aster and Conyza. The problem of delimiting these
three genera has been dealt with on several occasions by
Bentham & Hooker (1873), Bentham (1873). and Cronquist
(1941, 1947), among others. Consequently, this problem is
not discussed at length here, and the criteria enunciated by
Cronquist for separating these three genera will be followed.
Bentham and Hooker (1873) divided Erigeron into six
sections, based mainly on characters of the inflorescence,
style, pappus, ligules and habit. Four of these sections,
Leptostelma, Erigeron ( Euerigeron), Coenotus and Oritro-
phium are represented in South America. Section Coenotus
was transferred to the genus Conyza by Cronquist (1941).
Section Oritrophium, as established by Bentham and Hook-
er, is composed of perennial herbs with monocephalous
scapes, female flowers always ligulate and in one series, and
the style linear and subulate. Habit in the typical species is
like that of the New Zealand and Australian genus Celmisia.
ion Leptostelma was also established by Bentham and
Hooker and is based on the genus Leptostelma D. Don. It
1s composed of large herbs characterized by rather large
heads and an alveolate receptacle. Section Erigeron (Eueri-
geron) is represented in South America by perennial herbs,
small subshrubs, and a few shrubs. This is the largest and
Most variable of the North (Cronquist, 1947) and South
American sections of the genus.
We see, therefore, that Erigeron is represented in South
America by three relatively well distinguished groups. One,
section Erigeron, composed of over 25 species, is closely
related to the North American species. It shows great vari-
SOUTH AMERICAN ERIGERON 67
ation but in general this variability conforms to that of the
genus as represented in the northern hemisphere. The other
two groups, although apparently closely related to true Eri-
geron, show some characters that are absent in section Eri-
geron.
The object of this paper is to present some detailed obser-
vations on the morphology of species hitherto classified as
belonging to sections Oritrophi d Leptostelma and to
reevaluate the taxonomic position of the South American
sections of Erigeron.
MATERIALS AND METHODS
All observations were made on herbarium material. In addition to
studies of dried and boiled material, fragments of both leaves and
heads were cleared with sodium hydroxide and stained according to
Foster’s technique, as explained elsewhere (Solbrig, 1960), and per-
manent mounts prepared.
Materials of the Arnold Arboretum (A) and Gray Herbarium (GH)
of Harvard University; Museo de Ciencias Naturales, La Plata, Ar-
gentina (LP); Museo Botanico, Cérdoba, Argentina (corp); and In-
stituto Miguel Lillo, Tucuman, Argentina (LIL) were available for
this study.*
OBSERVATIONS AND RESULTS
Studies were made of the general habit; shape, venation
and pubescence of leaves; secondary inflorescences; shape,
arrangement and pubescence of involucral bracts; corollas,
styles, achenes and pappus of both ligulate and tubular
flowers; anthers and pollen; and receptacle. In addition, a
few general habitat preferences are recorded.
GENERAL H.
ABIT
All species of South American Erigeron studied are per-
ennial. Erigeron maximus and E. Tweediei, of section Lep-
tostelma, are gigantic herbs, up to 4 m, high in the case of
E. maximus and 1 to 1.50 m. in the case of E. Tweediei. The
latter species has a basal rosette of large leaves, which is
absent in E. maximus. Both species have profuse foliage,
greenish stems, and heads arranged in loose cymose second-
ary inflorescences. The habit of species of section Leptostel-
ma is very much in accordance with the humid, tropical
environment in which the plants grow.
Members of the section Oritrophium on the other hand are
very different. They have a crowded basal rosette, with stiff,
more or less lanceolate leaves, and a monocephalous floral
— Tam grateful to the directors and curators of the institutions mentioned for
ing available the material in their custody.
68 OTTO T. SOLBRIG
scape not more than 30 cm. high. Three of the species of this
section, E. pellitus, E. crocifolius and E. hieracioides, are
covered with a thick indumentum. A great variety of habits
is found in section Erigeron. The most common type, which
is found along the Andes, is somewhat similar to section
Oritrophium, with a basal rosette of leaves and a monoceph-
alous or polycephalous leafy scape. The development of hairs
on the leaves and involucral bracts is variable. Other species
in this section are subshrubs, with a woody base, with or
without a basal rosette of leaves. In these species, the ar-
rangement of the capitula is usually secondary compound.
Finally, in the Juan Fernandez and Galapagos Islands some
truly shrubby species (E. fruticosus, E. tenuifolius, and E.
lancifolius) are found, ranging from about a meter (E.
fruticosus) to a meter and a half (E. tenuifolius) in height.
It may be noted that sections of Oritrophium and Leptos-
telma can be separated from section Erigeron on characters
of habit alone. In spite of this, their external appearance is
not radically different from what is found in section Eri-
geron but constitutes two extremes within the general varia-
tion found in South America Erigeron.
LEAVES
As already pointed out, there is a marked difference in the
leaves of species belonging to each of the three sections.
Those of section Oritrophium are stiff and coriaceous, linear
to lanceolate in shape, with entire margins, varying in length
but not longer than 25 em. in the largest basal leaves. The
cauline leaves are very reduced, usually only about half a
centimeter long. Leaves of species of section Erigeron are
30 cm. in length and 10 em. in width, variably lanceolate and
with a cut or serrate margin.
As to the surface of the leaves, in section Oritrophium a
almost glabrous upper surface is present and the lower
surface is covered with a thick mesh of interwoven hairs,
which gives ita silvery aspect. In some cases the upper sur-
face also is covered with hairs, This peculiar type of pubes-
cence is not found in any of the other sections but instead is
characteristic of the genus Celmisia. Leaves of species of
Sections Leptostelma and Erigeron have a dull surface, gla-
SOUTH AMERICAN ERIGERON 69
brous or pubescent. If they are pubescent they are usually
equally so on both surfaces but never to the extent found in
the lower leaf surfaces of species of section Oritrophium.
The venation of the leaves is reticulate, with the veinlets
free at their termination. No appreciable difference could be
found in any of the sections.
INFLORESCENCE!
Differences in the inflorescence can sometimes be impor-
tant in the taxonomy of the Compositae. In the present
instance, this character can be used to some extent in separ-
ating sections Oritrophium, Leptostelma and Erigeron.
While most Species of Erigeron in South America have a
usually cymoid or racemose,
members of the section Oritrophiwm have their capitula
borne at the end of sparsely leafy scapes. On the other hand,
some members of section Erigeron, such as E. andicola, also
have their heads borne solitary at the end of slender scapose
branches. The scapes of species of section Oritrophium,
such as E. pellitus, are much more robust and rigid, while
those of section Erigeron are relatively slender. An excep-
tion to this is E. hieracioides (section Oritrophium), which
has scapes approximating those of E. andicola.
The inflorescence of E. maximus and E. Tweediei can be
roughly characterized as an inverted compound cyme. After
the terminal head has started to form and the growth of the
branchlet has come to an end, one to three lateral branches
develop from the axillary buds nearest to the head on the
central axis. Eventually heads will develop at the ends of
these branchlets but usually not until the tips of the lateral
branchlets have grown past the end of the main one. This
process may take place then in each of the lateral branchlets
again, leading to double-compound inflorescence. The end
result is that the last formed heads are at the top of the in-
florescence, rather than at the lower part as in the cymose
inflorescences of members of section Erigeron. In E. maxi-
mus this inflorescence type is quite distinct while in E. Twee-
diei it is more compact and there is some —— between
this type and a more typical cymose inflorescen
If the pattern of inflorescence Seat 3 is s carefully
‘“*Througnout this paper, inflorescence refers to the arrangement of the capitula,
of the flowers.
which are the real inflorescences, rather than the arrangement
use of the word inflorescence follows common usage in the Compositae.
70 OTTO T. SOLBRIG
as O
th | {
\ \
/
iV
Soe of inflorescence
1 poten maximus. sag . Karwinskianus. 3. E. plladarstel 4. Celmisia pellita.
5. Erigeron pinnatus. 6. E. Poeppigii. 7. E. Gayanus. 8. E. andicola, 9. E. brevi-
caulis.
For further explanation see text.
observed, an evolutionary series can be detected. If a simple
cymose inflorescence, common in section Erigeron and such
SOUTH AMERICAN ERIGERON TL
as is found in E. Gayanus (fig. 7) is first observed, it may
be seen that one tendency is towards a reduction in the num-
ber of heads, correlated with a reduction in the size and num-
ber of the leaves of the scape. At the same time, the leaves
of the scape become more distinct from those of the basal
rosette (loss of petiole, shorter and narrower lamina, etc.),
as exemplified by E. andicola (fig. 8). An extreme in this
adaption is represented by E. rosulatus and E. pulvinatus
where even the scape has disappeared, the terminal heads
being borne singly at the ends of very short and leafy stems.
Whether E. pellitus (fig. 4) and the other species of section
Oritrophium constitute another specialization within this
trend is difficult to decide but this possibility seems rather
doubtful.
Another trend has been towards a type with less differ-
entiation between basal and cauline leaves (e.g., E. Poep-
pigii, fig. 6) together with the formation of a tighter
inflorescence. The suffruticose species, such as E. Karwin-
skianus (fig. 2), show a pattern essentially similar to E.
Poeppigii, although obscured somewhat by the woody type
of growth. The basal rosette has di
ante all the cauline leaves are well developed. The inflores-
cences are borne toward the ends of the branches and are of
the same cymose type as in E. Poeppigii. In some cases, as
in E. Berterianus (fig. 3), we find a more reduced inflores-
cence. Whether the type of inflorescence found in members
of section Leptostelma (fig. 1) is an elaboration of the one
found in the shrubby species is hard to determine with any
degree of certainty.
INVOLUCRAL BRACTS
In general, the involucral bracts are rather uniform with-
in the Astereae. They are distinct structures, herbaceous or
somewhat woody, usually imbricate in two or more rows.
Sometimes, as in the genus Grindelia, they might take a
more specialized aspect, but we do not find in Astereae large
spines as in Cirsium or Centaurea, or any such other special-
izations in the bracts as are common in other tribes,
Mutisieae or Cynaroideae. Small variations, therefore, be-
come of taxonomic value, such as the arrangement of the
bracts in the involucre, size, shape, pubescence, number of
bracts, and degree of imbrication.
Characters of the involucre are important and have been
OTTO T. SOLBRIG
quist, 1947), but a certain degree of variation in involucral
used extensively to separate Aster from Erigeron (cf. Cron-
72
QS = SS SRY Ar)
bap Wr PAL ee)
PM WASIINDIS IM
, 1920, GH). isic
Anderson 229, GH). 15. Erigeron Tweediei (Osten
SSS SLL
;
'
LLL ZZ
cock 21997, cH). 14. C. gracilenta (
1475 0, GH).
SOUTH AMERICAN ERIGERON 73
characters is present from species to species within a genus
and in some cases within a particular species, as is common
in all biological material.
The species of Erigeron have herbaceous bracts which can
be non-herb at the base. They are of dif-
ferent sizes, with the outer ones shorter, loosely imbricate,
and usually in two loose series. Pubescence varies greatly.
Some species are glabrous and others are covered with a
thick cap of hairs. The inner surface of the bracts is always
glabrous. As to the shape, there is no appreciable difference,
all species having loosely lanceolate bracts.
The involucral bracts of the South American species of
Erigeron do not show any noticeable departure from this
pattern, with the exception of section Oritrophium. In this
group, the bracts are somewhat larger and less herbaceous
and appear to be intermediate between typical Erigeron and
Aster (fig. 13).
Pubescence of the involucral bracts is another character
that has been used widely in Erigeron, especially in the sep-
aration of species (Cronquist, 1947). In the material stud-
ied, we found a great deal of variation in the amount of
pubescence of the involucral bracts (fig. 10-15) which vary
from completely glabrous to thickly pubescent. In addition,
there is a qualitative character-difference in the trichomes
of section Oritrophium and those of sections Erigeron and
Leptostelma. The trichomes of the latter two sections are
multicellular and uniseriate, as are those of section Oritro-
phium, but the trichomes in the involucral bracts of section
Oritrophium are formed by a few elongated cells (sometimes
branched), all of about the same width and length and taper-
ing gradually towards the tip of the hair, whereas the tri-
chomes of the involucral bracts of species of the other
sections are formed by a few (3 to 6) short and wide cells
at the base, followed by a series of cells several times longer
always a very clear difference. In addition to the types of
are absent in species of Oritrophium.
74 OTTO T. SOLBRIG
FLOWERS
It is well known that species of Erigeron have two types
of flowers in the capitulum: the ray flowers, which are pistil-
late and have a ligulate corolla; and the disk flowers, which
are hermaphroditic and tubular. Flowers furnish some
important characters helpful in the delimitation of species.
In the present case, some substantial differences in flower-
characters exist between section Oritrophium and the other
South American species of Erigeron. Therefore, the pappus,
corolla, anther, pollen, style, achene, venation of the flowers,
and receptacle will be analyzed in some detail.
PAPPUS. The pappus is formed by a large number of
bristles about the same length as the tubular corolla and
arranged in one series. This holds true for all three sections.
COROLLA. The only species-difference present in the ligu-
late corollas is the length of the ligule, which varies from
about as long as the tube to about five times longer. Species
of section Oritrophium have rather long ligules but it is in
Erigeron maximus and E. Tweediei of section Leptostelma
that the longest ligules in South American Erigeron are
found. The width of the ligule varies somewhat but no pat-
tern can be discerned, the variation being between the
expected parameters for the genus. A similar uniform sit-
uation holds true in the tubular corollas, where the only dif-
ference is the presence of microscopic, uniseriate and
multicellular hairs on the outer surface of the tubular corol-
las of species of section Oritrophium.
ANTHERS. The anthers are uniform and are of a typical
asteraceous type with lanceolate upper appendages and no
lower appendages. An exception is Erigeron pellitus of sec-
tion Oritrophium, which has some very short appendages at
the base, usually called “tails”. Presence of “tails” on the
anthers is the character used to separate the tribe Inuleae
from the Astereae but it is also found in at least one genus
of Astereae, the genus Celmisia.
_ POLLEN. Pollen of E. andicola (section Erigeron), E. max-
tmus (section Leptostelma) and E. ifolius (section Ori-
trophium) was investigated. No qualitative character of
importance could be detected, aside from slightly larger
spines on grains of E. ifolius. Mi based on
samples of 25 grains are shown below:
SOUTH AMERICAN ERIGERON 75
Po>
PLP PIO
SES
2797279223 3
DOLL ZELLEEE,
ai DEEZ ZZ ZEEE Zs
ODI. IIS Lod
(Bang 913, GH). 17-
Mexia 4341,
it rigeron andicola (E. W. D.
and Mary Holway, 1920, GH). 21. Celmisia gracilenta (Anderson 229, GH).
76 OTTO T. SOLBRIG
22.26. Achenes of flowers. 22. Celmisia hieracioides (Pennell 12875,
GH). 23. C. pellita (Hitchcock 21997, GH). 24. Erigeron andicola (E. W. D. and
Mary Holway, 1920, GH). 25. E. rosulatus (Bang 913, GH). Fig. 26. Venation of
rigeron 1H).
SOUTH AMERICAN ERIGERON i §
Species Minimum Maximum Mean
E. andicola 25 29 26.84
E. crocifolius 28 33 30.32
E. maximus 24 31 27.32
Expressed in micrometer units for comparative purposes only
STYLES. The style has been, since Cassini, the most im-
portant single character used in the taxonomy of Compositae.
Each tribe and many genera are supposed to have distinct
and peculiar stylar appendages, so that the tribe and some-
times the genus to which a plant belongs can readily be de-
termined by examining the style. Actually it is not as simple
as this because a great deal of variation exists. A careful
study of variation in stylar characters in all genera of Com-
positae is needed, but in order to be of value, any study of
stylar characters must take into account the cellular details
as well as the overall shape of the style, as has already been
pointed out by Carlquist (1959).
The styles of the ligulate corollas are very similar in all
the species under consideration and in most species of As-
tereae. They consist of two elongate, flattened branches,
slightly pointed at the tip, with a border of stigmatic papillae
all around the margin of the inner surface of the stigmatic
branch.
The styles of the tubular flowers are of two types. In the
first, in section Oritrophium, the stigmatic branches are nar-
row, elongate, and gradually attenuate towards the tip. Col-
lecting hairs are more or less evenly scattered along the
outer surface. The collecting hairs are formed by long nar-
row cells of even width and are rounded at the tip. The stig-
matic papillae are small cells, usually pointed, which form a
narrow strip along the inside border of the stigmatic branch-
es (fig. 17). This type of style does not belong to the so
called “Erigeron type’, but is of a kind which is found in
some Pacific Basin genera of Astereae, such as Tetramolopi-
um, Vittadinia, Podocoma, etc.
The type of tubular flower style found in sections Eriger-
on and Leptostelma is as expected in the genus Erigeron.
The stigmatic branches are broader than in styles of species
of section Oritrophium and are of even width, te
in a “triangular” tip, characteristic of Erigeron. The col-
78 OTTO T. SOLBRIG
lecting hairs are appressed over the outer surface of this
“triangular” tip and are short and somewhat rounded (figs.
18 & 20). The stigmatic papillae form two wide borders
along both sides of the two stigmatic branches from the point
of bifurcation to the start of the area of collecting hairs.
Some variation can be observed in the downward extension
of the collecting hairs but in general the styles are quite
uniform. An extreme form of variation is found in Erigeron
pulvinatus and E. rosulatus, where the collecting hairs ex-
tend down along the whole outer surface of the stigmatic
branches and consequently restrict the papillae to the inner
surface of the stylar branches (fig. 16). The type and den-
sity of the collecting hairs, as well as the shape of the stig-
matic branches, indicate that E. pulvinatus and E. rosulatus
are correctly classified in section Erigeron.
ACHENE. A difference exists between section Oritrophium
and sections Erigeron and Leptostelma in the shape of the
achene. Achenes of species of section Oritrophium are elong-
ate and somewhat fusiform, tapering at both ends but partic-
ularly at the lower one (figs. 22 & 23). In contrast, the
species of sections Erigeron and Leptostelma have shorter
and more cylindrical achenes. All the species investigated
had somewhat flattened, two-ribbed achenes, flatter in spe-
cies of Oritrophium than in those belonging to other sections.
As far as pubescence of the achenes is concerned, we find
a great amount of variation, ranging from species with glab-
rous achenes to species with densely pubescent ones. The
trichomes are of three different types: (1) short, uniseriate,
multicellular, formed by a few isodiametric cells; (2) long,
uniseriate hairs with elongated cells; and (3) biseriate, mul-
ticellular hairs. In some cases hairs are found which are
biseriate at the base, turning uniseriate toward the apex.
as characteristic pattern can be discerned in any of the sec-
ons.
Another aspect to consider is the fertility of the achenes.
In sections Erigeron and Leptostelma, the achenes of both
the ligulate and tubular flowers are fertile, although shrunk-
en and unfertilized achenes are found occasionally in both
the ligulate and tubular flowers. In species of Oritrophium,
the achenes of the tubular flowers are usually very shrunken
and inviable while those of the ligulate flowers are fertile.
Sometimes one can find tubular achenes which are full and
SOUTH AMERICAN ERIGERON 79
apparently normal. In view of the great reduction of the
stigmatic papillae in section Oritrophium, it can be specu-
lated that only occasionally do pollen grains germinate and
fertilize the ovules to produce a fertile achene. This would
mean that section Oritrophium represents an intermediate
stage between hermaphroditism and unisexuality in the tubu-
lar flowers, but it is also possible that the achenes are always
sterile and that t of the achene might
be due to causes other than fertilization.
FLORAL VENATION. No appreciable difference exists in the
venation of the tubular flowers of the species under study.
The ray flowers, on the other hand, show a great amount of
variability, even among flowers of the same head. Therefore,
only the tubular flowers will be described.
There are usually two bundles in the achene (in a few
cases three were observed) which pass into the corolla where
they give rise to five corolla bundles, with adjacent pairs
uniting at the tips of the corolla lobes, and to two stylar
traces (fig. 26). In addition, a third trace can be seen in the
achene, the ovular trace. No ovular trace can be detected in
species of section Oritrophium, which enhances the suspicion
that the achenes in this section might always be sterile.
RECEPTACLE. Section Leptostelma is supposed to have a
“paleaceous” receptacle, while the other species of Erigeron
are supposed to possess a naked receptacle. Close inspection
shows that this difference is more apparent than real. All
South American species of Erigeron have an alveolate recep-
tacle, with the borders of the depressions extended into a
thin microscopic membrane. In E. maximus and E. Twee-
diei this membrane is less than half a millimeter larger and
cannot properly be called a ‘‘palea”. All species investigated
had a flat or slightly convex receptacle.
In conclusion, it is interesting to point out that many small
differences exist in characteristics of the capitulum and flow-
ers between Oritrophium and sections Erigeron and Leptos-
telma. Only in characters widespread in the tribe, such as the
pappus and the corolla, is agreement found. In view of the
uniformity and stability of flower characters i in Compositae,
these diff. a idered to be si;
GEOGRAPHIC DISTRIBUTION
Species of section Oritrophium grow in dry, almost bar-
ren high altitude plateaus in the northern Cordillera de los
80 OTTO T. SOLBRIG
Andes, locally known as the “paramos”. Species of Oritro-
phium are found at altitudes of 3,000 to 4,000 m. in north-
western Bolivia, Peri, Ecuador, Colombia and western
Venezuela. In Colombia is found the greatest development
B
<
ae ‘
= ,
Ny ee yh S ¢ »
ieeey
Yrt--
Oniraopmum. oN ee Woe
a
CO
ee
ary
\
ss
h\ re
%
\ “oa
axe LePToSsTELMA
, Qe
~*
\\\ ; o%
M =a Wes
Ae 23 2
7 Z
<5 Ss.
fl t x a te i)
<<
f
i ait,
“-G
Lex
seg ‘ @a
WW section Eniceron
O Cexwisia REPENS
@ Cems Hieraciowes
® Cecmisia crocciroua
x CeLMsia PELLITUS
QeEniceron Tweens
@ERceron waximus
Fic. 27. Geographical distribution of the South American seetions of Celmisia and
SOUTH AMERICAN ERIGERON 81
in number of species and also the largest number of herbari-
um collections comes from that area. All species of section
Oritrophium are typical high altitude plants and grow in the
di ypes of tal niches. On the con-
trary, ‘members of section Leptostelma are found in wet and
marshy localities in southern and southeastern Brazil (states
of Minas Gerais, Sao Paulo, Parana and Rio Grande do Sul),
northeastern Argentina (state of Misiones), Paraguay, in
the lowlands of eastern Bolivia (Santa Cruz de la Sierra)
and in the Departamento de San José in southern Uruguay.
The plants are found at altitudes below 1,000 m.
The areas where members of sections Oritrophium and
Leptostelma grow, the high Andes of northwestern South
America and subtropical forests of eastern South America,
have not been explored exhaustively and it is very likely that
the range of species can be determined with greater accuracy
than is depicted on the map of fig. 27 when botanical knowl-
edge of those areas is more complete.
Section Erigeron is found in South America along the
Andes frem Ecuador to Tierra del Fuego, at altitudes rang-
ing from sea level to about 5,000 m. Species belonging to
this section are also found in the Sierras Pampeanas of
Argentina, which are older geologically than the Andes, and
also along the coast of Chile and southern Pert, extending
northward to the area around Arequipa. Finally, a few spe-
cies are found in the Juan Fernandez and Galapagos Islands.
In all of this region, species of section Erigeron are found in
rather dry, rocky habitats, even though they sometimes spill
over into more mesic environments.
Sections Erigeron and Oritrophium overlap somewhat in
their ranges and ecological preferences in Bolivia, Perti and
Ecuador, while section Leptostelma seems to be separated
both geographically and ecologically. Cabrera (1957) des-
cribed two species of Erigeron from eastern Brazil of which
I have not seen any specimens. The description indicates
that the plants are similar to the North American Erigeron
annuus of section Phalacroloma and unlike any Brazilian
species of Erigeron. The two Brazilian species described by
Cabrera differ from Erigeron annuus in pappus details and
have to be placed in section Oligotrichium (Cronquist, 1947).
Neither section is represented in South America. It is always
possible that a mutant of EZ. annuus, of relatively recent
82 OTTO T. SOLBRIG
introduction, is the plant in question. Erigeron annuus,
which has spread as a weed in both the United States and
Europe, is a triploid (2n = 27) apomict. Some North Amer-
ican weedy Compositae, as for example Heterotheca latifolia,
have been introduced into similar areas of Brazil (Wagen-
Knecht, 1960).
DISSCUSSION
SECTION ORITROPHIUM. Morphologically, section Oritro-
phium is similar to Celmisia, a characteristic genus of the
flora of New Zealand, found also in Tasmania and Australia.
The similarities between Celmisia and section Oritrophium
have been pointed out by Bentham, 1873; Bentham & Hook-
er, 1873; Gray, 1884; and Cronquist, 1948. Bentham (1873)
comments as follows: “Around Erigeron may be grouped
the following slightly divergent genera, designated as much
by their geographical areas as by any structural characters,
and yet natural enough to be readily recognized: 1. The
Antarctic and Australian genera Pleurophyllum and Cel-
misia, closely connected with the Andine section Oritrophium
of Erigeron, differ from it chiefly in the shape of the achene,
which is more that of Olearia . . .”. Gray (1884) says“. . .
section Oritrophium (which must belong either to Celmisia
or Aster) . . .”. Hoffmann (1893) separates Celmisia from
Erigeron (including section Oritrophium) on account of the
tailed
rs.
TABLE 1. COMPARISON OF DIFFERENCES IN MORPHOLOGICAL CHARACTERS
IN THE GROUPS UNDER STUDY
Character Celmisia Celmisia Erigeron Erigeron
sect. Celmisia sect. Oritro- seet. Erigeron sect. Leptostelma
phium
General habit — Seapose with Scapose with Seapose with Perennial herbs
basal rosette basal rosette, up to 4m high
suffrutex or it
rosette
Monocephalous Monocephalovs Mono- or Polycephalous
Polycephalous
Very thickly Very thickly Pubescent to Thinly pubescent
pubescent pubescent. glabrous
Subulate Subulate Triangular Triangular
Elongated elongated prismatie prismatie
with oceasionally never with never with
“tails” ‘tails’
SOUTH AMERICAN ERIGERON 83
As may be seen in table 1, neither the achenes nor the
anthers of section Oritrophium differ from those of Celmisia.
It is therefore proposed that section Oritrophium of Eriger-
on be transferred to the genus Celmisia.
SECTION LEPTOSTELMA. Species of section Leptostelma are
easily separated from other species of Erigeron on the basis
of vegetative characters. The floral characteristics do not
differ appreciably. Consequently it is difficult to decide
whether Leptostelma merits generic rank.
A chromosome count for E. maximus has recently been
reported (Turner & Irwin, 1960). Unfortunately, the con-
dition of the material did not permit an accurate determina-
tion and it was reported as 40 + 4. The basic number for
Erigeron is x = 9 (Raven, Solbrig, Kyhos and Snow, 1960;
Montgomery and Yang, 1960). Polyploidy is known in North
American species but, with the exception of the apomict E.
annuus (n = 13; 2n = 26, 27), all counts reported to date
are multiples of 9 (Montgomery and Yang, 1960). No counts
other than Turner’s exist for South American species. There-
fore, if the chromosome number of E. mazinvws is not 36, it
might be an indication of a different generic stock.
Nevertheless, for the present, it is felt that, although on
the basis of vegetative and distributional characters Leptos-
telma might be set up as an independent genus, it is better
to wait until new information, especially cytological data, is
available, before taking positive action.
PHYTOGEOGRAPHY. The inclusion of Oritrophium in Cel-
misia raises an interesting phytogeographical problem. Cel-
misia is an important element of the flora of both the North
and South Islands of New Zealand, being one of the three
largest genera and the largest genus of Compositae. The
Compositae is the dominant family in New Zealand (Coc-
kayne, 1928). That genera and even species are common
to New Zealand and southern South America is a well known
fact, but floristic connections between New Zealand and the
high Andean areas of northwestern South America are not
frequent. In this case, one could probably explain the distri-
bution of Celmisia, section Oritrophium, by postulating a
migration across Antarctica and along the Cordillera de los
Andes, with an extinction of the genus in the areas where it
is now absent. The asteraceous genus Lagenophora (sub-
tribe Bellidineae) might be cited as providing a clue in the
84 OTTO T. SOLBRIG
direction of the postulated explanation. Lagenophora is
found in Australia, New Zealand, New Guinea, Hawaii and
South America. In the latter, it has a disjunct distribution,
with one species in the area of the subantarctic rain forest
and a second species in the high mountains of Venezuela
(Cabrera, 1954). Also, the fern family Loxsomaceae might
be cited. This small family has only two genera, Loxsoma,
with one species endemic in New Zealand, and Loxsomopsis,
with three species, found along the Andes from Bolivia to
Costa Rica (Copeland, 1947). As is evident, it covers largely
the same territory as Celmisia.
The inclusion of section Oritrophium in Celmisia clarifies,
to some extent, a picture of the relationships of Erigeron in
the southern hemisphere. Erigeron is undoubtedly a genus
of northern affinities, with a probable origin in western
North America. It has expanded into the Old World and
secondary centers of speciation can be postulated for Asia
and Europe. It has also migrated into the southern hemis-
phere. There are two possible ways of explaining this latter
migration. One, by postulating a migration along the Cor-
dillera de los Andes ; the second by long-range seed dispersal.
Either hypothesis cannot be categorically proved or dis-
proved and more data needs to be accumulated, pertaining
to such things as breeding systems, before we can weigh
both hypotheses in the light of our knowledge of Tertiary
climate and conditions.
Erigeron is but one of a group of genera such as Grindelia,
Haplopappus and Solidago in South America, which have
migrated from western North America. Another group of
South American Astereae, including Vittadinia, Lageno-
phora, Podocoma and Celmisia, is of Australasian origin.
Finally, some genera, such as Conyza, seem to be pantropical
in distribution. A better understanding of the origin of this
distribution will aid greatly our knowledge of the phylo-
genetic relationships within the tribe Astereae.
SUMMARY
Erigeron L. section Erigeron.
Approximately 25 to 30 species in South America.
specimens: Erigeron andicola DC., Holway, 1920 (GH);
E. brevicaulis Phil., Johnston 5960 (GH); E. cinerascens Sch. Bip.,
Venturi 8633 (GH, LIL) ; E. fruticosus DC., Bock 54 (cH); E. Gayanus
Remy, and Solander s.n. (GH) ; E. Karwinskianus DC., Montero
2020 (GH); E. Philippii Sch. Bip., Johnston 5911 (GH); E. pulvinatus
SOUTH AMERICAN ERIGERON 85
Wedd., Bang 913 (GH); Shepard 22 (GH); E. tenuifolius Hook. f.,
Stewart 733 (GH).
Erigeron L. section Leptostelma (D. Don) Benth. & Hook., Gen. Pl.
2: 280, 1873; Leptostelma D. Don ex Sweet, Brit. Fl. Gard., Ser. 2, t.
38.
Two species in eastern South America: E. maximus Link et Otto ex
DC., Prodr. 5: 284, 1836: E. Tweediei Hook. et Arn., Compan. Bot.
Mag. 2: 50, 1836.
Selected specimens: E. maximus Link et Otto, Mexia 4341 (GH);
E. Tweediei Hook et Arn., Osten 1475 b (GH).
Celmisia Cass. section Celmisia.
Some 50 species in New Zealand, Tasmania and Australia.
Selected specimens: Celmisia bellidioides Hook. f., Hector s.n. (GH) ;
C. discolor Hook. f., Cheeseman sn. (GH); C. glandulosa Hook. f.,
Cheeseman s.n. (GH); C. ———s Hook. f., Hunnewell 13,781 (GH);
C. sessiliflora Hook. f., Hector s.n. (GH); C. Sinelairii Hook. f., Kirk
sn, (GH); C. ana Hook. f., Hunnewell 13,780 (GH).
Celmisia Cass. section Oritrophium (HBK.) comb. nov.; Aster section
Oritrophium HBK. (pro parte), Nov. Gen. et Sp. 4: 89, 1820; Erigeron
sect. Oritrophium (HBK). Benth & Hook., Gen. Pl. 2: 280, 1873.
Four species in northwestern South America.
Celmisia crocifolia (HBK.) Sch. Bip. Bonplandia 4: 50, 1856; Aster
crocifolius HBK., Nov. Gen. et Sp. 4: 89, 1820; Erigeron crocifolius
(HBK.) Weddell, Chl. And. 1: 191, 1857.
Celmisia hieracioides (Wedd.) comb. nov.; Erigeron hieracioides
Weddell, Chl. And. 1: 194, 1857.
Celmisia pellita (HBK.) Sch. Bip. Bonplandia 4: 50, 1856; Aster
pellitus HBK., Nov. Gen. et Sp. 4: 91, 1820; Erigeron pellitus (HBK.)
Weddell, Chl. And. 1: 190, 1857.
Celmisia repens (HBK.) Sch. Bip. Bonplandia 4: 50, 1856; Aster
repens HBK., Nov. Gen. et Sp. 4: 90, 1820; Erigeron repens (HBK.)
Weddell, Chl. And. 1: 191, 1857.
specimens: Celmisia erocifolia (HBK.) Sch. Bip. Hitchcock
21656 (GH); C. hieracioides (Wedd.) Solbrig, Pennell 13875 (GH);
C. pellita (HBK.) Sch. Bip. Hitchcock 21997 (GH); C. repens (HBK.)
Sch. Bip. Weberbauer 6130 (GH) ; Pennell & Hazen 9853 (GH).
LITERATURE CITED
BENTHAM, G. 1873. Notes on the Classification, History and Geo-
graphic Distribution of Compositae. Jour. Linn. Soc. Bot. 13:
335-577.
. and J. D. Hooker, 1873. Genera Plantarum, Vol. 2(1),
1279 pp., London.
CABRERA, ae L. 1954. Origen y Evolucién de la Flora del Parque
N. al de Nahuel Huapi. Natura (Argentina) 1: 43-58.
. 1957. Compositae Brasilienses Novae. Arquivos do
Jardim Botanico (Rio de Janeiro) 15: 71-76.
CaRLQuist, S. 1959. Studies on Madinae: Rawue: Cytology, and
Aliso 4: 171-236.
86 OTTO T. SOLBRIG
Cassini, H. 1816-1830. Dictionnaire des Sciences Naturelles. 70 vols.
roel
‘aris.
Cockayne, L. 1928. The Vegetation of New Zealand. 456 pp. Leipzig.
CopeLanp, E. B. 1947. Genera Filicum. 247 pp. Waltham, Mass.
Cronquist, A. 1943. The Separation of Erigeron from Conyza. Bull.
Torr. Bot. cong 70: 629-632.
——————.. 1947. Revision of the North American Species of Erig-
eron, North of Mexico. Brittonia 6: 121-302.
Gray, A. 1884, Synoptical Flora of North America. Vol. 1 (2), 480
pp. New York.
HarLinG, G. 1951. Embryological Studies in the Compositae. Part
Ill. Astereae. Acta Horti Bergiani 16: 73-120.
HOSE MANS ae nas Compositae. In Engler, A. und K. Prantl, Die
4(5): 4,
Del someay <5 1959. Chromosomal Evolution in the Subtribe Aster-
inae. Evol. 13: 188-193.
Montcomery, F. H. anp S. YANG. 1960. Cytological Studies in the
Genus Erigeron. Canad. Jour. Bot. 38: 381-386.
Raven, P. H., 0. T. Sotpric, D. W. KyHos AND R. SNow. 1960. Chro-
mosome Numbers in Compositae. I. Astereae, Amer. Jour. Bot
47: 124-132.
Sonpric, O. T. 1960. Leaf Venation and Pubescence in the Genus
a (Compositae). Jour. Arnold Arb. 41: 259-269.
- L. anp H. S. IRwIN. 1960. Chromosome Numbers in the
Compas Il. Meiotie Counts for Fourteen Species of Brazilian
itae. Pcgnipaeriolgy 122-126.
acces B. L. 1960. Revision of Heterotheca, section Hetero-
theca (Camposiias): Riadora 62: 61-76, 97-107.
‘CONTRIBUTIONS FROM THE GRAY HERBARIUM _
OF HARVARD UNIVERSITY _ ae
: Studies in the Guttiferae. I. :
A Synopsis of Hypericum Section Myriandra ~
By : oe
= Taxonomic Fern Notes. II. =
CONTRIBUTIONS FROM THE GRAY HERBARIUM
OF HARVARD UNIVERSITY
Edited by
Reed C. Rollins and Robert C. Foster
NO. CLXXXIX
Studies in the Guttiferae. I.
A Synopsis of Hypericum Section Myriandra
By
PRESTON ADAMS
Taxonomic Fern Notes. II.
Pityrogramma (including Trismeria) and Anogramma
By
RoLLa TRYON
Published by
THE GRAY HERBARIUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASS., U.S. A.
Issued May 14, 1962
=
oes
eee
ct ggm ee
Se
oe
STUDIES IN THE GUTTIFERAE. I
A SYNOPSIS OF HYPERICUM SECTION MYRIANDRA'
PRESTON ADAMS?
Hypericum L. is an easily recognized cosmopolitan genus
with more than three hundred species classified among
eighteen sections (Keller, 1925). One of these sections,
Sect. Myriandra (Spach) Endl. (as delimited in the present
work), is restricted to eastern North America. The thirty
species of this group have never been studied as a sectional
unit. Coulter (1886, 1897), in his treatments of the family,
recognized many of the species involved but assigned them
to the Linnaean genera Ascyrum and Hypericum. Subse-
quent studies have been regional and floristic (e.g. Fernald)
1950; Gleason, 1952; Small, 1933) or else they were con-
cerned with special groups (Svenson, 1940; Adams, 1957).
Nomenclatural notes on several species were contributed by
Lott (1938)
new approach to the systematics of Hypericum was in-
itiated by Robson (1956) who surveyed the patterns of floral
vascularization present in Hypericum and related genera.
Information from his anatomical studies aided him in under-
standing the taxonomy of Hypericum in tropical and south-
ern Africa (Robson, 1957). His work stimulated my
interest in pursuing studies in the systematics of Hypericum,
especially on the group of species here included within Sect.
Myriandra. Through the kindness of Dr. Robson (now at
Kew) and the Edinburgh University Library, I obtained a
microfilm of his vascularization study which had been pre-
sented to that institution as a doctoral dissertation and is
not yet published. Data from his manuscript supplemented
by my morphological and cytological studies (Adams, pate
furnished the basis for a re- an
Ascyrum L. and Oe Small to the larger genus peste
cum (Adams & Robson, 1
Information derived ue anatomy, morphology, eve
gy, and phytogeography apt been used to construct this
synopsis of the syst Sect. Myriandra.
}Financial support of this stad was provided by the following sources: the —
Fund for Field Study in Systema‘ sou egret! at Harvard University, established
Mr. F. W. Hunnewell of Welles! a National Institute of ee grant-in-aid
through the Department of Biology, “eta University; and a research grant
(RG-6305) to Dr. R. K. Godfrey of Florida State University cies the Division of
2Present address: Department of Botany and Bacteriology, DePauw University,
G , Indiana. :
4 PRESTON ADAMS
A key to the species is accompanied by brief synonymy,
citations of type and representative specimens, habitat in-
formation, statements of geographical ranges, and comments
on interspecific relationships. Much more study of the group
remains to be done although the main outlines of the taxono-
my are clear. Detailed investigations of variation, both
within and among species, and the anatomy of the leaves,
sepals, and bark are in progress.
ACKNOWLEDGEMENTS
I am indebted to the following persons for their many helpful
criticisms and suggestions: Reed C. Rollins, R. K. Godfrey, Robert C.
Foster, Rolla Tryon, C. E. Wood, C. E. Kobuski, Lloyd H. Shinners,
George Argus, Arthur S. Barclay, K. A. Wilson, and J. L. Thomas. The
Latin descriptions were prepared by Richard Evans Schultes. Special
collections of specimens were provided by Delzie Demaree, R. B. Chan-
nell, H. F. L. Rock, Wilbur H. Duncan, R. K. Godfrey, R. Kral, Charles
B. Heiser, and Charles Schweinfurth. Mrs. Lazella Schwarten was
most helpful in locating and obtaining bibliographic items. Laboratory
space and equipment were made available by Jonathan J. Westfall at
the University of Georgia during the summer of 1958.
Much helpful information was contributed by Norman K. B. Robson
who has done intensive study of Hypericum. The loan of his unpub-
lished manuscript from the Library of Edinburgh University is much
appreciated.
pecimens from the following herbaria have been used in this study
(abbreviations according to Lanjouw & Stafieu, 1959) : Arnold Arbore-
(DUKE) ; Chicago Natural History Museum (Fr); University of Florida
(FLAS) ; Florida State University (Fsv) ; University of Georgia (GA);
vay Herbarium (GH); State University of Iowa (1A); Indiana Uni-
versity (ax) 5 Louisiana State University (Lsv) ; —— of Michi-
ical Club (NEBC) ; Tulane University (No) ; New York Botanical
Garden (Ny); University of Oklahoma (oKL); Oklahoma State
University (OKLA) ; Ohio State University (os); University of Penn-
PENN) ; Academy of Natural Sciences (PH); Purdue Univer-
sity (PUR); Southern Methodist University (smu); U.S. National
Herbarium (us); University of Tennessee (TENN); University of
Texas (TEx); University of Arkansas (UARK); Virginia Polytechnic
_ Institute (vpr); ee of Wisconsin (wis); and West Virginia
University (wv:
_ ‘THE COLLECTING oF HYPERICUM
"The ultimate writing of a definitive monograph on Hyperi-
STUDIES IN THE GUTTIFERAE. I. 5
cum Sect. Myriandra will be greatly aided if collectors will
pay close attention to the following suggestions. Probably
the most important is that samples of the stem and bark
(of the woody species) be included with the usual herbarium
specimens. Bark features are of paramount importance in
showing definitive characteristics of several species, especial-
ly the H. fasciculatum complex. Fruiting specimens are not
available for H. densiflorum from the Coastal Plain of the
Carolinas, apparently having been overlooked by collectors
even though the species has been known from that region
for at least one hundred years. Specimens of H. apocyni-
folium and H. sphaerocarpum in mature fruit are needed.
Carefully prepared plants (with accompanying rhizomes)
of H. ellipticwm would be most useful; the majority of the
herbarium specimens I have seen appear to have been yanked
from the soil! Observations (with specimens) of the winter
and early apie’ aspect of H. ellipticum, H. sphaerocarpum,
and H. dolab in are greatly needed. Collections of the
Hypericum spec es growing in Alabama and Mississippi
would be most ie So: in compilation of accurate distribution
maps. Specimens and observations on sympatric occurrences
of the following pairs of species would be of utmost value.
H. frondosum and H. prolificum, H. lobocarpum and H. pro-
lificum, H. lobocarpum and H. densiflorum, H. ellipticum and
H. adpressum, and H. stragulum ( = Ascyrum multicaule)
and H. hypericoides.
CHROMOSOME NUMBERS
Meiotic chromosome counts are available for | twenty-four
of the thirty species in 1 Sect. J (Nielsen,
1924; Hoar & Haertl, 1932; Adams, 1959a; H. L. Striplin,
rsonal communication). ‘All species studied possess a hap-
loid number of nine. Haploid chromosome numbers of eight,
twelve, and sixteen are known in other Hypericum species
native in eastern North America (but not in members of
Sect. Myriandra). The number nine is not confined to se
section, Psi present in several extra-North America
species also.
HYPERICUM L., Gen. Pl. ed, 5, 341. 1754
Sect. myRIaNDRA (Spach) Endl., Gen. Pl. 1033. 1840. Based on
Myriandra Spach, Hist. Nat. Vég. 5:434. 1836, as a genus.
Shrubs and perennial herbs. Leaves with or without an
at base; lamina expanded, or much reduced and the leaves eeatiaike:
translucent secretory glands present as dots or elongate sacs, black
ones absent. Trichomes absent. Sepals 5 or 4 (rarely 3 or 6), persis-
tent long after fruit maturity, or deciduous at capsule le dehiscence, with
6 PRESTON ADAMS
or without an articulation at base. Petals 5 or 4 (rarely 3 or 6), yellow,
deciduous soon after anthesis. Androecium of numerous stamens
arranged in a ring or narrow band (i.e. not fasciculate), deciduous or
persistent. Gynoecium of 5, 4, 3, or 2 fused carpels; placentation
pseudo-axile to parietal; styles same number as carpels, closely ap-
pressed to each other their entire length at anthesis, afterwards often
separating slightly; stigmas minute; ovules numerous to few. Seed
coats finely to coarsely reticulate, occasionally obscurely striated, raphe
inconspicuous, or forming a keel.
TYPE SPECIES (of section Myriandra) : H. prolificum L.
The circumscription of Endlicher’s Sect. Myriandra is
enlarged here in accordance with information yielded by
recent investigations. Robson (1956), from intensive studies
of the floral vascularization, concluded that the as
formerly classified within the genera Ascyrum L., Croo.
Small, and the sections Myriandra (Spach) Endl. "and Be
thydium (Spach) Endl. of the genus f Hypericum L are clos
ly related. This idea was by
morphological, geographical, and cytologi tigati
(Adams, 1959a). Recently Adams & Robson (1961) | trans-
ferred to Hypericum the species formerly placed in the
genera Ascyrum and Crookea.
changes involving Endlicher’s sections are proposed here.
Within Hypericum Sect. Myriandra there are two groups
of species which are distinguished by the presence or absence
of an articulation at the base of the leaves and sepals and, to
a lesser extent, we the degree of persistence of the withered
stamens after anthesis. The rank of subsection is used for
each of these groupings. The species of each subsection are
closely allied although several evolutionary lines are discerni-
ble within each.
The relationships of Sect. Myriandra are with Sect. Web-
bie (Canary Islands and Madeira) and Sect. Campylosporus
— saucy a and tropical Africa). Studies of floral
1956) and
(abso, 1956: Adams, 1959a) support this idea. These
ree sections
shrubby habit. Sect. Muriandra is also related to the cosmo-
politan Sect. Brathys although less closely (Robson, personal
communication). Features common to both include presence
of pale glands, lack of black glands, persistent sepals and
_ Stamens, androecium composed of a ring or narrow band of
_ humerous stamens, and parietal placentation.
STUDIES IN THE GUTTIFERAE. I. 7
Plants belonging to species of rami ontg Sect. Myriandra
be d by tion of the following charac-
teristics: woody stems (except ne species) ; styles which
are closely appressed their entire length at anthesis ; minute
stigmas (i.e. not capitate) ; presence of translucent or pellu-
cid-puncate secretory glands (dots and/or elongate sacs) in
the stems, leaves, sepals, and petals ; and the absence of black
glands. In addition, the leaves and sepals of the plants of
sixteen species (Subsect. Centrosperma) possess a small
groove or articulation at the base (see Fig. 1). Articulated
leaves occur in other species of Hypericum but the combina-
tion of grooved leaves and sepals is known to me in only one
of these, H. styphelioides A. Rich., a plant of Cuba and
British Honduras.
KEY TO THE SPECIES
A. Leaves clasping.
B. Flowers with 5 sepals and 5 petals; inflorescence a compound
dichasium; bark becoming corky, especially on older stems ............
tifolium
B. a with 4 sepals and 4 petals; flowers utary: “bark thin,
corky 27. H. petali
A. hace sessile, not clasping.
Cc. aa tetramerous (except in occasional flowers) : flowers
solita:
D. Gyn i a 2-carpellate, 2-styled.
E. Pedicels elongate, reflexed soon after Prigrpn subtending
bractlets at the base of the pedicel ......., 30. H. suffruticosum.
E. Pedicels short, erect even at frui' pr ei subtending
bractlets een eee to the sepa!
F. Plants erect, usually with a single stem, freely branched
well above the ground level, often becoming 1. 0-1.5° m =
(Fig. 3) 28. H.h
. Plants decumbent, with several prostrate stems arising
from a primary rootstock near ground level, each with
numerous erect branchlets and forming o% compact mats,
rarely over 0.3 m tall (Fig. 3) -- . H. stragulum.
D. Gynoecium 3-carpellate, 3-styled oe 2 or 4 carpels
and styles, especially in H. microsepalum
G. Outer pair of sepals only slightly Taree than the inner;
leaves linear to broadly oblong or oblanceolate, late, usually less
than 1.2 cm long and 0.5 em wide; margins of the leaf and
vy
Ae
:
By
elliptic-oblong or ovate-cordate, usually more than 1.0 em
long and 0.5 cm wide; margins of the ov oa coe sa amid
thin and hyaline, the edge thus like a honed i
act
8 PRESTON ADAMS
adventitious shoots from horizontal roots often aoe
(Glades, Highlands, and DeSoto counties, Florida )..
26. H.
. Outer pair of sepals broadly ovate or suborbicular, 10-15
mm wide; stems usually simple, or sparingly branched;
adventitious shoots lacking. (Eastern Texas to Kentucky,
Florida, and New Jersey) 25. H. stans.
C. Perianth pentamerous (except in occasional flowers) ; inflores-
-many-flowered dichasium (except H. frondosum and
H. buckleyi which often have solitary flowers).
I. Gynoecium oie amare 5-styled (often 3, 4, or 6 carpels and
styles present also).
J. Capsules deeply lobed; am: flowered (usually
15 or more) terminal ea Gichakinin and with similar
dichasia in upper 2-4 leaf axils, producing a panicle-like
aspect. (Mississippi Embayment region, — Arkansas,
Louisiana, Mississippi, and Tennessee) .... 2. H. lobocarpum.
J. Capsules unlobed or only slightly so; inflorescence a few-
flowered (3 or 7, rarely more) terminal, simple or compound
dichasium, rarely with similar dichasia in upper 1 or 2 leaf
axils. (Dune areas on the shores of the Great Lakes; inland
occasionally, ater in central Wisconsin and the Ottawa
River OF Quebec) s2.:stisic.sssssiccssevsiccsnstevetsoose 1. H. kalmianwm.
I. Gynoecium prone Sa Batvied (rarely 2, 4, or 5 present also).
K. Mature leaf with an expanded blade, not needlelike, usually
over 2 mm wide; sepals usually broadened, i.e. linear-elliptic,
oblong-ovate, or ‘Spathulate.
Flowers terminal and solitary, or occasionally in 3- flow-
ered dichasia.
M. Leaves without an articulation at base (see Fig. 2),
elliptic, elliptic-obovate, or obovate, usually less than 2.5
cm and 12 wide; plants decumbent, stems pros-
of North Carolina and Georgia) ............ 18. H. buckleyi.
M. Leaves with an iculation or groove at base (see Fig.
), te-] late, usually well over
3 cm long and 1 em a ike plants erect, stems not pros-
trate. (Caleareous soils, especially in the cedar glades
of central Tennessee and adjacent northern Alabama) ..
5. H. frondosum.
L. Flowers in 3-many-flowered terminal and/or subterminal
dichasi:
N. Shrubs, stems often 1-2 m tall.
0. Leaves and sepals without a basal articulation or
groove (see Fig. 2); inflorescence appearing naked
due to the much reduced bracts.
P. Leaves or or oblong- lanceolate, usually
je eee a
“Ty persistent long after fruit mavuritys seeds less
STUDIES IN THE GUTTIFERAE. I. 9
than 0.8 mm long; placentation “enys a
. H. cistifolium.
P. Leaves elliptic to orate lanoiie bi ‘yell over
1.0 em wide; stamens more than 75, usually decidu-
ous before fruit asics seeds over 1.2 mm long;
placentation pseudo-axile, i.e. the placentae project
inwardly toward the ovary axis, especially in the
lower half.
Q. Inflorescence a many-flowered (rarely 3) dicha-
sium; mature capsules 3.5-7 mm long, 3-4.5 mm
wide; raphe of seeds well- werecs forming a
conspicuous keel = iflorum.
Inflorescence a 3-flowered al occasionally
ae 6 flowers, or reduced to 1; mature capsules
6-14 mm long, 4.5-7 mm wide; Paphe of seeds only
slightly developed, forming a low ridge, but not
a conspicuous kee! 17. H. apocynifolium.
O. Leaves and sepals with a basal articulation or groove
(see Fig. 1) ; ee appearing leafy due to the
foliaceous bract:
R. Largest leaves 15-8. 0 em long (rarely over 3.2 em) ;
seeds 0.7-0.8 mm long; placentation parietal.
(Coastal Plain, North Carolina to me Florida,
westward to southeastern Texas) ...- at galioides.
a
sie at ovary base.
s. Mature capsules less than 6 mm long and 3 mm
inflorescence presenting an obpyramidal aspect;
seeds reddish-brown, 0.8-1.3 mm long and 0.27-
0.35 mm wide wasn. 3. H. densiflorum.
S. Mature capsules exceeding 7 mm in length and 3.5
mm wide, lance-ovoid; flowers in 3-7- (rarely 1-)
flowered terminal and 1-7-flowered subterminal
the inflorescence usually narrow and
compactly th ; seeds dark brown or ee hak
1.0-1.7 mm long and 0.32-0.50 mm wide ..
Sy 2 prob icum
N. Perennial herbs, usually less than 1 m tall.
definite
10 PRESTON ADAMS
somewhat spathulate; larger leaves 1.5-3.5 cm long;
plants short and slender, rarely becoming 0.5 m
tall 3. H, ellipticum.
- Leaves elliptic below, gradually becoming linear-
oblong to linear-lanceolate above; larger leaves 3.5-
7.5 em long; plants tall and stout, commonly becom-
earl OB van adh cesta a iar 22. H. adpressum.
T. Plants without rhizomes, but often bearing adventi-
tious shoots from horizontal roots, bases often woody;
young stems obviously flattened, the 2 primary wings
well-developed; placentation parietal; seeds over 1.5
mm long; seed coat coarsely reticulate; raphe well-
developed, forming a conspicuous keel.
V. Sepals equal or nearly so; stamens 45-85; seeds 2.0-
2.7 mm long; seed coat very coarsely reticulate, the
‘Wansyerse striae much more peoneny than the
20. H.
eq
V. Sepals get pneguah the eine pair larger; anees
mens 1: 3 seeds 1.5-1.8 mm long; seed coat
isc shai the pea striae much
more prominent than the transverse ....
a Wage : dulabriforme:
K. Mature emit and sepals linear-subulate or needle-like.
WwW. st leaves usually less than 11 mm long; sepals 4.5
mm long or less (occasionally to 5.5 mm).
X. Mature capsules 5.7-9.5 mm long (rarely less than 5.5
mm); plants often with decumbent stems, producing a
low matted form, usually less than 0.5 m tall; nodes of
ems
on
reticulate, the alveolae square-hexagonal (100X magni-
fication) and in longitudinal rows, the longitudinal
ridges imparting an obscure striated appearance; fresh
leaves usually dull green above, hardly glossy, if at all
12. H. reductum.
X. Mature capsules 3.5-5.5 mm long; plants almost in-
variably erect, often reaching 1.5 m in height; stem
base (stems
more than 4.5 mm long (occasionally only 4 mm).
Y. Bark smooth, metallic-silvery, exfoliating in large thin
STUDIES IN THE GUTTIFERAE. I. 11
inent than the transverse, thus the surface appearing
furrowed. (Bay and Washington counties, Florida) ..
Wocths Eepkine,
Y. Bark variously roughened, usually exfoliating in irreg-
ular strips or flakes; leaves, sepals, and young stems
scarcely or not at all glaucous; seeds less than 1 mm
long; seed coat finely and evenly reticulate.
Z. Plants with decumbent stems, often forming low mats
(rarely over 0.5 m tall), or at least reclining and
straggly. (Along the Fall Line, mostly in the Caro-
linas) 13. H. Uoydii.
Z. Plants definitely erect. (Coastal Plain, North Carolina
to Florida and Mississippi
a. Bark soft and spongy-t pean with conspicuous
vertically-aligned, cord-like laticifers; youngest
stems terete or nearly so, the primary wings poorly
developed; fiowers mostly in terminal and sub-
terminal 3-flowered dichasia (or rarely with termi-
nal and subterminal solitary flowers). (Wakulla
Co., westward to Santa Rosa Co., pgp
. chapmanii.
. Bark usually not Sener y ‘but if 0) ee
the laticifers
flattened, the primary wings well-developed; f = ers
mostly in 7-32-flowered terminal dichasia, the
below not bearing flowers or more frequently ces
similar compound dichasia.
b. Plants slender, less than 1 m tall, the stem rarely
over 0.8 cm thick; apices of sepals and leaves
long-acuminate; resin glands in leaves and sepals
often darkening upon drying; flowering during
May. (Bay, Gulf, Franklin, Liberty, and Wash-
ington counties, Florida) ............----.- 14. H. exile.
b. Plants stout, usually over 1 m tall (often to 4 m),
the stem 1-5 cm thick; apices of sepals and leaves
mucronate; resin glands in leaves and
usually remaining clear on drying; flowering
throughout the summer, or all year in lower
Loans Florida.
.. Bark spongy-thickened, exfoliating in grayish-
reddish tissue-thin layers; leaves with a deep
lower surface ( on living
leaves) ; mature capsules elliptic-oval ; terminal
dichasium usually compactl: rmi-
12 PRESTON ADAMS
ark thin, even on oldest arene not spongy,
exfoliating in flakes or narrow strips; leaves
merely slightly concave below, not grooved,
papillae inconspicuous, poorly developed; mature
capsules oblong; terminal dichasium usually
openly paniculate, inal y-flowered
dichasia present in upper 3-7 leaf axils; leaves
of axilary branchlets usually not crowded ........
10. H. nitidum.
Sect. MYRIANDRA, subsect. CENTROSPERMA Keller, in Engler
& Prantl, Pflanzenfamilien 3 (6-6a) 214. 1895.
Shrubs. Leaves and sepals with an articulation or groove
at the base (Fig. 1). Sepals 5 (rarely 4), deciduous after
fruit dehiscence. Petals 5 deta 4). Withered stamens
deciduous soon after anthes:
: H. prolifi eh
Fifteen species, nine of which are distributed on the Coast-
al Plain of the southeastern United States, Cuba, and British
Honduras, and the remaining six in the central and eastern
United States and adjacent Canada.
1. Hypericum kalmianum L., Sp. Pl. 2:783. 1753
Hypericum bartramium Miller, Gard. Dict. ed. 8, no. 10.
1768.
TYPE: LINN; photograph in A library; sketches and notes
from type by H. K. Svenson, GH.
REPRESENTATIVE SPECIMENS : ONTARIO: Great Duke Island,
Grassl 5554 (MICH). ILLINOIS: Lake Co., Waukegan, Glea-
son & Shobe 331 (DUKE). INDIANA: Starke Co., Bass Lake,
Deam 20133 (IND). MICHIGAN: Alpena Co., Alpena, Fernald
& Pease 3420 (GH). WISCONSIN : Jackson Co., shore of Black
River, Fassett 15716 (wis).
Dunes and rocky shores about Lakes Michigan, Huron,
and Erie; inland in central Wisconsin and the Ottawa River
of Quebec. Flowering late June through early August.
relative, Hyp bocarpum, by various features, espe-
cially of th le and i (see that species). An
sapeaserasnent problem involving the origin
of H. kalmi and its distribution during the Pleistocene
is discernible (McLaughlin, 1931; Hugh H. Iltis, personal
communication).
oO Bee = leh
Gatti Bot. Gaz. 11:275. 1886
STUDIES IN THE GUTTIFERAE. I. 13
Hypericum oklahomense Palmer, Jour. Arn. Arb. 5:128.
TYPE: TENNESSEE: Carroll Co., Hollow Rock, A. Gattinger
August 1867 (Lectotype, F, with dissections and sketches by
Gattinger ; isotype, GH).
REPRESENTATIVE SPECIMENS: ARKANSAS: Pulaski Co., Lit-
tle Rock, Demaree 8207 (A, BKL, GH, NY). KENTUCKY: Callo-
way Co., between Murray and New Concord, Smith &
Hodgdon 4083 (GH, US). LOUISIANA: Webster Parish, 4
miles west of Minden, Correll & Correll 10310 (DUKE, F, GA,
MO, NY, PH). OKLAHOMA: LeFlore Co., Page, Palmer 22228
(GH, holotype of H. oklahomense Palmer; isotypes, A).
TENNESSEE: Chester Co., Sharp et al 9410 (TENN).
Creek and river bottoms, stream banks, moist pinelands,
and ditches, Arkansas and southeastern Oklahoma, south-
ward into eastern Texas, southern Louisiana, and southern
Mississippi, northward into western Tennessee and Massac
Co., Illinois (Mohlenbrock & Voigt, 1959). Flowering mid-
June through early August, occasionally earlier (late May)
or later (September and October).
This species is distinguished from its closest relatives,
Hypericum densiflorum and H. kalmianum, by several
characteristics, especially the hol of the g i
Carpel number, predominantly five in the first two species,
readily separates H. lobocarpum from H. densiflorum (Table
1).
TABLE 1. Comparison of carpel numbers in the gynoecia
three species of Hypericum
species number of carpelmumber (in Ye) gynoecia
collections studied 3 4 § 6 counted
H. lobocarpum 23 O08 -4.0 192 > Wt O08 2014
H. densiflorum 49 0.2 92.1 6.9 08 0.0 3197
H. kalmianum 25 00° 10." 225 162 OF 389
Ovary lobing, best observed in the mature capsule, is pro-
nounced in H. lobocarpum, present slightly in H. densiflorum,
but virtually absent in H. kalmianum. The mature capsule
dimensions overlap, but significantly different trends in
proportion are discernible. The fruits of H. lobocarpum,
often larger than those of H. densiflorum, never reach the
relatively great size attained by H. kalmianum. Capsules 6-8
mm long and over 3 mm wide are produced by H. lobocarpum
frequently but never occur in its eastern relative, H. densi-
florum. Fruits of many individuals, otherwise definitely re-
14 PRESTON ADAMS
ferable to H. lobocarpum, may be the same size as the small-
est ones produced by H. densiflorum, a situation which has
influenced the interpretation of the former as a variety of
the latter (Svenson, 1940; Fernald, 1950; Gleason, 1952;
Gillespie, 1959).
Nowhere does Hypericum lobocarpum overlap the range
of H. densiflorum, the known stations of each being at least
eighty miles apart (Winston Co., Mississippi and Tuscaloosa
Co., Alabama, respectively). Further north in Tennessee the
known populations of these two species are separated by
some one hundred twenty miles (Carroll and Coffee coun-
ties).
3. Hypericum densiflorum Pursh, Fl. Am. Sept. 376. 1814
Hypericum foliosum Jacq., Hort. Schoenbrun. 3:299. Tot
not Aiton, Hort. Kew. ed. 1, 3:104. 1789.
Hypericum prolificwm L. var. densiflorum (Pursh) Gray,
Man. Bot. ed. 2, 50. 1856.
Hypericum glomeratum Small, Bull. N. Y. Bot. Gard. 1:281-
- 1899.
Hypericum interior Small, Bull. Torrey Club 28:359. 1901.
Hypericum revolutum Keller, Bot. Jahr. 58:194. 1923, not
Vahl, Symb. Bot. 1:66. 1790.
TYPE: “On the dry ridges and savannahs of the Virginia
mountains,” but probably no longer extant ( Svenson, 1940).
REPRESENTATIVE SPECIMENS : GEORGIA: Whitfield Co., Har-
per 2032 (GH, NY, isotypes of H. revolutum Keller). NEW
JERSEY: Cumberland Co., South Vineland, Bassett & Long
12 August 1923 (GH, PH). NoRTH CAROLINA: Avery Co.,
Adams 117 (DUKE, FSU, GH, NY, US); Grandfather Mt.,
Huger August 1896 (Ny, type of H. glomeratum Small).
: GA,
MO, SMU). TENNESSEE: Jefferson Co., Dandridge, Rugel July
Adams 109 (NCU).
Abundant in wet meadows, lake margins, open stream
banks, moist pinelands, bogs, and roadside ditches (especial-
ly on the Coastal Plain of the Carolinas) ; also on dry road
£ nts and rocky hillsides. The Coastal Plain of New
Jersey, Delaware, and Maryland, southward (except south-
STUDIES IN THE GUTTIFERAE. I. 15
from extreme th t P Ivani tl 1 to
northern Georgia, eastern Teuweean and central Alabama.
Flowering late June through early September. Harper’s re-
port (1928) of Hypericum galioides from Jefferson, Tusca-
loosa, and Bibb counties, Alabama, is probably based on
plants of H. densiflorum. His comment that “some of the
specimens [were] ten feet tall” is fairly conclusive, since
plants of H. galioides rarely exceed 1.5 m in height. Speci-
mens of H. densiflorum have been collected in Tuscaloosa
County and in adjacent Jefferson County. Hypericum gali-
oides does not grow in Alabama north of Geneva and Escam-
bia counties, adjacent to the Florida border, according to
Harper (1928).
Hypericum densiflorum, superficially similar to H. gali-
oides, is closely related to H. lobocarpum (which see) and, to
a lesser degree, H. prolificum. It may be distinguished from
Ht. prolificum se narrower leaves and more floriferous in-
Pp oecia are the rule in flowers
of H. Sendihomin plants but four, five, and even two carpels
have been observed, even on the same individual. Mature
seeds from the Carolina Coastal Plain populations of this
species have not been available for study.
4. Hypericum prolificum L., Mant. Pl. 1:106. 1767
Myriandra spathulata Spach, Hist. Nat. Vég. 5:440. 1836.
Hypericum spathulatum (Spach) Steud., Nomencl. ed. 2,
789. 1840, not Keller, Bot. Jahr. 58:195. 1923.
TYPE: sheet number 943.20, selected by H. K. Svenson
(1940), LINN; photograph, A library.
REPRESENTATIVE SPECIMENS: ARKANSAS: Drew Co., De-
maree 17648 (A, FSU, MO, NY, PH). NORTH CAROLINA: Union
Co., 1.2 miles north of Sturtevant, Fox & Whitford 3888
(NCSC, SMU, TENN). PENNSYLVANIA: Chester Co., Strafford,
Bartram 1390 (PH).
Rocky slopes, granite outcrops, dry rocky creek beds, road-
side embankments, and abandoned fields. Widespread over
central and eastern United States from Pennsylvania south-
ward to northern Georgia, around the northern rim of the
Mississippi Embayment to southeastern Oklahoma, north-
ward to Iowa and Michigan. Abundant at the summit of
Brasstown Bald, Union Co., Georgia, at an elevation of
4768 feet. Hardy around the Boston, Massachusetts area,
16 PRESTON ADAMS
having become well established at Wellesley and in Essex
County. Flowering late June (southward) through early
September.
This species is very variable but still retains a distinctive
facies. The ovate to lance-ovate, relatively thin-walled cap-
sules, while varying in size, cannot be mistaken for those of
any other species (except H.
Hypericum prolificum is aeoneaiaied torn the related H.
lobocarpum by a less floriferous inflorescence and unlobed
capsules. However, the separation of this species from the
closely related H. frondosum is more difficult. Hypericum
prolificum usually has a smaller, thicker-walled capsule,
smaller petals and seeds, and a more complex inflorescence.
Plants of these two species may hybridize, at least in culti-
vation. A putative hybrid between them has been described
as H. X VanFleetii, according to Rehder (1940). Specimens
in the herbarium of the Arnold Arboretum which bear this
epithet upon their labels are readily referable to H. prolifi-
cum. A highly variable population possibly originating from
hybridization between cultivated plants of these species
grows in a pasture on the Harbison farm at Highlands,
Macon Co., North Carolina. Intensive study of this popula-
tion and those of both species in the Central Basin of middle
Tennessee is greatly needed.
After study of a “vast amount of herbarium-material”
Fernald & Schubert (1948) were unable to identify plants
of this species with the type of H. prolificum in the Linnaean
Herbarium. Consequently, they revived the relatively un-
used epithet H. spathulatum (Spach) Steud. for this species.
Other investigators (Svenson, 1940, 1952; Adams, 1959b)
have amply demonstrated, however, that Fernald & Schu-
Bert s name changing was unnecessary, pointing out that
the “aberrant” appearance of the type specimen (i.e. its
“unusually revolute leaves”) is due merely to wilting prior
to collecting and to insufficient pressure during the drying
process. ee rs caerencge used by Fernald (1950),
isa later
5. Mee eta , Fl. Bor.-Am. 2:81. 1803
Hypericum aureum Bartram, REE: 383. 1791, not Lourei-
ro, Fl. Cochinch. 2:472. 1790.
Hypericum amoenum Pursh, Fl. Am. Sept. 2:375. 1814.
STUDIES IN THE GUTTIFERAE. I. 5 als
Hypericum rugelianum Kunze, Linnaea 24:177. 1851.
Brathydium rugeli (Kunze) C. Koch, Hort. Dendr. 66.
1853.
Brathydium aureum (Bartram) C. Koch, Hort. Dendr. 66.
1853.
Hypericum splendens Small, Bull. Torrey Club 29:291. 1901.
291. 1901.
TYPE: “Hab. in rupibus, ad flumen Tennasee,” Michaux
(P; photograph, GH; sketches from type by H. K. Svenson,
GH).
REPRESENTATIVE SPECIMENS: ALABAMA: Morgan Co., 2
miles south of Tennessee River, south of Huntsville, Godfrey
57520 (FSU). GEORGIA: Dade Co., Cloudland Canyon, east of
Trenton, Cronquist 5293 (GA, GH, NY, SMU); DeKalb Co.,
Stone Mt., Small July 4, 1893 (NY, type of H. splendens
Small). TENNESSEE: Wilson Co., 2 miles north of Lebanon,
Godfrey 59498 (FSU).
A showy plant in flower, Hypericum frondosum is abun-
dant in the cedar glades of the Central Basin in middle
Tennessee. It has also been collected at widely scattered
localities, especially on river bluffs, in southern Kentucky,
extreme eastern Tennessee, northern Alabama, and central
and southwestern Georgia. Adventive plants have been col-
lected in Connecticut, New York, and Massachusetts. Flow-
ering late May through early July.
When best developed, plants of Hypericum frondosum are
readily distinguishable from the related H. prolificum
(which see). The separation becomes difficult with non-
fruiting specimens and cultivated material.
The characteristics of the large-flowered Hypericum which
grows on the slopes of Stone Mountain, DeKalb Co., Georgia,
are within the H. frondosum variation pattern. Small, after
several visits to that famous granite exposure, became so
impressed with these plants that he described them as a new
species, H. splendens. My studies of this population at vari-
ous times during the growing season strongly suggest, how-
ever, that these plants represent merely a geographically
isolated stand of H. frondosum.
6. Hypericum galioides Lam., Encyc. Méth. Bot. 4:161.
1797, not Freyn. & Sint., Bull. Herb. Boiss. 3: 103. 1895
Hypericum azillare Lam., Encyc. Méth. Bot. 4:160. 1797.
18 PRESTON ADAMS
Hypericum fasciculatum Michx., Fl. Bor.-Am. 2:80. 1803,
not Lam., Encye. Méth. Bot. 4:160. 1791, not Tapeyr,
Hist. Abr. Pl. Pyr. 450. 1813.
Hypericum michauzii Poir., Suppl. 3:696. 1813.
Hypericum ambiguum Elliott, Sketch. 2:30. 1821.
Myriandra michauzxii (Poir.) Spach, Hist. Nat. Vég. 5:437.
1836.
Hypericum galioides, var. pallidum Mohr, Contrib. U. S. Nat.
Herb. 6:621. 1901.
Hypericum spathulatum Keller, Bot. Jahr. 58:195. 1923, not
Steud., Nomencl. ed. 2,789. 184
TYPE: “Caroline,” Fraser, P-LA; sketch and notes from
type by H. K. Svenson, GH.
REPRESENTATIVE SPECIMENS: FLORIDA: Gadsden Co., 3.5
miles west of Greensboro, Adams 362 (DUKE, FSU, GA, GH,
NCSC, NY, TENN, US); Liberty Co., Adams 370 (FLAS, MO,
NCU, SMU). GEORGIA: Harper 1155 (A, GH, NY, US, isotypes
of H. spathulatum Keller). SOUTH CAROLINA: Berkeley Co.,
Adams 89 (IND).
Abundant on open stream banks, seepage areas, swampy
clearings, river bottoms, floodplains, hammocks, pond shores,
moist roadside ditches, edges of borrow pits, and low pine-
lands on the Coastal Plain from North Carolina southward
to northern Florida and westward to southeastern Texas
(excepting the Delta region of Louisiana). Flowering late
May through early August, occasionally to late October.
This species, equated by Lott (1938) and Svenson (1940)
with the other members of the Hypericum fasciculatum com-
plex, is distinguished by adult leaves with well-developed
blades. The leaf margins may inroll on dried specimens,
simulating the linear-subulate leaf on plants of the H. fasci-
culatum group. When best developed the lamina is 3-6 mm
wide. Small (1933), Svenson (1940), and Gillespie (1959)
H. galioides with the more northern H. densiflorum,
especially with the populations of the latter in northwestern
Georgia and eastern Tennessee. Superficially, the leaves are
similar but the leaf length/width ratio in the Georgia-Ten-
nevsee H. densiflorum is different, being more narrow in
proportion to the length than those of H. galioides. Also, the
parietal placentation and shorter seeds (with a finer seed
coat ere pi of H. — contrast with these features
as present in g , the ranges of
STUDIES IN THE GUTTIFERAE. I. 19
both species overlap on the outer Coastal Plain of the Caro-
linas but the two are readily identifiable, even from a moving
automobile, and do not intergrade.
The reason for the usage of the name Hypericum galioides
rather than H. axillare for this species (both published by
Lamarck in his 1797 work) is as follows. Early American
botanists, e.g. Pursh (1814) and Elliott (1824), were able
to identify Lamarck’s H. galioides fairly definitely but were
unsure concerning H. azillare. For example, Pursh (1814)
considered this name to be a synonym of H. fasciculatum
Lam. Later authors, including Torrey & Gray (1838),
Coulter (1886, 1897), Chapman (1897), Small (1903, 1913,
1933), and Svenson (1940), have employed the well-ground-
ed H. galioides. I see no reason to change.
THE HYPERICUM FASCICULATUM COMPLEX
The following eight species are closely related. All have
adult leaves which are linear-subulate or needle-like, the
blade being poorly developed. The gynoecia are deeply lobed,
3-carpelled (rarely 2 or 4), and possess parietal placenta-
tion. With the exception of Hypericum lissophloeus, the
seeds are remarkably similar, differing mainly in the seed
coat markings. The asymmetrical, apiculate petals are also
much alike in shape but differ somewhat in size. The cap-
sules of each species exhibit a characteristic shape which is
obviously related to the overall proportions of the fruit at
maturity.
Taxonomic opinion regarding the H:
group has varied considerably. Coulter (1897 ), puzzled by
the seeming “intergradation” of characters, recognized a
single species, H. fasciculatum Lam. Chapman (1897), who
possessed field experience with the complex, was able to
distinguish two species, H. fasciculatum and a shorter-leaved
plant which he called H. aspalathoides Willd. Small (1903,
1913, 1933) seems to have followed Chapman’s treatment.
Svenson (1940) considered these taxa as varieties of H.
galioides Lam., a perfectly distinct although closely related
species. Svenson’s conception of H. galioides also included
plants of H. densiflorum Pursh (Svenson, 1940, plate 587,
figure 7). It is understandable, therefore, why he concluded
that the plants represented by the names H. fasciculatum
5 er
20 PRESTON ADAMS
sufficient properly prepared sp of the H.
group no doubt caused Svenson to conclude further that
“some of these variations of H. galioides are of doubtful
geographic significance and are probably not the equivalent
of subspecies of zoological usage.”
In recent years observant collectors in northern Florida,
especially the Panhandle region, have been greatly puzzled
by the Hypericum fasciculatum complex (R. K. Godfrey, R.
Kral, and E. L. Tyson, personal communication). Their
attempts to understand the group using the treatments of
Small (1933) and Svenson (1940) have met with frustra-
tion; the reasons for this have now become clear (see below).
I was introduced to the perplexities of H. fasciculatum an
its relatives in June of 1958 while collecting in Florida with
R. K. Godfrey. My field experience that summer combined
with scrutiny of herbarium specimens led me to conclude
that “the group is composed of at least four very striking
variation patterns” and that “an inseparable intergrading
series most probably does not exist” (Adams, 1959a). It was
evident to me also that these “variation patterns” were most
probably biological entities worthy of specific recognition.
Intensive field observation of the Hypericum fasciculatum
complex by both R. K. Godfrey and myself, especially in
northern Florida and southern Georgia, during the past two
years, has revealed that the long-standing taxonomic con-
fusion is due primarily to the presence of eight morpho-
logically distinct taxa, each with a distinctive geographic
range and specific habitat requirements. Close study, in the
laboratory and in the field, has revealed no evidence of inter-
gradation among these several taxa, despite their close asso-
ciation in the same habitats. Their flowering periods, while
distinctive for each species, often overlap. Insects, chiefly
bumblebees and honey bees, have been seen to visit the
flowers of two or more species in succession. There is no
evidence of polyploidy even though the chromosomes of only
five taxa have been studied.
It is my considered opinion that each of the eight kinds of
plants in the Hyp i fascicult compl ts a
H. galioides “form an inseparable intergrading series.” In-
Piha fascial
tedly, they are closely related and may well be phylogenetical-
ly “young” species, compared to the other members of
STUDIES IN THE GUTTIFERAE. I. 21
Hypericum Sect. Myriandra. All evidence suggests that the
eight species share a common ancestry and that they are
related through H. galioides to the other members of Sub-
sect. Centrosperma.
-7. Hypericum lissophloeus P. Adams, sp. nov.
Arbuscula usque ad 4 m alta, trunco ad 4.5 cm in diametro.
Rami juveniles, folia et sepala conspicue glauca. Cortex
laevis (velut nitidus), statu juvenile castaneo-fuscus, statu
adulta argenteo-metallicus, in plagulis magnis crispis decor-
ticans. Folia basi articulata, lineari-subulata, lamina vix ex-
pansa, valde et carnoso-marginata, 9-17 mm longa, 0.5-0.75
mm lata, subtus minute papillata. Incrementa vegetativa
indeterminata, cum floribus solitariis vel subinde dichasio
tribus floribus. Sepala basi articulata, lineari-subulata, 7-8
mm longa, 0.5-0.75 mm lata, ante capsulae dehiscentiam
decidua. Petala flava, inaequalia, 10-12 mm longa, 5-6 mm
lata. Stamina 170-221, 8-9 mm longa. Gynoecium 6-9 mm
longum, 3- (vel raro 4-) carpellatum, placentatione parietale,
cum 3 (vel raro 4) stylis. Capsula matura 6-7 mm longa, 2.5-
3.5 mm lata. Semina fusca ad brunnea, carina aliquid humile,
1-1.6 mm longa, 0.5-0.6 mm in diametro, testa grossissime
reticulata, striis longitudinalibus quam transversalibus
prominentioribus, adspectu rugosa.
Shrub, to 4 m tall, with stem to 4.5 cm thick. Young stems,
leaves, and sepals conspicuously glaucous. Bark smooth (as
if polished), chestnut-brown when young, metallic-silvery on
older stems, exfoliating in large thin curled plates. Leaves
articulate at base, linear-subulate or needle-like, lamina poor-
ly developed, edges thickened and turned abruptly downward
(inrolled on drying), 9-17 mm long, 0.5-0.75 mm wide,
minutely papillate beneath. Vegetative shoots often continu-
ing growth throughout the season, producing solitary flowers
(or occasionally 3-flowered dichasia) from the leaf axils.
Sepals articulate at base, linear-subulate or needle-like, 7-8
mm long, 0.5-0.75 mm wide, deciduous before fruit maturity.
Petals yellow, asymmetrical, 10-12 mm long, 5-6 mm wide.
Stamens 170-221, 8-9 mm long. Gynoecium 6-9 mm long,
carpels 3, rarely 4, placentation parietal, styles 3, rarely 4.
Mature capsule 6-7 mm long, 2.5-3.5 mm wide. Seeds tan
to dark brown, raphe developed into a moderately low keel,
10-1.6 mm long, 0.5-0.6 mm wide, seed coat very coarsely
22 PRESTON ADAMS
reticulate, soqiaecone! striae much more prominent than
a furrowed Suga Noe Specific
epithet apt de Greek meaning “smooth bark.”
TYPE: FLORIDA: Bay Co., shores of Merial Lake, ca. 10
miles north of Panama City, Godfrey & Triplett 59844
(Holotype, GH; isotypes, DUKE, F, FLAS, FSU, GA, IA, ILL, K,
MSC, MO, MT, NCU, NCSC, NY, PAM, SMU, TENN, TEX, UC, US,
USF, VDB, VPI, WIS).
REPRESENTATIVE SPECIMENS: FLORIDA: Bay Co., shores of
Merial Lake (exact type locality), Adams 685 (DUKE, FLAS,
GA, K, MSC, NY, SMU, US) ; same locality, Adams 734 (F, GH,
ILL, K, MO, NCU, TEX) ; Washington Co., border of Long Pond
north of Redhead, Godfrey 60793 (FSU).
Known only from Bay and Washington counties, Florida.
Abundant in sandy soil on the shores of sinkhole ponds and
lakes. Frequently growing in water to 1.5 m deep. Flower-
ing p Sporadically from late May until October.
hl is distinguished from its closest
melativens in the H. fascicuJatum complex by many features,
including a smooth, polished, metallic-appearing bark which
exfoliates like birch (on older stems) ; the slender, wand-
like, lax or drooping younger stems; a large seed size and
a furrowed seed coat; and glaucous leaves, sepals, and young
stems. It grows in company with one or both of two other
species of the H. fasciculatum group. Plants of H. reductum,
low, much-branched, and suffrutescent, grow about the bases
of H. lissophloeus plants on the upper pond or lake shores
out of the water. It is sometimes associated with H. fasci-
culatum either in the water or on upper drier shores, in the
latter situations the three occurring together. Since the
three grow together and are readily identifiable, no inter-
gradation being apparent, this is taken as additional evi-
dence of their specific nature.
“8. Hypericum chapmanii P. Adams, nom nov.
Based on Hypericum arborescens Chapman, FI. S. U. S.,
ed. 2, suppl. 2, 680. aitbcaatiicas Symb. Bot. 2:86. 1791.
TYPE: FLORIDA: round Apalachicola, Chap-
man, but not yet located. Bilt. Herb. no. ‘5735a, apparently
| in 1893, bears the name Hypericum arborescens
and illustrates Chapman’s conception of this taxon (A, NY)-
REPRESENTATIVE SPECIMENS: FLORIDA: Bay Co., northeast
STUDIES IN THE GUTTIFERAE. I. 23
of Vicksburg, Adams 513 (FSU); Franklin Co., 8.6 miles
west of Apalachicola, Adams 508 (FLAS, FSU) ; Gulf Co., 3.6
miles of Wewahitchka, Adams 340 (DUKE, GH, PAC, PUR,
SMU, VDB) ; Liberty Co., 8 miles south of Hosford, Adams
287 (FSU) ; Santa Rosa Co., Tyson 485 (FLAS).
Abundant in flatwoods depressions, margins of cypress
ponds, and borrow pits, from the Ochlocknee River (between
Wakulla and Liberty counties, Florida) westward to Santa
Rosa County, Florida. Not known to grow more than thirty-
five miles inland from the Gulf of Mexico. Flowering early
June through middle July.
Hypericum chapmanii stems are characterized by a soft
bark with conspicuous, vertically-aligned, resin-filled latici-
fers occurring at discrete levels between layers of cork often
3 or 4 mm thick. Upon being torn apart, the layers of cork
appear striated or fluted owing to the large size of the latici-
fers running through them. Within, the cork is reddish-
brown to almost cinnamon-colored; on the surface it
weathers to gray. The resin in the laticifers, clear and
nearly colorless when fresh, gradually hardens, turning dark
brown or almost black. As the stem grows in circumference
the bark cracks externally and the cork layers disintegrate
and fall away, leaving the more resistant strips of hardened,
resinous laticifers, the whole presenting a stringy, ragged,
unkempt appearance. Frequently the bark attains a thick-
ness of 3 or 4 cm, especially near the bases of older stems.
The soft spongy feature is evident even on young stems
under 5 mm thick. The soft bark is sometimes used by mice
to line their nests (E. L. Tyson, personal communication).
These animals also build their nests in the branches of large
plants of this species, according to Tyson.
When best developed, plants of Hypericum chapmanii
usually have a single stem which produces a dwarf tree-like
aspect, a feature which doubtless influenced Chapman in his
choice of the name “art ” Individuals often attain
a height of 3 or 4 m and a thickness of 10-15 cm, especially
near the base. The strongly ascending, light green leaves
have a deep channel or groove on their lower surface along
each side of the midvein. Numerous tiny (but obvious
under 10X magnification) white papillae cover the surface
of these canalicula. The flowering season begins about June
1 and extends through late July. A second crop of flowers
24 PRESTON ADAMS
is never produced in the fall, at least not during the two
years I have observed these plants.
The distributional area includes or overlaps those of the
following related species: H. caoaaenecrs H. nitidum, H.
exile, H. reductum, H. lissophl and ji
However, observations in the field and on specimens of
fresh and dried plants in the laboratory during the past two
years have revealed no evidence of intergradation.
9. Hypericum fasciculatum Lam., Encyc. Méth. Bot.
4:160. 1797
Not Michx., Fl. Bor.-Am. 2:80. 1803, not Tapeyr., Hist.
Abr. Pl. Pyr. 450. 1813.
agin” aspalathoides Willd., Sp. Pl. 3:1451. 1803, nom.
Myron bruise Spach, Hist. Nat. Vég. 436. 1836, nom.
[ious ‘golivides Lam. var. aspalathoides (Willd.) T. &
G., Fl. N. Am. 1:672. 1840
Bupecnon galioides Lam. var. ase sadn (Lam.) Sven-
son, Rhodora 42:12. 1940.
TYPE: “Caroline,” Fraser, P-LA; photograph, GH ; sketches
and notes from type by H. K. Svenson, GH.
ILLUSTRATION: Svenson, 1940, plate 587, figure 1.
REPRESENTATIVE SPECIMENS: FLORIDA: Liberty Co., 7.7
miles east of Bristol, Adams 357 (DUKE, FLAS, FSU, GH, NCSC,
SMU, NCU, USF) ; Polk Co.,7 miles west of Frostproof, Adams
393 (FLAS, FSU, USF) ; Wakulla Co., 1 mile north of Panacea
bridge, Adams 305 (FLAS, FSU, DUKE); Washington Co.,
Long Pond, north of Redhead, Godfrey 60792 (FSU).
GEORGIA: Echols Co., 8 miles west of Fargo, Adams 820
(FSU, GA, GH) ; Lee Co., 3.5 miles south of Smithville, Adams
3 (BH, GA, IA, IND, MSC, MT, NA, NCU, NY, SMU, US).
Common around the margins of cypress ponds and small
lakes, in low pinelands and ditches, throughout Florida,
southern Georgia, southeastern South Carolina, southern
Alabama (Harper, 1928), and southern Mississippi. Flower-
ing all year in southern Florida, beginning in late April-
early May in northern Florida, sporadically through the
summer and until November in northern Florida and
Plants of Hypericum fasciculatum have a spongy-thick-
STUDIES IN THE GUTTIFERAE. I. 25
ened bark which exfoliates in tissue-thin sheets. Vertically-
aligned, resin-filled laticifers occur in definite layers between
strata of cork. The resin in the laticifers is liquid and
nearly colorless when fresh but gradually hardens to a
reddish-brown color. As the stem grows in thickness the
resinous layer cracks, exposing the cork which peels away in
brown or cinnamon-colored tissue-like layers. The laticifers,
being more resistant to weathering, remain longer but event-
ually exfoliate in elongate flakes or strips.
Both Hypericum fasciculatum and its relative, H. chap-
manii, possess corky barks. However, each is distinctly
different, being easily recognizable, even by touch. The bark
of H. fasciculatum never attains the softness nor the thick-
ness of H. ch ii. Anatomically, the cork strata of the
former species do not become as thick (i.e. have fewer cell
layers) as do those of the latter. Since the laticifers of H.
fasciculatum are relatively small and thread-like, the inter-
vening layers of cork are smooth, in marked contrast to the
conspicuously striated or fluted aspect produced by the
large resin tubules in H. chapmanit.
Other characteristics of Hypericum fasciculatum which
distinguish it from H. chapmanii include darker green,
spreading to only slightly ascending leaves, a different in-
florescence (see key), strongly flattened young stems, earlier
initiation of flowering, development of a second crop of
flowers in the fall (see below), smaller maximum height and
thickness of stems, and much more extensive geographic
range.
The distributional range of Hypericum fasciculatum over-
laps or encompasses that of six related species. In the Bay-
Gulf-Franklin-Liberty county region of Florida, plants of
this species grow in close iation with individuals of H.
Ie, hl, Hck - I
i p ‘ hyphyll H. nitidum,
_H. exile, and H. reductum. No evidence of intergradation
among these species has been discove: :
At fruit maturity plants of Hypericum fasciculatum may
produce a second increment of vegetative growth, the young
shoots frequently bearing flowers at their tips. In the north-
ern portion of the range this second crop of flowers rarely
matures, usually being killed by frost in November.
The seedling leaves of plants of this species possess a
strikingly different shape than the adult ones. These juvenile
26 PRESTON ADAMS
leaves are obovate, elliptic, or elliptic-oblong, with the blade
well-developed, but gradually become modified in form and
structure at successive nodes up the stem. Eventually, adult
leaves with a linear-subulate or needle-like shape are formed.
“Stump sprouts,” originating when older stems are cut near
the base (as along mowed highway right-of-way), show a
similar leaf develop EO lly, a mature plant will
suddenly form juvenile Jeaves (see illustration in Svenson,
1940, plate 587, figure 2). Reversal to juvenility may occur
also in greenhouse plants. This phenomenon has been re-
sponsible for some of the confusion concerning H. fascicula-
tum and the related H. galioides, the juvenile leaves of the
former having been interpreted as evidence of intermediacy
between these species.
10. Hypericum nitidum Lam., Encyc. Méth. Bot. 4:160. 1797
Myriandra nitida (Lam.) Spach, Hist. Nat. Vég. 5:436.
183
Hypericum cubense Turcz. [?], Bull. Soc. Nat. Mosc. 31:384.
1858, see discussion below. Type: Linden 1696, but not
yet located.
Hypericum galioides Lam. var. cubense Griseb. [?], Cat. Pl.
Cub. 39. 1866, see discussion below. Type: Wright
2126 (GH, NY).
TYPE: P-JU; photograph, GH ; fragment of type, GH.
REPRESENTATIVE SPECIMENS: FLORIDA: Bay Co., just east
of Callaway, Adams 350 (DUKE, F, FLAS, FSU, GA, GH, K, MO,
NCSC, NY, SMU, US). GEORGIA: Colquitt Co., Adams 33 (FSU,
GA, GH, NA) ; Irwin Co., 11 miles southeast of Fitzgerald,
Adams 555 (FSU, GA) ; Brooks Co., 2.8 miles east of Barney,
Adams 376 (DUKE, FLAS, FSU, GH, IA, K, MO, MSC, NCSC, NCU,
SMU, TEX) ; Thomas Co., 2 miles north of Pavo, Adams 334
(DUKE, FSU, GA).
Moist soil along open stream banks and pond margins,
low pinelands, highway fill excavations, and roadside ditches.
Southern Georgia (excepting the coastal counties) nearly to
the Fall Line and southward into the Panhandle of Florida
from Liberty to Escambia counties. Also in Baldwin Co.,
Alabama, Lexington Co., South Carolina, and Brunswick Co.,
North Carolina. Flowering early June through early
August, occasionally later.
Plants of Hypericum nitidum are usually very abundant
STUDIES IN THE GUTTIFERAE. I. 27
locally, often forming dense thickets. In Bay, Gulf, and
Liberty counties, Florida, these plants are often 2 m or
more in height and frequently 3-4 cm in diameter near the
base. The bark is thin, dark brown or reddish, and exfoli-
ates in flakes or narrow strips. Plants of this species greatly
resemble those of H. brachyphyllum; indeed, the two species
seem to differ mainly quantitatively, the latter appearing
to be a “miniature” of the former. However, they seem
amply distinct morphologically, being easily distinguished
even when growing in the same habitats. Plants of H. niti-
dum, besides having much longer and somewhat wider
leaves, are typically much taller and the stems thicker in
diameter. Furthermore, those of this species usually begin
to flower at least two weeks earlier than those of H. brachy-
phyllum. In addition, each of these species possesses a dis-
tinctive geographic range, although their distributions
overlap throughout the Panhandle of Florida and the lower-
most portion of Georgia. Hypericum nitidum grows farther
north toward the Fall Line in Georgia but does not occur
in Florida southeast of Franklin County. Further intensive
study is greatly needed to establish the exact morphological
limits of each of these species.
The plants of the Hypericum fasciculatum complex which
grow in Cuba and British Honduras seem to be closest to
H. nitidum, with the exception of Grisebach’s H. limosum
(isotype, Wright 2125, GH) which greatly resembles H.
brachyphyllum. The external morphology of the leaves of
the Cuban specimens compares readily with that of the
United States H. nitidum. However, stem samples with bark
taken from near the base of the plants are needed before
the Cuban material can be definitely classified. Interestingly
enough, Asa Gray, upon his last visit to Paris in 1881, con-
cluded that Grisebach’s type of H. galioides var. cubense
(Wright 2126) is “exactly H. nitidum Lam.” (notes in
Gray’s handwriting, GH). An isotype of this taxon (GH)
is comparable to many specimens of H. nitidum from Florida
and Georgia.
ll. Hyperi brachyphyll (Spach) Steud., Nomencl.
1:787. 1840
Based on Myriandra brachyphylla Spach, Hist. Nat. Vég.
5:435. 1836.
28 PRESTON ADAMS
Hypericum limosum Grisebach [?], Cat. Pl. Cub. 39. 1866,
see discussion below.
TYPE: FLORIDA, Apalachicola, Drummond (Isotypes, GH;
K, two sheets, not seen).
REPRESENTATIVE SPECIMENS: ALABAMA: Covington Co., 7.4
miles south of Opp, Shinners 27453 (FSU). FLORIDA: Cal-
houn Co., 2.5 miles south of Blountstown, Godfrey 57582
(FSU); Franklin Co., west of Apalachicola, Adams 309
(Fsu) ; Levy Co., 14 miles south of Chiefland, Adams 388
(FSU) ; Santa Rosa Co., 6 miles east of Munson, Kral & Red-
fearn 2946 (FSU); Wakulla Co., 1 mile north of St. Marks,
Godfrey 57862 (FSU). GEORGIA: Coffee Co., 8 miles east of
Douglas, Adams 830 (Fsu, GA) ; Early Co., 9 miles southeast
of Blakeley, Adams 791 (DUKE, FSU, GA, USF). MISSISSIPPI:
Jackson Co., Ocean Springs, Demaree 32846 (FSU).
Fic. 1-2. Morphology of the leaf and a ases in cides Seet. Myriandra.
Fie. 1. Left. Hypericum brachyphyllum (Spa ach) Steud., below, bases of two opposite
leaves, each with a conspicuous articulation or groove at the junet
Above, bases o sacking sepals (two visible), each
with the receptas Fic. 2. Right. caraagtongls mudi run
Note the oe ace of an articulation. Magnificatio:
Abundant in low pine flatwoods, pond margins, borrow
pits, and roadside ditches, especially in the Florida Pan-
handle region and scattered stations along the Florida west
coast to Collier County, westward into southern Mississippi,
STUDIES IN THE GUTTIFERAE. I. 29
northward into lower Alabama and Georgia. Flowering
late June through early September, occasionally sporadically
to late November, or later in southern Florida.
Plants of Hypericum brachyphyllum characteristically
possess a rounded bushy aspect due to the production and
retention of numerous branches from the upper three-
fourths or one-half of the stem. The height is usually about
0.5 to 1.0 m although occasional individuals may reach 1.5
m. Stem diameter near the base rarely exceeds 1.5 cm. The
bark is relatively thin and tight, being composed of alternat-
ing strata of cork cells and thread-like laticifers, and ex-
foliates in small plates of strips, the whole presenting a dark
reddish or brownish appearance.
Although plants of Hypericum brachyphyllum often grow
in ane association with individuals of H. nitidum, H. exile,
Hi. H. i, and H, reductum, no inter-
gradation has been observed. Since it was described by
Spach in 1836 few, if any, authors have recognized the bio-
logical distinctiveness of this species. Coulter (1897) in-
cluded it within his concept of H. fasciculatum. Chapman
(1897), who knew the species in the type region, referred it
to H. aspalathoides Willd. I cannot be certain that Chapman
also knew the related H. reductum which grows along the
coast in the Apalachicola area. If so, then he did not dis-
tinguish between them. Small’s treatments (1903, 1913,
1933) were essentially the same as that of Chapman. Sven-
son (1940) made no mention of the plants now referred to
H. brachyphyllum but he did recognize H. reductum (as a
variety of H. galioides, however). He demonstrated also
that H. aspalathoides Willd. was merely a re-naming of H.
fasciculatum Lam.
That the plants of Hypericum brachyphyllum are amply
distinct from those of H. reductum has become evident to me
from field observations extending over the past two years.
In addition to the features listed in the key, each of these
two species possesses different (although slightly overlap-
ping) flowering periods, geographic ranges, and habitat
preferences. Plants of H. reductum begin to flower earlier
than do those of H. brachyphyllum. Flowers appear on
plants of the former species in late April in southern penin-
sular Florida and early June in western Florida. However,
those of the latter species do not begin to flower until the
30 PRESTON ADAMS
last week of June (except for sporadic individuals, especially
in the southern portion of its range). Furthermore, the
length of the flowering period is different in these two
species. That of H. reductum is usually completed by the
first of July whereas that of H. brachyphyllum extends
through August. There is a tendency for plants of both
species to produce a second crop of flowers during the fall
but this phenomenon is much more pronounced in the latter.
Each of these species has different habitat preferences,
the plants of Hypericum reductum being adapted to some-
what drier sites than those of H. brachyphyllum. Ecology,
no doubt aided by past geological events, has influenced the
creation of the distinctive distributional patterns which are
discernible in these species. The geographic range of H.
reductum appears to be composed of four isolated segments
(see below). In each of these, there seems to be a relation-
ship between the present distribution and the presence of
recent or fossil sand dunes. The distributional pattern pre-
sented by plants of H. brachyphyllum is more or less continu-
ous instead of being fragmented. The present distribution
does not show any relationship with coastal physiography,
the plants growing in low wet pinelands inland from the
ocean.
Additional evidence supporting my interpretation of speci-
fic status for each of these two taxa has been obtained by
observations on sympatric populations. During the course
of my field studies I have encountered three widely separated
stations where plants of both species were growing in close
association. Two of these are in heast
Georgia, one being three miles south of Pembroke in Bryan
County, and the other about four miles west of Eulonia in
McIntosh County. The third sympatric association is in
Franklin Co., Florida, about five miles east of Eastport. At
each of these places plants of each species were immediately
and easily distinguishable, even though growing within a
few feet of each other. The habitat differences were main-
tained at each site also, with H. reductum plants on the
slightly drier and better drained soils. Further analysis of
population samples made at these stations is in progress.
Grisebach’s Hypericum limosum, type: Wright 2125 (GH,
NY), appears to be closest to H. brachyphyllum. However,
STUDIES IN THE GUTTIFERAE. I. 31
better specimens must be available before a definitive inter-
pretation can be made.
12. Hypericum reductum P. Adams, stat. & nom. nov.
Based on Hypericum fasciculatum Lam. var. aspalathoides
T. &G., Fl. N. Am. 1:672. 1840, not H. aspalathoides Willd.,
Sp. Pl. 3:1451. 1803, nom. superfl.
Hypericum fasciculatum Lam. var. 8 T. & G., Fl. N. Am.
1:160. 1838.
Hypericum galioides Lam. var. reductum Svenson, Rhodora
42:14. 1940, nom. superfl. for var. aspalathoides T. & G.
TYPE: North Carolina, Wilmington, Curtis (Lectotype,
GH; isotype, NY, fragment). Cited by Torrey & Gray (1838)
under their description of H. fasciculatum var. 8.
ILLUSTRATION : Svenson, 1940, plate 587, figure 5.
REPRESENTATIVE SPECIMENS: FLORIDA: Bay Co., Panama
City Beach, Godfrey & Triplett 59815 (FSU) ; Franklin Co.,
Alligator Point, Adams 298 (DUKE, FSU, VDB) ; Highlands
Co., Hicoria, Adams 402 (DUKE, FLAS, FSU, GH, USF) ; Lake
Co., 4 miles north of Altoona, Adams 599 (F, FSU, GH, MO) ;
Marion Co., 1 mile east of Lynne, Adams 610 (FSU, GA, GH) ;
Oklaloosa Co., 4 miles east of Ft. Walton, Thorne & David-
son 17336 (FSU). GEORGIA: Bryan Co., 2.6 miles west of
Pembroke, Adams 540 (FSU) ; Bulloch Co., 17 miles south-
east of Statesboro, Adams 535 (FSU, GA, GH, NY); Evans
Co., 5.2 miles west of Claxton, Adams 544 (FSU) ; Tatnall
Co., 3 miles northwest of Reidsville, Cronquist 5342 (FLAS,
GA, GH, NY, SMU, US). NORTH CAROLINA: New Hanover Co.,
Carolina Beach, Godfrey 1260 (BKL, DUKE, FLAS, GA, GH, MO,
NCU, NCSC, NY, SMU, TENN, US).
The range of Hypericum reductum appears to be composed
of four allopatric segments. In the Wilmington region of
southeastern North Carolina plants of this species grow
among the dunes and dune hollows along the coast. Several
stations are also known further inland in that area. It is
very abundant over many acres of longleaf pine, scrub-oak
sand ridge between Wilmington and Southport (R. K. God-
frey, personal commuication). A second segment occurs in
the triangle formed by the towns of Swainsboro, Jesup, and
Savannah, Georgia. Here the plants grow on the sands of
fossil dunes (e.g. east side of the Canooche River, 8 miles
east of Pembroke, Bryan Co.) and on the sandy soil of dry
32 PRESTON ADAMS
pinelands, roadside embankments, and borrow pits. The
third population occurs in central peninsular savbees from
Putnam County (east of G High-
lands and Charlotte counties. The fourth ly of the
range is along the coast of the Gulf of Mexico from Franklin
Co., Florida, westward to Escambia County. In nothern Bay
and southern Washington counties, Florida, H. reductum
grows in the sandy soil on the shores of sinkhole ponds and
lakes, its branches frequently touching the bases of H.
lissophloeus plants. The former species also Occurs in close
association with plants of H. fasciculat: i:
and H. brachyphyllum. Flowering late April through late
June in central and southern Florida, occasionally sporadic-
ally earlier in the spring or the fall; during June and oc-
casionally July and November in western Florida ; mid-June
through late July in Georgia; and early June through mid-
August in North Carolina.
Plants of Hypericum reductum possess a low, decumbent,
matted aspect, especially when growing among the loose
sands of dunes and dune hollows. In dense stands of grasses,
palmetto, gallberry, and other plants an erect habit may be
formed but a reclining and straggly tendency is still re-
tained. Other features include the short leaves, four well-
developed secondary wings on the young stems, and the
straight-sided, elongated, mature capsules.
Svenson (1940), following Torrey & Gray (1838), cited
Hypericum tenuifolium Pursh (Fl. Am. Sept. 2:377. 1814)
as a synonym of H. galioides var. reductum. Pursh’s descrip-
tion could be interpreted as fitting either H. lloydii or H.
reductum. The specimen which he cited (collected “In
Georgia” by Enslen) is not present in the Enslen Herbarium
at Vienna (K. H. Rechinger, personal communication). A
fragment of an Enslen collection of Hypericum is at Phila-
delphia but there is no indication on the sheet that it repre-
sents the same ein cited by Pursh. This specimen is
definitely H. red
413. aaa loydii (Svenson) P. Adams comb. nov.
Based on Hypericum galioides Lam. var. Lloydii Svenson,
Rhodora 54:207. 1952.
TYPE: SOUTH CAROLINA: Aiken Co., Graniteville, Eggert
in 1898 (Holotype, ny; isotypes, MO, US).
STUDIES IN THE GUTTIFERAE. I. 33.
ILLUSTRATION: Svenson, 1940, plate 587, figure 8.
REPRESENTATIVE SPECIMENS: ALABAMA: Tallapoosa Co.,
Harper 3691 (GH, PH, US). GEORGIA: Richmond Co., August,
Cuthbert July 17, 1899 (FLAS, NY). NORTH CAROLINA: Gran-
ville Co.,-Oxford, Gillespie 394.(DUKE, FSU, NCSC). SOUTH
CAROLINA: Lancaster Co., Elgin, House 2568 (MO, NY, US).
Abundant on the sandy soil of scrub oak-longleaf pine
sandhills, dry woods borders, granite outcrops, and roadside
embankments along the Fall Line in the Carolinas; also
Richmond and Heard counties of Georgia and Chilton
(Harper, 1928) and Tallapoosa counties, Alabama. Flower-
ing early June through mid-July, occasionally to late August.
A low rounded or straggly growth form, fairly long and
narrow leaves, and short widened mature capsules with
tapering sides mark Hypericum lloydii. This distinctive
plant is related to the other members of the H. fasciculatum
complex but it is not known to grow in association with any
of them.
The range of this species is essentially isolated from other
members of the Hypericum fasciculatum group. In North
Carolina its range approaches that of H. reductum within a
county or two but plants of the two species have not been
discovered growing together. Nevertheless, I believe that
the taxonomy of the group can be best understood by recog-
nizing H. Ioydii as a species. Morphologically it is as dis-
tinct as any of the other members of this complex.
- 14. Hypericum exile P. Adams, sp. nov.
Frutex exilis, pauci-ramosus, usque ad 1 m altus vel raro
altior, usque ad 1 em in diametro. Cortex tenuis, in laciniis
vel segmentis parvis i i Folia basi
articulata, patentia, Gucariaebalela: lamina vix expansa,
valde et carnoso-marginata, margine siccitate maxime in-
voluta, (10-) 16-26 mm longa, 0.5-0.8 mm lata, apice longi-
acuminata, subtus obscure ; papillsts: remus axillaribus
haud conspicuis. L di-
chasis similibus in 3-6 nodis
compacta, paniculoidea. Sepala “basi sea lineari-
subulata, 6-7 mm longa, 0.5-0.8 mm lata, apice longiacumi-
nata, ante capsulae dehiscentiam decidua. Petala flava,
asymmetrica, 6-7 mm longa, 3-4 mm lata. Stamina 80-100,
5-7 mm longa. Gynoecium 4.5-7 mm longum, 3- (raro 2 vel
34 PRESTON ADAMS
4) carpellatum, placentatione parietale, cum 3 (raro 2 vel 4)
stylis. Capsula matura 6-7 mm longa, 1.5-2.5 mm lata.
Semina rufo-brunnea, carina minuta, 0.4-0.6 mm longa,
testa subtiliter reticulata.
Slender, sparingly-branched shrub, rarely over 1 m tall
and usually less than 1 em thick. Bark thin, exfoliating in
small irregular flakes or strips. Leaves articulate at base,
spreading, linear-subulate or needle-like, lamina poorly de-
veloped, edges thickened and turned abruptly downward
(inrolled on drying), (10-) 16-26 mm long, 0.5-0.8 mm wide,
apices long-acuminate, lower surface obscurely papillate,
axillary branchlets poorly developed. Shoots terminating
in a 3-7 flowered dichasium, usually with similar dichasia
from upper 3-6 nodes, inflorescence compact, panicle-like.
Sepals articulate at base, linear-subulate or needle-like, 6-7
mm long, 0.5-0.8 mm wide, apices long-acuminate, deciduous
before fruit maturity. Petals yellow, asymmetrical, 6-7 mm
long, 3-4 mm wide. Stamens 80-100, 5-7 mm long. Gynoeci-
um 4.5-7 mm long, carpels 3, rarely 2 or 4, placentation
parietal, styles 3, rarely 2 or 4. Mature capsules 6-7 mm
long, 1.5-2.5 mm wide. Seeds reddish-brown, raphe poorly
developed, keel minute, 0.4-0.6 mm long, 0.2-0.3 mm wide,
seed coat finely reticulate. Specific epithet from Latin mean-
ing “slender.”
TYPE: FLORIDA: Gulf Co., 2.5 miles east of Port St. Joe,
Adams 456 (Holotype, GH; isotypes, DUKE, FLAS, FSU, GA, K,
MO, NCSC, NCU, NY, SMU, US).
REPRESENTATIVE SPECIMENS: FLORIDA: Franklin Co., 10
miles west of Apalachicola, Adams 473 (FSU) ; Gulf Co., 3
miles east of Jort St. Joe, Adams 740 (FLAS, FSU, GA, GH,
SMU); Liberty Co., 6 miles north of Wilma, Adams 485
(FsU).
Known only in the sandy soil of open pinelands, Liberty,
Franklin, Gulf, Bay, and Washington counties, Florida.
Flowering throughout May.
Plants of Hypericum ezile, as the epithet suggests, are
, with a single stem which dichotomizes 25-40 cm
above the ground. The inflorescence is relatively narrow
and compact, with a panicle-like aspect. Leaf length, seem-
ingly long for such a small slender plant, is strikingly
reminiscent of H, lloydii. This hitherto undescribed species
STUDIES IN THE GUTTIFERAE. I. 35
occurs in close association with H. fasciculatum, H. chap-
manii, H. brachyphyllum, and H. nitidum but they do not
intergrade.
15. Hypericum myrtifolium Lam., Encye. Méth. Bot.
4:180. 1797. Not Spach, Hist. Nat. Vég. 5:399. 1836.
Hypericum rosmarinifolium Lam., Encye. Méth. Bot. 4:159.
1797.
Hypericum glaucum Michx., Fl. Bor.-Am, 2:78. 1803.
Hypericum sessiliflorum Willd. ex Spreng. Syst. Veg. 3 7346.
1826.
Myriandra glauca (Michx.) Spach, Hist. Nat. Vég. 5 :442.
1836.
TYPE: P-LA; photograph, GH.
REPRESENTATIVE SPECIMENS: FLORIDA: Duval Co., near
Jacksonville, Curtiss 265 (BKL, F, GH, NY, PH, US) ; Lake Co.,
Eustis, Nash 708 (A, MICH); Leon Co., 4 miles south of
Tallahassee, Adams 581 (FLAS, FSU). GEORGIA: 6 miles east
of Adel, Adams 47 (GA).
Sandy and peaty soil on the margins of evanescent ponds,
in moist pine flatwoods, grass-sedge bogs, and low roadside
ditches on the Coastal Plain of Georgia (nearly to the Fall
Line), most of peninsular Florida, and westward to Jackson
Co., Mississippi. Flowering mid-May through late July,
occasionally also in October.
This is the only species of Subsect. Centrosperma in which
the plants have clasping leaves. Other distinctive features
include the glaucous sepals, leaves, and young stems, gnarl
and woody caudex-like rootstock, moderate development of
a corky bark, subcoriaceous, ovate, elliptic-ovate, or cordate-
lanceolate leaves, and pyramidal-ovoid, strongly triquetrous,
black, glossy-shiny mature capsules. It is superficially simi-
lar to H. tetrapetalum (Subsect. Pseudobrathydium), both
having clasping leaves, and is often confused by the casual
observer. Besides its articulated leaves and sepals, H. myrti-
folium is readily distinguished from this species by a pen-
tamerous perianth, spongy-thickened bark, and compound
dichasial inflorescences.
Usage has firmly established the epithet Hypericum myrti-
folium over H. rosmarinifolium (both published by Lamarck
in his 1797 work).
Sect. MYRIANDRA, subsect. PSEUDOBRATHYDIUM Keller, in
36 PRESTON ADAMS
Engler & Prantl, Pflanzenfamilien, 3 (6-6a) 214. 1895,
excluding description.
Hypericum, Sect. Myriandra, Subsect. Brathydium (Spach)
Keller, 1. ¢., based on Brathydium Spach, Hist. Nat. Vég.
5:442. 1836, as a genus.
Hypericum, Sect. Myriandra, Subsect. Suturosperma Keller,
bie,
Hypericum, Sect. Isophyllum (Spach) Coulter, Bot. Gaz.
11:82. 1886, based on Isophyllum Spach, Hist. Nat. Vég.
52432. 1836, as a genus.
Ascyrum L., Gen. Pl. ed 5. 342. 1754 (excluding A. filicaule
Dyer), as a genus.
Hypericoides Adanson, Fam. Pl. 2:443. 1763, as a genus.
Crookea Small, Fl. Southeastern U. S. 786, 1335. 1903, asa
genus.
Shrubs and perennial herbs. Leaves and sepals without
an articulation or groove at the base (Fig. 2). Sepals 5 or 4,
persistent long after fruit maturity (deciduous in H. nudi-
florum). Petals 5 or 4. Withered stamens persistent long
after anthesis (except in H. nudiflorum and H. apocyni-
folium).
TYPE SPECIES: H. buckleyi M. A. Curtis.
Fifteen species distributed mostly in the central and
southern United States.
16. Hypericum nudiflorum Michx., Fl. Bor.-Am.
2:78. 1803
Hypericum mediflorwm Darby, Man. Bot., part 2, 35. 1841.
TYPE: “Hab. in Carolina. Goose Creek, Berkeley Co.,
[South Carolina ?],” P; photograph, GH.
REPRESENTATIVE SPECIMENS: ALABAMA: Etowah Co.,
Gadsden, Vasey 385 (F, GH, NY, PH, US). FLORIDA: Wakulla
Co., Godfrey 57135 (FSU, GH). GEORGIA: Elbert Co., 11.9
miles west of Elberton, Duncan 11699 (GA, NCSC). NORTH
CAROLINA: Ware Co., 3 miles south of Raleigh, Godfrey 4617
(DUKE, GH). TENNESSEE: Sequatchie Co., west of Dunlap,
Svenson 9554 (BKL, DUKE, MO, PH, TENN). VIRGINIA: Prin-
cess Anne Co., Macon’s Corner, Fernald & Long 4943 (A,
NY, US).
Locally abundant on stream banks and in rich deciduous
woods, southeastern Virginia, the inner Coastal Plain and
outer Piedmont in the Carolinas, the Piedmont of Georgia,
STUDIES IN THE GUTTIFERAE. I. 37
the Cumberland Plateau of Tennessee, the Panhandle of
Florida, and westward to southern Mississippi. Flowering
early June through late July, occasionally in late August.
Closely related to the following species.
17. Hypericum apocynifolium Small, Bull. Torrey Club
25:616. 1898
TYPE: TEXAS, Texarkana, A. A. Heller August 1897 (Lec-
totype, NY).
REPRESENTATIVE SPECIMENS: LOUISIANA: DeSoto Parish,
2 miles west of Hunter, Correll & Correll 10178 (DUKE, GH).
ARKANSAS: Drew Co., Monticello, Demaree 16231 (NY).
GEORGIA: Decatur Co., Butler’s Creek, Thorne & Davidson
17211 (GA).
Stream banks and moist woods, widely scattered localities,
southern Arkansas, western Louisiana, extreme northeast-
ern Texas, the Flint River drainage in extreme southwestern
Georgia, and the Apalachicola River bluffs in Gadsden Co.,
Florida. Flowering in June.
Vegetatively, plants of Hypericum apocynifolium cannot
be distinguished from those of H. nudiflorum. The differ-
ences listed in the key, i.e. simpler inflorescence, larger fruit,
and better-developed keel on the seed, are constant. Pending
acquisition of additional specimens and further study, I
think this taxon merits continued recognition as a species.
Svenson’s conception of this species included several
diverse elements (Svenson, 1940). One of the collections
which he cited (R. M. Harper 1755, NY) is referable to
H. frondosum Michx. A second collection (R. M. Harper
1501, NY, US) comprises plants of H. myrtifolium Lam.
Finally, the L pe (NY) cited ‘ac Small
in the original d tion of H. y mg to
that species and not to HE lobocarpieen Gattinger, as pees
believed.
18. Hypericum buckleyi M. A. Curtis,
Am. Jour. Sci. 44:80. 1843 [as Buckleii*]
TYPE: “In montibus Carolinae et Georgia,” S. B. Buckley
38 PRESTON ADAMS
(Lectotype, GH; isotype, NY, in less satisfactory condition).
REPRESENTATIVE SPECIMENS: GEORGIA: Rabun Co., Rabun
Bald, Pyron & McVaugh 895 (GA, US). NORTH CAROLINA:
Haywood Co., Mt. Pisgah, Bilt. Herb. no. 1319 (A, GH, MO,
NCU, US, WIS) ; Macon Co., Highlands, Bilt. Herb. no. 1319b
(F, MICH, NCSC, NY, PH). SOUTH CAROLINA: Greenville Co.,
1 mile from Caesar’s Head, Canby June 15, 1881 (A, F, MO,
NY, PH, US).
Endemic to the Blue Ridge Mountains of southwestern
North Carolina and adjacent Georgia at high elevations
(3,000 to 5,000 feet). Usually in seepage areas and moist
¢revices, sometimes along road embankments and ditches.
Among the North Carolina stations are Cold Mt., Mt. Pisgah,
Sassafrass Mt., Satulah Mt., Standing Indian Mt., Thomas
Bald, Wayah Bald, and Whitesides Mt. The only known
Georgia localities are Blood Mt., Brasst Bald, Hig
Bald, and Rabun Bald. Also near Caesar’s Head in Green-
ville Co., South Carolina. Flowering early June through
mid-July.
Hypericum buckleyi superficially resembles H. stragulum
in its decumbent growth form, elliptic to obovate leaves, and
a tendency for its flowers to be solitary. The former species
is readily distinguished by its pentamerous perianths, 3-
carpelled gynoecia, and much larger fruit.
19. Hypericum cistifolium Lam., Encyc. Méth. Bot.
4:158. 1797
Hypericum opacum T. & G., Fl. N. Am. 1:163. 1838.
Hypericum punctulosum Bertol., Misc. Bot. 13:18. 1853.
TYPE: P-LA; photograph, GH.
REPRESENTATIVE SPECIMENS: FLORIDA: Duval Co., near
Jacksonville, Curtiss 253 (F, GH, MO, PH, US) ; Franklin Co.,
Lanark, Gillespie G10-55-8 (DUKE, FSU, NCSC, SMU): Lee
Co., Ft. Myers, Hitchcock 12 (Ny,US). GEORGIA: Miller Co.,
1 mile west of Colquitt, Thorne 5873 (GA, GH, US) ; Sumter
Co., near Leslie, Harper 441 (A, BKL, F). LOUISIANA: Tan-
gipahoa Parish, 2 miles west of Robert, Correll 10511 (F,
GH, NY, PH). MISSISSIPPI: Harrison Co., Biloxi, Pollard 1002
(F, GH, MO, NY, US). NORTH CAROLINA: Brunswick Co.,
Godfrey 48428 (GA, SMU). SOUTH CAROLINA: Charleston Co.,
5 miles northwest of McClellanville, Godfrey & Tryon 1114
(DUKE, F, GH, MICH, MO, NY, PH, TENN).
STUDIES IN THE GUTTIFERAE. I. 39
Frequent in moist soil of pine flatwoods, seepage slopes,
grass-sedge bogs, margins of swamps and marshes, ditches,
and road embankments. Coastal Plain, North Carolina
tk 1 through peninsular Florida, and westward along
the coast of the Gulf of Mexico to eastern Louisiana. Flower-
ing early April, especially in southern Florida, through mid-
September, occasionally to mid-October.
Perhaps the most distinctive feature of this species is the
peculiarly lobed gynoecium. The carpel walls (between the
margins) are greatly indented or depressed, nearly meeting
in the center along the ovary axis to produce a trilocular
appearance in cross-section. In contrast, the lobed gynoecia
of other species in Sect. Myriandra are the result of indenta-
tion of the carpel margins (with their associated placentae)
so that a multilocular aspect is produced.
20. Hypericum sphaerocarpum Michx., Fl. Bor.-Am.
2:78. 1803
Brathydium sphaerocarpum (Michx.) Spach, Hist. Nat-
Vég. 5:444. 1836.
Brathydi h ium Spach, Hist. Nat Vég. 4:445.
1836.
STS S hy. Spach, Hist. Nat. Vég. 4:445.
P
1836.
Hypericum chamaenerium (Spach) C. Koch, Hort. Dendr.
66. 1853.
Hypericum turgidum Small, Fl. Southeastern U. S. 788.
1903.
Hypericum sphaerocarpum, var. turgidum (Small) Svenson,
Rhodora 42:17. 1940.
TYPE: “Hab. in Kentucky. Route de Louisville,” Michaux
(P; photograph, A; sketch from type by H. K. Svenson, GH).
REPRESENTATIVE SPECIMENS: ALABAMA: Morgan Co., Val-
hermoso Mt., Harper 39 (A, GH, NY, US) ; Madison Co. Canby
14 (NY, type of H. turgidum Small). ARKANSAS: Logan Co.,
Iltis 5344 (MICH, SMU). INDIANA: Jasper Co., Friesner
14557 (GA, MO). KANSAS: Osage Co., 3 miles south of Lyn-
don, Horr & Franklin E323 (FLAS, IND, NCSC). MISSOURI:
Oregon Co., near Thomasville, Palmer & Steyermark 41694
(A, MO, NY). TENNESSEE: Bedford Co., 5 miles northwest of
Shelbyville, Adams 71 (DUKE, F, FSU, 1A, IND, K, TEX, VPI).
Locally abundant on limestone outcrops, in cedar glades,
40 PRESTON ADAMS
rocky woods, prairie strips along railroads, and sandy
stream banks, from southern Ohio southward to the Black
Belt region of central Alabama, around the northern rim
of the b t to Arkansas
and eastern Oklahoma, northward to central Iowa, southern
Wisconsin, and northern Indiana. Flowering mid-June
through mid-August.
Closely related to Hypericum dolabriforme, this species
is readily distinguished by its nearly equal sepals, smaller
number of stamens, and larger seeds. The seed coats are
coarsely reticulate, with the transverse striae much better
developed than the longitudinal ones. Fewer than eight
seeds per capsule are produced by plants of H. sphaerocar-
pum as well as H. dolabriforme. All other members of the
section have capsules in which numerous seeds are matured.
Much variation in the mature capsule is present in this
species. Typically, the fruits are about as wide as long but
in some plants the width is greater than the length, pro-
ducing a depressed-globular form. In others, the capsules
are longer than wide. This variation does not appear to be
correlated with geography, judging from herbarium speci-
mens. Further investigation is necessary.
21. Hypericum dolabriforme Vent., Hort. Cels. 45
t. 45. 1800
Hypericum procumbens Desf. ex Willd., Sp. Pl. 3:1450. 1803.
Hypericum procumbens Michx., Fl. Bor.-Am. 2:81. 1803.
Brathydium grandiflorum Spach, Hist. Nat. Vég. 5:443.
1836.
Hypericum Bissellii Robinson, Rhodora 4:135. 1902. Type:
Bissell 4025 (GH).
TYPE: “Trouvée par Michaux sur les collines trés-arides
du Kentucky,” presumably the Delessert Herbarium, G.
RERESENTATIVE SPECIMENS: GEORGIA: Catoosa Co., 10
miles west of Ringgold, Cronquist 5614 (GA, GH, IND, MO,
NY, PH, SMU, US). TENNESSEE: Bledsoe Co., Pikeville, Sven-
son 9354 (BKL, DUKE, PH) ; Knox Co., Kearney July 9, 1894
(f, NCU). KENTUCKY: Wayne Co., Monticello, Smith &
se 4013 And GH, NY, US) ; Nelson Co., 12 miles south
McFarland 50 (IND, MO, NY, PH, TENN, WIS).
STUDIES IN THE GUTTIFERAE. I. 41
Abundant on limestone outcrops, in cedar glades, and dry
rocky beds of intermittent streams from north-central Ken-
tucky southward through eastern Tennessee to northwestern
Georgia. Flowering mid-June through mid-August, occa-
sionally in early September.
Grossly unequal sepals, a large number of stamens, small-
er seeds, and a finer seed coat reticulation are among the
characteristics which distinguish Hypericum dolabriforme
from H. sphaerocarpum, its closest relative. While these
two species occupy similar habitats and their geographic
ranges overlap, especially in Kentucky, they are not known
to grow in close association.
22. Hypericum adpressum Barton, Comp. Fl. Phila.
2:15. 1818
Hypericum bonaparteae Barton, Fl. N. Am. 3:95. 1823.
Hypericum fastigiatum Elliott, Sketches 2:31. 1821, not
HBK, Nov. Sp. & ge r 195. 1821.
Hypericum adp tigiati (Elliott) T. & G.,
FI. N. Am. 1:673. 1840, Type: Elliott (CHARL).
a fastigiatum (Elliott) C. Koch, Hort. Dendr. 66.
aaa, ipressum, var. i Robinson, Rhodora
4: 135-137. 1902. binge Kennedy et al 15 Sept. 1901 (GH).
Hypericum adp (Robinson) Fer-
nald, Rhodora 51: res "1949,
TYPE: Pennsylvania, “close to the Schuylkill, and not far
above Breck’s Island,” Barton (PH).
REPRESENTATIVE SPECIMENS: GEORGIA: Dougherty Co.,
west of Pretoria, Thorne 5709 (GA, IA). INDIANA: Jasper
Co., 2.5 miles southeast of Tefft, Deam 45934 (GH, IND).
MASSACHUSETTS: Barnstable Co., Flax Pond, Kennedy, Wil-
liams & Fernald 234 (BKL, F, MICH, NCSC, PH, TENN, US,
WIS). NORTH CAROLINA: Northampton Co., near Margaretts-
ville, Heller 1155 (GH, NY, PH). SOUTH CAROLINA: Jasper
Co., 1.7 miles south of Tillman, Ahles 15675 (NCU). TEN-
NESSEE: Coffee Co., south of Manchester, Svenson 8783
(DUKE, PH, WIS). VIRGINIA: Sussex Co., Stony Creek, Fer-
nald & Long 10727 (BKL, DUKE, GH, MO, US).
, pond margins, and wet roadside ditches at
widely scattered stations along the Atlantic coast from Cape
Cod and Nantucket, Massachusetts, southward through
42 PRESTON ADAMS
Long Island, New York, New Jersey, Delaware, southeastern
Virginia, northeastern North Carolina, the outer Coastal
Plain of South Carolina, Screven and Dougherty counties of
Georgia, middle Tennessee, southeastern Missouri, north-
eastern Illinois, and northwestern Indiana. This distribu-
tional pattern is peculiar and unlike any other species of
Hypericum in eastern North America. Flowering early July
through early September.
Hypericum adpressum is closely related to H. ellipticum
but differs in its much taller and more robust stems, stouter
rhizomes, larger leaves, and different placentation (i.e. the
placentae project inwardly much less). Spongy-thickened
stems, especially in the lower portion, occur on plants grow-
ing in standing water. This condition is obviously due to
the direct effects of the environment (Bicknell, 1913; Sven-
son, 1940).
23. Hypericum ellipticum Hooker, Fl. Bor.-Am. 1:110. 1830
Hypericum ellipticum, forma submersum Fassett, Rhodora
41 :376. 1939.
Hypericum ellipticum, forma foliosum Marie-Victorin, Nat.
Canad. 71:201. 1944.
TYPE: Canada, Mr. Cleghorn (Lectotype, K, not seen).
The Todd and the Richardson specimens also cited by Hooker
(1830) have not been located.
‘ATIVE SPECIMENS: MARYLAND: Garrett Co.,
Mountain Lake Park, Steele 56 (GH, US). MICHIGAN: Chip-
pewa Co., Sugar Island, MceVaugh 8751 (MICH, NCSC). NEW
HAMPSHIRE: Cheshire Co., Alstead, Fernald 338 (F, GH, NY,
PH, US). NEWFOUNDLAND: Fernald & Wiegand 5843 (GH,
NY). QUEBEC: Algoma District, Taylor et al 1349 (GH)-
NOVA SCOTIA: Yarmouth Co., Kemptville, Fernald & Linder
21857 (GH, MO, PH, Us).
on pond and lake shores, stream banks, mead-
ows, river flats and sand bars, and swamps, eastern New-
foundland, Nova Scotia, New England, southwestern
Pennsylvania, West Virginia, northeastern Tennessee, the
Lake Superior region of Wisconsin, Minnesota, and Michi-
gan, and the Georgian Bay area of Ontario. Flowering late
June through early September.
This species is closely related to Hyperi lp
Plants of both taxa have prominent rhizomes, a feature
STUDIES IN THE GUTTIFERAE. I. 43
absent in all other members of Sect. Myriandra. Floral
morphology and the size, color, and testa marking of the
seeds are similar also. Each species is easily recognizable
by leaf shape and size, placentation, and growth habit.
Tetramerous perianths, red or purplish petals, leaves, and
stems, and peculiar submersed aquatic forms with simple
stems and round to ovate “feather-veined” leaves (forma
submersum Fassett; type: Fassett 19172 WIS) are seen
occasionally. Following fruit maturity the axillary branch-
lets may resume growth, even overtopping the infructescence
(forma foliosum Marie-Victorin ; type: Marie-Victorin et al
56602, MT, isotypes, F, GH, MO, PH).
24. Hyperi i lum (T. & G.) Gray ex
S. Watson, Biblio. Index to N. Am. botany 1:456. 1878
Based on Ascyrum microsepalum T. & G., Fl. N. Am.
1:157. 1838.
dy dii Spach, Hist. Nat. Vég. 5:432. 1836.
Not H. drummondii (Grev. & Hook.) T. & G., FL N. Am.
1:165. 1838, which is based upon Sarothra drummondii
Grev. & Hook., Bot. Misc. 3:236. 1833.
Crookea microsepalum (T. & G.) Small, Fl. Southeastern
U. S. 786, 1335. 1903.
TYPE: Florida, Apalachicola, Drummond (GH).
REPRESENTATIVE SPECIMENS: FLORIDA: Franklin Co.,
Adams 125 (FSU); Jefferson Co., Godfrey 60629 (FSU) 5
Taylor Co., Adams 807 (FLAS, FSU, GH, K, smu). Without
exact locality, Dr. Alexander (NY).
Abundant in low pine flatwoods of the Florida Panhandle
from Taylor to Bay counties and northward into southern
Georgia (Calhoun and Atkinson counties). Flowering late
February through late April, often sporadically in May and
November. i
The features which mark this distinctive species include
the following: a very early spring flowering time, four
nearly equal sepals, four large showy petals, 2 : ate
gynoecium, and parietal placentation. In addition, a tend-
ency exists for some of the flowers on many to be
pentamerous, with five nearly equal sepals and five petals.
Not infrequently other flowers (on these same plants) will
show tetramery in their calyces and pentamery In their
corollas. The reverse situation is often encountered also.
44 PRESTON ADAMS
This phenomenon is general throughout the range of the
species and can be considered another distinguishing feature.
This unstable floral situation could be interpreted as indi-
cating that the species is und i luti y transition
from the presumably primitive pentamerous condition to
the more advanced 4-parted one.
25. Hypericum stans (Michx.) Adams & Robson,
Rhodora 63:15. 1961
Based on Ascyrum stans Michx., Fl. Bor.-Am. 2:77. 1803.
Ascyrum cuneifolium Chapman, Fl. Southeastern U. S. ed.
2, suppl. 2. 680. 1892. Type: Chapman 1835 (NY).
TYPE: “Hab. in Va.” (Presumably at P).
REPRESENTATIVE SPECIMENS : FLORIDA: Columbia Co., Lake
City, Nash 2489 (F, FLAS, GH, MICH, NCU, 0S). GEORGIA:
Douglas Co., 2 miles east of Villa Rica, Cronquist 5559 (GA,
NO, NY, PH, SMU, US). KENTUCKY: McCreary Co., McFar-
land & James 48 (DUKE, IND, MO, PENN, TENN, WVA).
MISSISSIPPI: Jackson Co., Ocean Springs, Seymour 75 (NCU,
SMU, TEX). VIRGINIA: Princess Anne Co., near Virginia
Beach, Heller 1268 (F, GH, NY, PENN, PH, US).
Dry to moist flatwoods, bogs, meadows, bottomlands, road-
side ditches, and shores of ponds and lakes. Long Island,
New York, southward through New Jersey, eastern Vir-
ginia, the Carolinas, to central Florida, westward to eastern
Texas and central Arkansas. Also on the Cumberland
Plateau through Tennessee into McCreary and Laurel coun-
ties of Kentucky (see Adams, 1957, for distribution map).
Flowering late July through mid-October, occasionally in
June and November.
This species is most closely related to Hypericum edi-
Somanum, endemic in south peninsula Florida. The distin-
guishing features are those presented in the key and further
discussed in the treatment of that species (which see).
26. Hyperi disoni (Small) Adams & Robson,
Rhodora 63:15. 1961
Based on Ascyrum edisonianum Small, Man. Southeastern
868. 1933.
Fl
TYPE : “21 miles east of Arcadia, Florida,” Hand 118 (NY).
REPRESENTATIVE SPECIMENS: FLORIDA: Highlands Co., 21
_ miles east of Arcadia, Adams & Testasecca 100 (DUKE, F,
STUDIES IN THE GUTTIFERAE. I. 45
FLAS, FSU, GA, MICH, MO, NCSC, NY, PH); same locality,
Adams 140 (GH) ; Glades Co., Fisheating Creek, Brass 14824
(GH).
Locally abundant on sandy soil of low open prairies, lake
and pond shores, and roadside ditches, Highlands, Glades,
and DeSoto counties, Florida.. The known-stations are in
central and southern Highlands County from Lake Josephine
southward to the Venus area and westward along the
Arcadia highway (Fla. 70) to about the DeSoto County line.
Also collected along Fisheating Creek in adjacent Glades
County. Flowering probably throughout the year.
Closely related to Hypericum stans, this species may be
easily recognized by the characteristics listed in the key.
Other features include a much better developed laticifer
system in the bark of H. edisonianum (especially noticeable
on older stems) and the larger and more numerous resin
dots in the leaves of H. stans.
Dense thickets, often 1.5 m in height, are commonly pro-
duced by plants of Hypericum edisonianum. A contributing
factor is the tendency for numerous vegetative shoots to be
produced at frequent intervals along horizontal roots of the
plants. This form of reproduction, present in several other
species of the section, is not known to occur in the closely
related H. stans.
The range of Hypericum edisonianum appears to be essen-
tially allopatric from that of its relative H. stans. The
closest known stations of the latter species are at least
thirty-five miles to the south (Immokalee in Collier County)
and southwest (Ft. Myers in Lee County). Northward the
nearest collection of H. stans was made at Haines City in
Polk County, some sixty miles distance.
27. Hyperi t talum Lam., Encye. Méth. Bot.
4:153. 1797
Ascyrum tetrapetalum (Lam.) Vail in Small, Fl. Southeast-
903.
TYPE: “Hab. in Florida.” P-LA; photograph, GH.
REPRESENTATIVE SPECIMENS: FLORIDA: Clay Co., 2 miles
north of Orange Park, Moldenke 160 (DUKE, MO, NY, PENN,
us) ; Dade Co., Small, Mosier & Small 6703 (FLAS, GH, TENN,
wvA) ; Duval Co., near Jacksonville, Curtiss 245 (F, PH, US);
Lake Co., Eustis, Nash 1977 (DAO, MICH, NCU, PH). GEORGIA:
46 PRESTON ADAMS
Irwin Co., 4 miles north of Ocilla, Wilbur 3074 (FSU, NCSC,
SMU) ; McIntosh Co., Sapelo Island, Duncan 17970 (GA).
Sandy soil of pine flatwoods, peaty lake and pond margins,
marshes, and wet roadside ditches, on the outer Coastal Plain
of southeastern Georgia and peninsular Florida; Pinar del
Rio, Cuba. Flowering late January (in southern Florida)
through late April, ionally sy lically in July, August,
and September.
This species is distinguished from the related Hypericum
stans by ovate-cordate, clasping leaves which are similar to
the sepals and by the dichotomous branching pattern. From
the superficially similar H. myrtifolium this species can be
readily separated by its tetramerous perianth, terete cap-
sules, solitary flowers, and thin, non-corky bark.
28. Hi i
YP
hyperi (L.) Crantz, Institut. rei
herbariae 2:520. 1766
Based on Ascyrum hypericoides L., Sp. Pl. 2:788. 1753.
See Adams (1957) for complete synonymy.
TYPE: “Hab. in Jamaica,” Patrick Browne (LINN, speci-
men no. 944.2, designated neotype by Adams in 1957; photo-
graph, GH).
REPRESENTATIVE SPECIMENS: ARKANSAS: Ashley Co.,
Parkdale, Demaree 16398 (DAO, NY, 0S, PENN, TENN, TEX).
FLORIDA: Lake Co., Eustis, Nash 1609 (F, GH, MICH, MO, NY,
swamp margins, and hardwood slopes. New Jersey south-
ward through peninsular Florida, westward to southeastern
Missouri, eastern Oklahoma, and eastern Texas ; the eastern
escarpment in Mexico from Nuevo Le6n and Tamaulipas
or
Hyp
erieum
Left, Hypericum stragulum Adams &
Robson
(Aseyrum multicaule Michx.
one plant,
hypericoides (L.) Crantz (Aseyrum hypericoides L.), a single plant with one main
wround level. Hall, Co,, Ga., ca, 8 miles west
f Flowery Branch,
(Photograph by courtesy of Dr
branche
stem. Bra
Wilbur
decumbent
n
H, Duncan.)
ching well abs
NI SaICGALS
‘I ‘SVYSAILLAD AHL
48 PRESTON ADAMS
southward to Chiapas and the mountains of Guatemala and
Honduras; Bermuda, the larger Bahamas, Cuba, Jamaica,
Hispaniola, and Puerto Rico (see Adams, 1957, for distribu-
tion map). Flowering late June through late October (at
least in the United States).
Geographically the most widespread and occupying the
most diverse range of habitats, plants of Hypericum hyperi-
coides nonetheless retain the facies characteristic of the
species. Considerable variation is present in the leaf and
sepal dimensions but this appears to be simply a response
to habitat differences (Adams, 1957). An erect bushy
growth form, two large outer sepals and two minute inner
ones, four pale yellow, nearly symmetrical, early-falling
petals, and a 2-carpelled gynoecium distinguish this species.
Its closest relative is H. stragulum (Ascyrum multicaule
Michx.), the erect growth form being the principal discern-
ible difference (see Fig. 3).
29. Hypericum stragulum Adams & Robson, Rhodora
63:15. 1961
Based on Ascyrum multicaule Michx., Fl. Bor.-Am. 2:77.
1803, not Hypericum multicaule Lam., Encyc. Méth. Bot.
4:178. See Adams (1957) for complete synonymy.
TYPE: “Hab. in Va., Carolina.” P; photograph, GH.
REPRESENTATIVE SPECIMENS : ARKANSAS: Cross Co., Crow-
ley’s Ridge, Demaree 19579 (GH, MO, OKL, TENN). ILLINOIS:
Pope Co., Golconda, Palmer 16698 (NY, PH). KANSAS:
Cherokee Co., Hitchcock 1012 (MICH, US). MISSOURI: Ozark
Co., Tecumseh, Palmer 32882 (GH, MO). NORTH CAROLINA:
Jackson Co., Godfrey & O’Connell 51631 (DUKE, NCSC).
TENNESSEE: Knox Co., Knoxville, Ruth July 1895 (F, GH,
MO). TEXAS: Henderson Co., Eustace, Lundell & Lundell
9574 (GH, MICH, NY, SMU). WEST VIRGINIA: Cabell Co.,
Gilbert 548 (DUKE, F, PENN, PH, SMU, TENN, WVA).
Adams, 1957, for distribution map). Flowering early July
through early August.
STUDIES IN THE GUTTIFERAE. I. 49
A decumbent, low, matted, rounded growth form distin-
guishes Hypericum stragulum from the closely related H.
hypericoides. The two species often grow in close association
but each is readily identifiable (see Fig. 3). The decumbent
habit appears in the seedlings of H. stragulum. The erect
seedling soon loses its upright position and becomes nearly
prostrate. One or more axillary branchlets, usually close to
the base of the stem, begin growth and become almost pros-
trate. However, in young plants of H. hypericoides the stem
remains erect, eventually producing an erect habit unlike
that of H. stragulum. These two taxa require further exten-
sive study.
30. Hyperi ti Adams & Robson,
Rhodora 63:15. 1961
Based on Ascyrum pumilum Michx., Fl. Bor.-Am. 2:77.
1803, not Hypericum pumillum Sessé & Moc., Fl. Mexic. ed.
2, 177. 1894,
Ascyrum pauciflorum Nutt., Gen. 2:15. 1818. Not Hyperi-
cum pauciflorum HBK, Nov. Gen. 5:192. 1822.
TYPE: ‘Hab. in Georgia.” Presumably at P.
REPRESENTATIVE SPECIMENS : FLORIDA: Columbia Co., Lake
City, Nash 2211 (GH, MICH, MO, NY, US) ; Duval Co., near
Jacksonville, Curtiss 246 (F, FLAS, PH). GEORGIA: Brooks
Co., 3 miles northeast of Pavo, Adams 37 (GA). MISSISSIPPI:
Farrisoi Co., Biloxi, Tracy 4489 (F, MICH, MO, NY, 0S, US).
SOUTH CAROLINA: Colleton Co., 11 miles northwest of Wal-
terboro, Adams 84 (FSU).
Dry sandy soil of pinelands on the Coastal Plain of south-
ern Georgia, northern and western Florida, southern Ala-
bama, and southern Mississippi. Also St. Tammany Parish,
Louisians. Bladen Co., North Carolina, and Berkeley and
Beaufort counties of South Carolina (see Adams, 1957, for
— map). Flowering mid-March through mid-
une.
Morphologically, plants of this species exhibit more reduc-
tion than any other member of Sect. Myriandra. The low,
inconspicuous habit, early-falling petals, obsolete or nearly
absent inner pair of sepals, bicarpellate gynoecia, and rela-
tively small leaves are all interpreted as evidences of extreme
reduction. Although morphologically a well-isolated species,
Hypericum suffruticosum is related to H. stragulum (As-
cyrum multicaule) and, to a lesser degree, H. hypericoides.
50 PRESTON ADAMS
The elongate pedicels which become reflexed soon after an-
thesis and the position of the bractlets at the pedicel base
are among the features which serve to distinguish H. suf-
fruticosum from these two species.
Dusi0us AND REJECTED EPITHETS
Hypericum fulgidum Raf., Fl. Ludov. 88. 1817, description; Herb.
Raf. 55. 1833, name. This epithet may refer to H. fasciculatum L:
but the description is insufficient for a definite determination.
Hypericum rostratum Raf., Fl. Ludov. 88. 1817. Rafinesque furn-
ished this name and a Latin description for a H. ypericum which Robin
(1807) had seen on his travels in Louisiana. These two descriptions
Suggest a plant similar to H. lobocarpum Gattinger. Since there is
no type specimen and since Rafinesque received his information in-
Myriandra galioides (Lam.) Spach, Hist. Nat. Vég. 5:437. 1836.
Torrey & Gray (1840) listed this name as a synonym of H. galioides
Lam. The description is too vague for a definite determination.
Myriandra ledifolia Spach, Hist. Nat. Vég. 5:441. 1836. Cited by
Coulter (1897) as a synonym of Hypericum prolificum L. but the
original description is hardly sufficient to make an exact determination.
LITERATURE CITED
ApaMs, W. P. 1957. A revision of the genus Ascyrum (Hyperica-
ceae). Rhodora 59:74-95.
aes | The taxonomy of Hypericum section Myrian-
dra (Hypericaceae). Unpub. Ph.D. Thesis, Harvard University.
. The status of Hypericum prolificum. Rhodora
61:250-251.
W. P. anD N. K. B. Rosson. 1961. A re-evaluation of the
generic status of Ascyrum and Crookea (Guttiferae). Rhodora
E. P. 1913. The ferns and flowering plants of Nantucket.
Bull. Torrey Club 40:609.
CHapMaN, A. W. 1897. Hypericaceae. Flora of the southern United
States. C: i
Courrer, J. M. 1886. Revision of the North American Hyperica-
» eeae. Bot. Gaz. 11:78-88.
——————. 1897. Hypericaceae in Gray, Synoptical Flora of
North America. Vol. 1, part 1, 282-297.
TAXONOMIC FERN NOTES. II. 51
Euuott, S. A sketch of the Botany of South Carolina and Georgia.
2:21-35, J. R. Schenck. Charleston.
FERNALD, M. L. 1950. Guttiferae in Gray’s Manual of Botany. Ed.
8. American Book Co. 1008-1015. New York.
FERNALD, M. L. AND B. G. SCHUBERT. 1948. Studies of American
types in British Herbaria. Part II. Some Linnaean Species.
Rhodora 50:167-1
GILLESPIE, J. P. 1959. "The Hypericaceae of Tennessee. Castanea
24 S 32,
GLEAsON, H. A. 1952. Hypericaceae in the New Britton and Brown
pendiorae Flora. 2:538-544. New York.
, R. M. . Hypericaceae. Economie Botany of Alabama.
II. Geological Survey of Alabama, Monograph 9. 270-273.
Hoar, C. S. anp E. J. Haertt. 1932. Meiosis in the genus Hyperi-
cum. Bot. Gaz. 93:197-204,
KELLER, Ropert. 1925. Ascyrum and Hypericum in Engler and
Prantl, Pflanzenfamilien 2 Aufl. 21:154-234.
LanJouw, J. & F. A. STAFLEU. rs Index Herbariorum, Part I.
The Herbaria of the World. Ed.
Lott, H. J. 1938. Nomenclatural ee on Hypericum. Jour. Arnold
Arb. 19: serine 279-281.
McLavucuiin, W. 1931. Preliminary reports on the flora of
Wisconsin. ae Hypericaceae. Trans. Wisc. Acad. Sci., Arts,
and Lett. 26:281-288.
MOHLENBROCK, R. H. AND J. W. Vorct. 1959. A flora of southern
Illinois. Southern Illinois University Press. Carbondale.
NieEtson, N. 1924. Chromosome numbers in the genus Hypericum.
Hereditas 5:378-382.
PursH, F. 1814. Flora Americae Septentrionalis. 2:373-379. White,
Cochrane and Co.
REHDER, A. 1940. Hypericaceae. Manual of Cultivated Trees and
Rosin, C. C. 1807. Flore Louisianaise. Vog. Louis.
1957. The genus is pera in Africa south of
the Sahara, Madasncene: and the Mascarenes. Kew Bull. No. 3,
433-446.
Sma, J. K. 1903. Hypericaceae. Flora of the southeastern United
States. 784-791. New York.
Hypericaceae. Flora of the southeastern United
1913.
States. 784-791. New York.
933. Manual of the southeastern
66-875.
Svenson, H. K. 1940. Plants of the southern United States. II.
Woody species of Hypericum. Rhodora 42:8-19.
1952. What is Hypericum prolificum? Rhodora
54:205-207.
Torrey, J. anp AsA Gray. 1838. Hypericaceae. Flora of North
rica. 1:155-169.
TAXONOMIC FERN NOTES. IL.
PITYROGRAMMA (INCLUDING TRISMERIA) AND ANOGRAMMA
ROLLA TRYON
During studies in Pityrogramma, Trismeria and Anog-
ramma' in prep. of treat of these genera as
they may occur in Peru, various matters pertaining to other
species, as well as the Peruvian ones, have been assembled.
These seem better presented in a separate paper.
Pityrogramma, Trismeria and Anogramma are all closely
related and have been usually maintained as distinct genera.
Each of these is adequately separated from allied genera
which are Coniog 2, G pteris, Bommeria, Hemi-
onitis and Trachypteris (Christ *) and, in addition,
Saffordia, Paraceterach, Cerosora and Pleurosoriopsis
(Copeland’). My conclusions are to unite Trismeria with
Pityrogramma but to maintain Anogramma as distinct
from it.
Throughout this paper frequent reference is made to two
publications by Karel Domin: “Generis Pityrogramma
(Link) species ac sectiones in clavem analyticam dispositae”,
Publ. Fae. Sci. Univ. Charles, no. 88. 1928 and “The hybrids
and garden forms of the genus Pityrogramma (Link)”,
Rozpr. II. Tr. Ceské Akad. 38*. 1929, For the sake of brevity
these will be referred to as Domin, 1928 and Domin, 1929
respectively.
I. THE GENUS PITYROGRAMMA
Pityrogramma proper consists of some 11 species and
closely related to these are three species that evidently rep-
resent three isolated and specialized derivatives from it,
or from a common ancestor. The species most “typical” of
Pityrogramma are P. chrysophylla, P. chrysoconia, P. de-
albata, P. “Eggersii”, P. sulphurea, P. tartarea and P. calo-
melanos. These all have concolorous rhizome scales (or if
darker and lighter areas are Present there is no well defined
central stripe) ; clustered petioles with a groove on their
upper side and with two (or in large leaves more) roundish,
pes
‘Supported, in bart, by a grant from the National Science Foundation (NSF-G1064) .
a a
TAXONOMIC FERN NOTES. II. 53
oval to C-shaped vascular bundles; fertile and sterile leaves
that are monomorphic or nearly so; a more than 1-pinnate
lamina; stalked pinnae; ceraceous indument on the lamina
(some variants of P. tartarea, P. calomelanos and P. “Eg-
gersii” are pubescent or glabrous) and sometimes also on
the rhizome, especially at its apex; the veins of the ultimate
segments arising from a single basal vascular branch; and
distinctive spores (Fig. 2c) that are tetrahedral-globose (or
somewhat flattened) and have the exospore with rather
broad and dark ridges on a lighter background, the ridges
forming prominent rings at the equator and mostly “hiero-
glyphs” or dots elsewhere (one variant of P. calomelanos
has nearly smooth spores).
To these species must be added P. argentea, P. Humbertii,
P. ferruginea and P. schizophylla. These are clearly closely
related to the species of the “central group” although they
differ in some ways from them. P. argentea, for example,
has spores with the whole exospore dark and with fewer
ridges and somewhat tuberculate. (I have seen no material
of P. Humbertii but it is evidently closely related to P.
argentea). Pityrogramma schizophylla has the lamina
glabrous and bears proliferous buds, and P. ferruginea has
the whole lamina densely and persistently lanate. The char-
acters by which these species depart from those of the
“central group” are not striking and together they all form
a relatively uniform genus.
In contrast, there are the three specialized species previ-
ously mentioned. These also have ceraceous indument on the
lamina and two vascular bundles (at least in large leaves)
near the base of the petiole. They each depart, however, in
several characters from the species of Pityrogramma previ-
ously mentioned.
Pityrogramma triangularis has the petiole terete, the
pinnae are sessile or adnate and the spores are light colored,
the exospore has a low, fine and sharply reticulate surface.
The rhizome scales have marked light margins beyond the
dark sclerotic central stripe.
Gymnogramma Lehmannii has the spores, grooved petiole
and rachis and two vascular bundles typical of Pityro-
gramma. However, the lamina is only pinnatisect (Fig. 6a)
and the ultimate segments (pinna-segments) are furnished
54 ROLLA TRYON
with many veins arising from the costa (Fig. 6b), the
petioles are spaced on the rhizome and the vascular bundles
near the base of the petiole are elongate S-shaped.
Trismeria trifoliata has the two vascular bundles typical
of Pityrogramma; they are, however, above the basal por-
tion of the petiole (Fig. 3b), rather C-shaped but with the
back of the C curved forward. The lamina architecture is
somewhat different, the fertile and sterile leaves are subdi-
morphic and the spores are light colored with a nearly
smooth exospore. These spores are quite similar to those
present in one variety of P. calomelanos (var. ochracea).
A relation of this species and Pityrogramma is shown by the
4 hybrids (discussed below) between them.
Comparing each of these three species, in their similar-
ities and diff; , with Pityrog it is evident that
they represent close relatives of that genus. Two courses are
possible for their classification: each may be placed in its
own genus or they may all be placed in Pityrogramma. The
former course would require the recognition of two new
genera (one for P. triangularis and one for Gymnogramma
Lehmannii) and this classification would emphasize the dif-
ferences between them and Pityrogramma. However, the
imilarities must be idered equally and these are more
than sufficient to place them all in Pityrogramma. Fur-
ther, if this is done, the diversity of that genus is increased
primarily in characters of the lamina architecture and
venation, the attachment of the pinnae, and of the shape
of the petiole and the vascular bundles. Such diversity can
well be accommodated in a single genus, especially when the
divergent charact ot lated in a group of species.
Accordingly, I am treating all of these species in Pityro-
gramma.
PITYROGRAMMA Link, Handb. Gewiichse, 3:19. 1833
Type species: P. chrysophylla (Sw.) Link (Acrostichum
chrysophyllum Sw.).
: Ceropteris Link, Fil. Sp. Cult. 141. 1841, nom. superfi.
illegit. Type species: the same as that of Pityrogramma.
Trismeria Fée, Mém. Fam. Foug. 5 (Gen. Fil.) : 164,
t. 14, f. 1, 2. 1852. Type species: Pityrogramma trifoliata
(L.) Tryon (T. aurea Fée, Op. Cit. 165, nom. superfi. illegit.
TAXONOMIC FERN NOTES. I. 55.
= Acrostichum trifoliatum L.). Copeland’s choicet of T.
argentea Fée as the type is surely not as suitable as that
of T. aurea by Maxon’, for the latter was not only more
ts the
fully illustrated by Fée, but Pp
— well known Acrostichum trifoliatum L.
na section Isgnog Hieron. Engl. Bot.
Jaded, 34: 2474, 1904. Type species: G. Lehmannii Hieron.
AMERICAN SPECIES OF PITYROGRAMMA
From the key and notes which follow it will be evident
that I take a very different view of the species of Pityro-
gramma than did Domin.* Specifically, I recognize, among
the American species sufficiently studied, 10 species where
Domin recognized 24. Some comments on the reasons for
this treatment will be offered.
The species I recognize are based primarily on characters
of the scales of the rhizome and petiole base, the color of the
petiole and rachis, the relative development of the two sides
of the pinna, the angle of the pinnules to the pinna rachis,
the shape of the ultimate segments, the margin and relation
of the veins ends to it and the spores. Other characters are
the densely lanate lamina ( in P. ferruginea) and the pres-
ence of proliferous buds on the lamina (in P. schizophyl’a).
The complexity of the lamina is used as a supporting char-
acter in P. dealbata, as is the shape of the lamina in some
species, and lack of indument in P. schizophylla. Indument
color, although sometimes mentioned in the key where it
may be useful, is not used to define the species. The use
of these characters defines species which are consistent with
our knowledge of the variability of these plants; and they
have ranges (either contiguous or disjunct) which are of
a pattern known in other species.
Domin used most of the kinds of characters previously
mentioned (although not always the same ones) and also
employed a considerable number of others, such as type of
indument, color of the ceraceous indument, disposition of
the pinnae and their shape, texture, size of segments and
of the leaf, revolute or plane margin, spacing of pinnules,
“Gen. Fil, 76. 1947.
®Pterid. in Sei. Surv. Porto Rieo & Virg. Isls. 6: 436. 1926.
*Domin, 1928.
56 ROLLA TRYON
toothing of segments, and relative length of petiole. Some
of these are subject to variation in plants of different sizes,
and others to variation under different environmental condi-
tions, correlations which may be seen in large suites of speci-
mens. Others, while evidently more stable, nevertheless do
vary in some species to an extent that they can not be relied
upon.
The type of indument on the undersurface of the lamina
and its color, if ceraceous, has been so overemphasized that
some additional discussion of this character is desirable.
The color of the ceraceous deposit on the under surface
of the lamina may be of little or no taxonomic importance.
It is true that some species such as P. dealbata and P. sul-
phurea are known in only one color form, but most others
have two or more colors, That the color may easily vary is
suggested by reports in the literature of a plant of one color
giving rise, by spores, to plants with another color. These
reports, however reasonable, must be subject to reservation
because of the possibilities of contamination of a “culture”
by spores of other kinds of Pityrogramma. There are some
reports of two colors on a single plant. For example T.
Smith’ discusses a plant of P. tartarea with leaves with
white wax and one leaf with golden wax. Hooker® reports
a leaf of P. calomelanos with one pinna half white and half
yellow. A specimen of P. calomelanos (Colombia, Daniel
292, US) has the indument on two leaves light yellow-orange
and on one it is roseate.
Wax color forms are present but not recognized in the
following species or varieties where the color is not cor-
related with any other character nor with geography: P.
lomel var. calomel: (white, lemon yellow, pale
roseate), P. tartarea var. tartarea (white, cream yellow),
P. chrysoconia (white, yellow), P. chrysophylla (white,
bright yellow), P. trifoliata (white, yellow) and P. “Eg-
gersit” (white, rose).
Some species that most commonly have ceraceous indu-
ment on the under surface of the lamina may be sometimes
glabrous or pubescent. Species that have pubescent variants
are P. trifoliata, P. tartarea and P. calomelanos. The latter
rather rarely may be glabrous.
“Gard. Chron. ns. 8: 599. 1877.
‘Gard. Ferns, t. 50, f. 3. 18962.
TAXONOMIC FERN NOTES. II. 57
That the presence and type of indument on the under
surface of the lamina may be a “superficial” character is
demonstrated by a plant of P. calomelanos from Peru.’ This
plant has 10 leaves with the following characters: the three
oldest leaves are glabrous, the next one is sparsely ceraceous
and the next two are white ceraceous, the following two are
pubescent and the youngest two are white ceraceous.
The type and color of indument, nevertheless, are used as
the sole characters for the varieties of P. calomelanos and
of P. tartarea. In these species the strong correlation of
the character with geography seems to provide it with an
importance it would otherwise not have. Even so, with fur-
ther knowledge, it may not be reasonable to maintain such
a treatment.
The extent to which hybridization among the species
occurs in nature is not known. Hybrids between P. trifoliata
and P. calomelanos and P. ferruginea are proposed in this
paper. It is perhaps significant that hybrids have been rec-
ognized only with P. trifoliata which is an especially distinc-
tive species in its lamina architecture. It is quite possible
that the difficulties of definition and identification that exist
between P. calomelanos and P. tartarea, and some other
species, may be due to hybridization. However, the rather
few “intermediate” plants may as well be ecological or
genetic variants of either species. Where two species grow
together or in close proximity, the presence or absence of
intermediate plants should be confirmed.
KEY TO THE SPECIES OF PITYROGRAMMA
(Four insufficiently known taxa, discussed at the end of the next
section, have not been keyed out.)
a. Petiole terete; rhizome scales with a dark, sclerotic central stripe
and narrow, thinner, pale margins. (Pinnae, except sometimes the
basal pair, adnate to the rachis, the basal ones enlarged.)...... ra
be J
a. Petiole grooved on the upper side; rhizome scales wholly sclerotic to
subsclerotic, concolorous or often with a dark basal or central
area. b.
b. Lamina pinnatisect; pinna-segments entire, about as broad as
long, broadly obtuse, with many veins arising from the costa of
the lamina. 8. P. Lehmannii.
b. Lamina 1-pinnate or more complex; pinnae elongate; veins of the
ultimate segments arising from a single basal vascular branch. c.
presen Sra eae
*Tingo Maria, Tryon & Tryon 5242 (GH).
58
ROLLA TRYON
ce. Vascular bundles of the petiole (at about 1/3 the distance to
the lamina) C-shaped with the back of the C curved forward. ....
1. P. trifoliata and hybrids of it.
The species has the pinnae entire, especially the apical ones, to
commonly 'rifoliolate below, or a few to rarely many 5- (very
rarely 7-) foliolate. The hybrids have the pinnae with many
segments.
Vascular bundles of the petiole (at about 1/3rd the distance to
the lamina) roundish, oval to C-shaped; pinnae with numerous,
innately arranged totes or segments. d.
d, Ultimate segments mostly shallowly (if at all) toothed,
pinnately many veined (with a strong midvein), broadest at
the base to lanceolate or elliptical; or rarely lacking a mid-
pag (or with a poorly developed one) and obovate to oval. e.
e. Primary rachis and secondary rachises densely and persis-
tently lanate, the petiole similar but sometimes deciduously
lanate. 6. P. ferruginea.
e. Petiole, primary rachis and secondary rachises glabrous,
ceraceous or thinly and irregularly pubescent. f.
f. Scales of the apex of the rhizome and the base of the
petiole with a long portion one cell wide below the
terminal cell. (Spores ated ridged, the ridges with
sharply defined margins
g. Small leaves a pidaaaapeiniahd, or bipinnate-lobed
only at the very base, large ones to tripinnate; ultimate
segments oblong, deltoid, lanceolate, elliptical to long-
triangular. h. i
h. Pinnae equilateral; pinnules ascending, those on the
basiscopic side more strongly so than those on the
acroscopic side; scales of the apex of the rhizome
and the base of the petiole with a long-conical
terminal cell. (Fertile segments with a sterile costu-
lar area absent or substantially so, or if present,
eng a sterile marginal area present that is as broad
costular one.)
1.
¢
PF:
h. Pinnae inequilateral; pinnules, on both sides of the
— at nearly ath a to the pinna-rachis (or
terminal
blackish, the ridges on the upper side of the rachis
[or in larger leaves of the pinna-rachis] lighter in
than the rachis proper; absent from the Lesser
Antilles). Eee .. 2. P. tartarea.
g. Small leaves rather fully bipinnate-pinnatif many
ultimate segments to Pana-
ma). regeeaesieiee
1 Geille of the apex Uf Tas hivices and the base of the
TAXONOMIC FERN NOTES. II. 59
petiole with only a short portion one cell wide below the
spherical terminal cell. i.
i. Rachis and the upper portion of the petiole (or all of it
except the base) straw colored to light brown, the
ridges on the upper side of the same color; spores
prominently ridged, the ridges with a defined
ietdes . P. chrysoconia.
i. Rachis and petiole yeddiah- brown to blackish; spores
heavily and closely ridged, the ridges with erose
margins that give a granular appearance to the spore.
(Fertile segments with a sterile costular area present,
a sterile marginal area absent, or if present, then only
half as broad as the costular one; as Rico and the
Lesser Antilles
d. Ulamate — usually mostly toothed ma deeply laciniate,
ew-veined (without a
midvein), ett to narrowly flabellate-cuneate. j.
j. Ultimate segments strongly laciniate into 1-veined lobes;
indument lacking on the lamina. (Lamina usually bearing
pinnae are more or less reduced. ............ 12. P. schizophylla.
a (Ultimate segments entire and several-veined to deeply cleft
into 2- several-veined lobes (these often strongly toothed) ;
ceraceous indument usually present on the under surface of
the lamina. 1.
1, Basal pinnae reduced; a pinnules usually strongly
benign Greater Anti
soars brown, vert but not Prag ose indu-
ulph
‘White OF PORE) .3, 5S Ae P. “Eggersii”.
1. Basal pinnae the largest or nearly so; apical pinnules
ona spreading; Africa, Madagascar Mascarenes.
n. Lamina lanceolate or deltoid-elongate with cune:
Parumeged lobes. BE b Bie lepinden.
. Lamina broadly — or pent: nts
rounded, not emarginate or cleft. ...... 14. P Humbertii.
NOTES ON THE SPECIES OF PITYROGRAMMA
In the following notes, the distribution of the American
species and varieties has been taken from the material in
the Harvard University Herbarium (a and GH) and in the
United States National Herbarium (US).
In some matters of taxonomy, or nomenclature, this study
is incomplete or inconclusive: a discussion of these matters
is included in the appropriate places. I have tried to include
60 ROLLA TRYON
all of the basionyms of species names that have been cer-
tainly or probably applied to native plants. Varietal names
are given only where they are pertinent to the nomencla-
ture. I have made no attempt to account for the names (in
various categories) that have been applied to material of
horticultural or presumed hybrid origin.‘°
1. Pityrogramma calomelanos (L.) Link, Handb.
Gewichse 3:20. 1833
A discussion of the difference between P. calomelanos and
P. tartarea is presented under the latter species.
la. P. calomelanos var. calomelanos
A tich I l L. Sp. Pl. 1072. 1753. LINN
sheet 1245.19, photo a, GH, is this species; maintained over
A. ebeneum L. by Lam. Encycl. 1:38. 1783.
Acrostichum ebeneum L. Sp. Pl. 1071. 1753. Lectotype:
Sloane, Hist. Jam. t. 53, f. 1. (Jenman, perhaps, made this
choice in Jour. Bot. Brit. For. 24:38. 1886. The Sloane fig-
ure should stand as the type of A. ebeneum because the Lin-
naean specimen, LINN sheet 1245.14, photo A, GH, is a small
plant of Pityrogramma tartarea).
Gymnogramma bidentata Presl, Rel. Haenk. 1:18, t. 2,
f. 3. 1825, ex char. et icon.
Gymnogramma. distans Link, Hort. Berol. 2:53. 1833. A
presumably authentic specimen “Hort. Berol. Sept. 1842”
det. Link is illustrated by Domin, 1929: t. 7. Link says ‘Hab.
in Brasilia?” and this may be of horticultural origin rather
than from a native source.
i as 7, Ls
y g va var. denudata Harr. Jour.
Linn. Soc. Bot. 16:37. 1877. Isotype: Pebas, Peru, Steere,
K!, photo GH.
Pityrogramma chamaesorbus Domin, 1928: 6. Type:
British Guiana, Schomburgk 390, K!, photo GH.
Pityrogramma imsularis Domin, 1928: 6. Syntypes:
Princes Island, Barter 1912; Fernando Po, Barter, Gustav
Mann. Tardieu-Blot, Flor. Madagasc. & Comores Fam. 5, 1:
117. 1958 ; and Alston, Ferns and Fern Allies W. Trop. Afr.
(FL. W. Trop. Afr., ed. 2, Suppl.), 38. 1959, both reduce this
species to P. calomelanos.
Pityrogramma distans (Link) Domin, 1929: 49.
"See Domin, 1955,
TAXONOMIC FERN NOTES. I. 61
Lamina ceraceous beneath, the wax white to pale (lemon)
yellow or pale roseate; or rarely the lamina glabrous.
Southern Florida; Greater and Lesser Antilles; Mexico
to Panama; French Guiana west to Colombia, Ecuador and
Galapagos Islands south to Bolivia and Argentina; Paraguay
and Brazil. Widely introduced in the Old World tropics.
lb. P. calomelanos var. aureoflava (Hook.) Weath. ex
Haney Man. Cult. Pi. 64. 1926.
var. aureoflava Hook. Gard.
Peers t. 50 text. 1862. Of the several specimens cited by
Hooker, I would take Seemann 948 as the one that may
best serve as a type but unfortunately I have not seen it.
Other specimens may (or certainly do) represent either
other species or else the pale yellow variant of P. calomelanos
which I refer to var. calomelanos.
Pityrogramma austroamericana Domin, 1928: 7. (Also
Kew Bull. 1929: 221.) Lectotype: Bolivia, Mandon 1549 bis,
K!, photo GH; isotype, GH! (Five other collections were cited
in a and one additional one in Sari
Pi icana (Do-
min) eee ne Midl. Nat. 12: "280. 1931.
Bailey did not properly transfer Hooker’s epithet to P.
calomelanos but C. A. Weatherby, then Editor of the Gray
Herbarium Index, accepted the name as cited above in the
Index and this may be taken as a correction of Bailey’s
publication.
Lamina ceraceous beneath, the wax bright yellow to
orange-yellow.
Costa Rica ; Venezuela, Colombia, Ecuador and Galapagos
Islands south to Bolivia and northwestern Argentina ; Brazil.
Introduced in the Hawaiian a aige Java and Tahiti. Re-
ported (as P. t icana) as duced in Africa by
Alston & Schelpe (Jour. So. Afr. Bot. 18:169. 1952).
le. Pityrogramma calomelanos var. ochracea (Presl) Tryon,
comb. noy. FIG. 1.
Acrostichum caudatum Cav. Deser. 242. 1802. Type:
“Panama”, Née, seen by C. Chr. at MA (Dansk Bot. Ark. 9°:
10. 1937, where it is figured, in part, t. 2, f. 4-5.). I believe
that this name belongs here (much less likely it is P. tartarea
var. Jamesonii), and that it was obtained in Andean South
America rather than in Panama.
62 ROLLA TRYON
Gymnogramma ochracea Presl, Rel. Haenk. 1:17. 1825.
Type: Peru, Haenke, PR (Herb. no. 24360), photo GH, US;
probable isotypes, K, fragment Ny!, B, fragment, NY!
Gy J di ides Karst. ex Mett. Ann. Sci. Nat.
V, 2:212. 1864. Syntypes: Colombia, Bogota, Karsten, B!,
photo GH, fragment NyY!; Colombia, Caqueza, 1700 m.,
Triana 606, duplicate: BM, photo and fragments us! *
Gymnogramma Hookeri J. Sm. ex Hook. & Bak. Syn. Fil.
381. 1868. Syntypes: Colombia, Purdie, fragment NY!;
Colombia, Schlim 609; Bolivia, Mandon 1547.
Gymnogramma Ballivianii Rosenst. Fedde Repert. 6:314.
1909. Type: Bolivia, Buchtien 1038, S-PA! fragment ex
Rosenst. Us! (Steinbach 8507, Buchtien 35 and other speci-
mens commonly so named are the same).
Ceropteris adiantoides var. peruviana Hieron. Hedwigia
48:221. 1909, ex char. & loc. Type: Peru, Stibel 1096, B.
Not P. calomelanos var. peruviana (Desv.) Farw. (sub P.
tartarea).
Ceropteris tartarea var. ochracea (Presl) Bonap. Notes
Pterid. 2:150. 1915.
Pityrogramma. ochracea (Presl) Domin, 1928: 8.
Pityrogramma adiantoides (Mett.) Domin, 1928: 8.
Pityrogramma Ballivianii (Rosenst.) Domin, 1928: 10.
Pityrogramma perelegans Domin, 1928: 8. Type: Tara-
poto, Peru, Spruce in 1855-56, K!, photo GH.
Pityrogramma caudata (Cay.) C. Chr. Dansk Bot. Ark.
9°:10. 1937.
Fic. 1. Pityrogramma calomelanos var. ochracea: la, pinna, X 1/2; Ib, pinna, X 1;
le, vaseular bundles in petiole, enlarged. Fic. 2. P. ferruginea: 2a, pinnae, X 1/2; 2b,
vascular bundles in petiole, enlarged; 2c, S i
bundles in petiole, enlarged; both from Peru, Tryon & Tryon 5 3 6.
peer Oe Noe KG “wematies, Gk keene cP bah
: Colombia, 3945.
TAXONOMIC FERN NOTES. II.
64 ROLLA TRYON
Presl’s name has been often misapplied to forms of this
species, or of P. tartarea, with yellow wax.
Lamina pubescent beneath, not ceraceous.
The spores of some specimens of var. ochracea are of the
typical Pityrogramma type with well defined dark ridges.
Other specimens have spores that are more spherical than
the usual type, although somewhat flattened in three planes
on the commissural face, and the tan exospore is smooth
or only slightly I ft It-is ptional to find such
different spores in the same species (or variety) but I have
been unable to find any character that correlates with the
spore type and there is every indication that the unusual
spores are mature.
Some specimens of var. ochracea with the “normal” spore type are
the following (all from Peru): HUANUCO: Macbride 4994 (us), Allard
22022 (GH). AYACUCHO: Killip & Smith 22667 (us). cuzco: Vargas
4023 (us), Scolnik 881 (US), Biies 1715 (US). Some specimens with
the smoothish spores are: PERU: (all from cUzco), West 8013 (UC),
Cook & Gilbert 1715 (us), Mexia 8089 (Gu, US), Herrera 1285 (US),
Tryon & Tryon 5366 (GH). BOLIVIA: R. S. Williams 1373 (cH), Krukoff
10094 (F, GH).
Honduras; Venezuela, Colombia, Ecuador, Peru and
livia.
2. Pityrogramma tartarea (Cay.) Maxon, Contrib.
U. S. Nat. Herb. 17:173. 1913
More difficulty exists in the separation of this species from
the previous one, P. calomelanos, than would be expected
from an examination of the common forms of each. Some
of this difficulty, it is true, is due to specimens that are single
leaves, but the characters of each do vary toward the other.
species are common, widely distributed, and grow in
a variety of habitats and the extent to which their variation
TAXONOMIC FERN NOTES. II. 65
rachis; pinnules obtusely lobed . pinnatifid; basal inferior
beatieren at about the same angle to the pinna-rachis on all
pinnae (or in large ae ee fees pada tertiary segments so
in relation to the p' P. tartarea.
This species has a less extensive range than P. calomelanos
and both the variety with yellow wax and the one with the
lamina pubescent are also more restricted than their counter-
parts in P. calomelanos.
2a. P. tartarea var. tartarea
Acrostichum tartareum Cav. Descr. 242. 1802. Type: near
Guamantanga, Peru, Née, seen by C. Chr. at MA (Dansk Bot.
Ark. 9*:10. 1937).
Hemionitis dealbata Willd. Sp. Pl. 5:131. 1810, nom.
— illegit. seapeeen tC tartareum Cav.).
Ges. Naturf. Freunde
Becks Mag. 5:329. 1811. Type: eon Jos. de Jussieu (Herb.
Jussieu no. 1009) P!, photo GH.
Gymnogramma dealbata Link, Hort. Berol. 2:52. 1833,
(epithet from Hemionitis dealbata Willd.), nom. superfi.,
illegit. (—Acrostichum tartareum Cav.), not Presl, 1825
(sub P. dealbata).
Ceropteris Stuebelii Hieron. Hedwigia 48 :223, t. 10, f. 9.
1909. Type: Colombia, Stiibel 12, B!, fragment US!; ex B, GH!
Pityrogramma peruviana (Desy.) Maxon, Contrib. U. S.
Nat. Herb. 17:173. 1913.
Pityrogramma Stuebelii (Hieron.) Domin, 1928: 7.
Pityrogramma tripinnata Domin, 1928: 7. Type: Mexico,
Parry & Palmer 1004 in 1878, K!, photo GH; isotype, Us!
Pityrogramma calomelanos var. peruviana (Desv.) Farw.
Am. Midl. Nat. 12:280. 1931.
Lamina ceraceous beneath, the wax white to pale (crearn)
yellow, rarely very sparse and almost colorless; rarely partly
ceraceous-pubescent.
Greater Antilles; Mexico to Panama; Venezuela, Colom-
bia, Ecuador and Galapagos Islands south to Bolivia; Brazil
(Rio Grande do Sul, Parana and vicinity of Itatiaya).
2b. Pityrogramma tartarea var. aurata (Moore) Tryon,
nogramma tartarea var. aurata Moore, Gard. Chron.
1870: 493. Type: Cultivated by Veitch from a collection by
Pearce in Peru. (The specimen of Pearce 218 at K!, photo
66 ROLLA TRYON
GH, Domin, 1929: t. 1, may be the type as Domin says; it
does clearly represent Moore’s name).
Pityrogramma praestantissima Domin, 1928: 6. Type:
Colombia, Kalbreyer 1909, K!, photo GH.
Pityrogramma Presliana Domin, 1928: 6. Type: Peru,
Matthews 1823, K!, photo GH (although Domin also cited
Pearce 218, he annotated that collection, K, as P. Presliana
var. aurata (Moore) Domin and later stated it to be the
type of that variety).
Lamina ceraceous beneath, the wax bright yellow. Colom-
bia, Ecuador and Galapagos Islands to Argentina.
2c. Pityrogramma tartarea var. Jamesonii (Baker) Tryon,
comb. nov.
Gymnogramma Jamesonii Baker, Syn. Fil. 516. 1874.
Type: Ecuador, “Pinchinchan” [Pichinchan] Andes, Jame-
son, K!, photo GH.
Gymnogramma xerophila Baker, Jour. Bot. Brit. For. 19:
206. 1881. Type: Colombia, Kalbreyer 1563, K!, photo GH.
Pityrogramma Jamesonii (Baker) Domin, 1928: 8.
Pityrogramma xerophila (Baker) Domin, 1928: 8.
Lamina pubescent beneath, not ceraceous. Colombia and
Ecuador.
3. Pity: h
(Desv.) Maxon ex
Domin, 1928: 10
Acrostichum chrysoconium Desv. Mém. Soc. Linn. Paris
6:212. 1827. Type: Peru, evidently Dombey, P, photo GH;
isotype, B!, photo GH, fragment vs!
G g i is Kl. Linnaea 20:413. 1847.
Type: British Guiana, Schomburgk 1154, B!, photo GH, frag-
ment NY!; isotype, K, fragment Ny!
Gymnogramma Ornithopteris Kl. Linnaea 20:413. 1847.
Type: Venezuela, Moritz 288, B!, photo GH, fragment Us!;
isotype, K!, photo GH.
mma flexilis Kl. Linnaea 20:414. 1847. Type:
Venezuela, Mérida, Moritz 287, B: isotype, K, fragment NY!
Ceropteris obtusa Fée, Mém. Fam. Foug. 8:80. 1857. Type:
Colombia, Schlim (873; isotype, P!, photo GH.
Pityrogramma Ornithopteris (K1.) Maxon ex Knuth,
Fedde Rep. Beih. 43:95. 1926.
—
™Domin, 1929: t. 1.
TAXONOMIC FERN NOTES. II. 67
Pityrogramma flexilis (K1.) Domin, 1928: 5.
Pityrogramma obtusa (Fée) Domin, 1928: 10.
Pityrogramma subn‘valis Domin, 1928: 5. (Also Kew Bull.
1929: 220). Type: Colombia, Santa Marta, Purdie in 1844,
K!, photo GH.
Jamaica (Blue Mt. Peak) ; Costa Rica to Panama; British
Guiana west to Colombia and south to Bolivia; principally
or wholly at the higher elevations.
4. Pityro~ramma dealkata (Presl) Tryon, comb. nov.
Gymnogramma dealbata Presl, Rel. Haenk. 1:18, t. 3,
f. 1. 1825. Lectotype: Panama, Haenke, PR (sheet 24358a),
photo GH (PR sheet 24358b, photo GH, is probably P. ealome-
lanos) .
Ceropteris Schaffneri Fée, Mém. Fam. Foug. 8:80. 1857,
ex char. & loc. Syntypes: Mexico, Schaffner 165a, 165b.
Pityrogramma Schaffneri (Fée) Weath. Contrib. Gray
Herb. 114:25. 1936.
The photograph of Presl’s type is a very close match for
some specimens of this species, for example, Costa Rica,
Seamman 7055 (GH) and Panama, Allen 147 (GH). This
species was long referred to as Gymnogramma peruviana
Desv. (or Pityrogramma or Ceropteris peruviana) but in
1936 C. A. Weatherby pointed out that Desvaux’s type was
actually a specimen of P. tartarea.
Mexico to Panama.
5. Pityrogramma chrysophylla (Sw.) Link, Handb.
Gewiichse, 3:19. 1833
Acrostichum chrysophyllum Sw. Schrad. Jour. Bot. 1800°:
14. 1801. The specimen at S-PA (Herb. Swartz!), is this
species.
Acrostichum aculeatum Desv. Ges. Naturf. Freunde Berl.
Mag. 5:310. 1811. See Weath. Contrib. Gray Herb. 114:15.
1936. Not L. Syst. Nat. ed. 10, 2:1320. 1759.
Ceropteris intermedia Fée, Mém. Fam. Foug. 11 (Hist.
Foug. Antill.) :30. 1866. Type: Guadeloupe, L’Herminier in
1861, P, photo GH.
k i Carruthers in Seemann, FI.
Vit. 370. 1873, echnrer tog from Brack. U. S. Expl. Exped.
16:24 (sub G. tartarea), 1854. Type: near Pago-Pago,
Island of Tutuila, Samoa, Brackenridge, Us!
68 ROLLA TRYON
Pityrogramma Brackenridgei (Carr.) Maxon, Carnegie
Instit. Wash. Publ. 341:124. 1924.
Pityrogramma caribaea Domin, 1928: 6. Syntypes: Dom-
inica, Imray 71, K!, photo GH; Dominica, Bradford 395, K!,
photo GH.
The white and yellow color forms, although especially
striking in this species, do not merit recognition.
The material I have seen from Samoa, which has con-
sistently been recognized as an endemic (C. Chr. Pterid.
Samoa, Bull. Bishop Mus. 177:57-58. 1943), is surely this
species. Porto Rico (rare) ; and the Lesser Antilles; intro-
duced in Samoa.
6. Pityr £,
i (Kze.) Maxon, Contrib.
U.S. Nat. Herb. 17:173. 1913. Fic. 2
Gymnogramma ferruginea Kze. Linnaea 9:34. 1834. Iso-
type: Quebrada de Chinchao, Peru, Jul. 1829, Poeppig, Diar.
1160, P!, photo GH.
Gymnogramma Bommeri Christ, Bull. Bot. Soc. Belg.
35:237. 1896. Syntypes: Costa Rica, “Biolley in Pittier
1066”; Costa Rica, Pittier 1169. I have not seen either of
these collections but am accepting Wercklé, Costa Rica, det.
Christ, Us! as an authentic representative of the name.
The material in Central America is more variable than
that in Peru. The pinnae are often petiolulate there, while
in Peru they are uniformly subsessile. Also the lamina may
be long-triangular or deltoid in Central America while in
Peru it is narrowly elliptic.
Guatemala to Panama; Peru.
. 7. Pityrogramma trifoliata (L.) Tryon, comb.
nov. FIG. 3.
L. Sp. Pl. 1070. 1753. LINN
sheet 1245.9, photo a, and Sloane, Hist, Jam. t. 45, f. 2, are
is species.
A tich FE Se ae
both th
Trismeria argentea Fée, Mém. Fam. Foug. 5(Gen. Fil) :
165, t. 14A, f. 1. 1852. Syntypes: Caracas, Moritz; Minas
Geraes, Pohl.
Trismeria aurea Fée, Mém. Fam. Foug. 5(Gen. Fil) :165.
1852, nom. superfi. illegit. (—Acrostichum trifoliatum L.).
Trismeria microphylla Fée, Mém. Fam. Foug. 5(Gen.
Fil.) :165. 1852, ex char. “Habitat in Peruvia”.
TAXONOMIC FERN NOTES. II. 69
Trismeria trifoliata (L.) Diels, Nat. Pflanz. 1!:265. 1899.
Southern Florida; Greater Antilles ; Mexico to Costa Rica
(notably absent from Panama) ; Venezuela, Colombia south
to Bolivia, Argentina and Chile (Arica) ; Paraguay, Uru-
guay and Brazil.
I am recognizing four putative hybrids of Pityrogramma
trifoliata, one of them with P. ferruginea and the other three
with the varieties of P. calomelanos. All of these have the
long, narrow lamina of P. trifoliata, the strongly ascending-
arching lower pinna-stalks and (at least in large leaves) the
same type of vascular bundles. There is a strongly developed
tendency in them toward long and entire segments. Char-
acters of the pinnae and the vascular bundles of two of the
putative hybrids, and their parents, are illustrated in Figs.
1-5. While these are of interest, I do not believe that a
binomial is justified for any of them. The hybrids are all
distinguished from P. trifoliata by their pinnae with numer-
ous segments. In P. trifoliata the pinnae are entire to 3-, 5-
or very rarely 7-foliolate.
The material has not been adequate to establish the degree
of fertility of the hybrids. An examination of the fertile
leaves available suggests that they may produce no viable
spores or relatively few of them. The collections Tryon &
Tryon 5440 and 5449 were both single plants.
7a. P. calomelanos var. calomelanos trifoliata
Gymnogramma longipes var. concolor Baker, Jour. Bot.
Brit. For. 16:301. 1878. Type: Paraguay, Balansa 334, K,
is probably this hybrid.
Ceropteris longipes var. argentea Rosenst. ex Hassler,
Trab. Instit. Bot. Farm. Buenos Aires 45:51. 1928. Type:
Paraguay, Hassler 3176, is probably this hybrid.
Petiole and rachis glabrous or ceraceous, segments white
ceraceous, or with nearly colorless wax, benea
ARGENTINA: Prov. Tucuman, July, 1912, Castillon (Inst. Lillo 41858)
(GH, in part). PARAGUAY: April 25, 1946, Rojas 13343 (@H), Decem-
ber, 1915, Rojas 1117 (GH); Pedersen 3158 (US). BRAZIL: Minas
Geraes, 1948, iors 1161 (Us), a 2220 (Us)
7b. P. eal var. Af trifoliata
Gymnogramma longipes Baker, Jour. Bot. Brit. For. 16:
301. 1878. Type: Paraguay, Balansa 333, K!, photo GH.
Trismeria longipes (Baker) Diels, Nat. ‘Piianz. a
1899.
70 ROLLA TRYON
Trismeria trifoliata var. subbipinnata Hieron. Hedwigia
48:223. 1909. Type: Ecuador, Stiibel 733, B, is probably
this hybrid.
Petiole and rachis glabrous or ceraceous, segments with
yellow wax beneath.
This is perhaps the same as 7a, the yellow wax being
derived from a yellow form of P. trifoliata rather than from
P. calomelanos var. aureoflava.
ADOR? ARGENTINA: Tucuman, Venturi 9646 (US). PARAGUAY:
December, 1902, Fiebrig 610 (GH).
7c. P. calomelanos var. ochracea X trifoliata FIG. 4
Gymnogramma Herzogii Rosenst. Med. Rijks. Herb. 19:
21. 1913. Type: Bolivia, Herzog 2000; isotype, US!
Petiole and rachis thinly pubescent to glabrous, segments
pubescent beneath.
PERU: Dept. Junin, Tryon & Tryon 5440 (GH); Dept. Cuzco, Biies
1936 (US). BOLIVIA: Herzog 2000 (us).
7d. P. ferruginea X trifoliata Fic. 5
Petiole and rachis closely and densely lanate, although
deciduously so. The only collection has the pinnules mark-
edly irregular in their length.
PERU: 10 km. sw. of San Ramén, Dept. Junin, Oct. 28, 1956, Tryon
& Tryon 5449 (GH).
8. Pityrogramma Lehmannii (Hieron.) Tryon,
comb. nov. FIG. 6
G J Leh it Hieron. Engl. Bot. Jahrb. 34:
474. 1904. Type: near Aquaclara, Popayan, Colombia, 1200-
1400 m., Lehmann 8944, B!, photo GH; isotypes, GH! US!
The unusual ch ters of this distinctive species have
been mentioned in the key and in the discussion of the genus.
It is known only from the original collection.
Colombia.
8.
Pityrogramma triangularis (Kaulf.) Maxon,
Contrib. U. S. Nat. Herb. 17:173. 1913
The four varieties of this species were treated by Weath-
erby* and in spite of additional knowledge his taxa and
their status are certainly still justified. The genetical and
cytological study of Alt and Grant", for example, is an im-
22: 113-120. 1920.
. Grant,
*Rhodora
“alt, K. S. and V. Cytotaxonomic observations on the goliback fern, Brittonia
12: 153-170. 1960.
TAXONOMIC FERN NOTES. II. 71
portant contribution to our understanding of this species
but it does not furnish a basis for different taxonomic
conclusions.
9a. P. triangularis var. triangularis
Gymnogramma triangularis Kaulf. Enum. Fil. 73. 1824.
Type: San Francisco Bay, California, Chamisso, probably
LE; isotypes, P, (Chamisso 1875) photo GH, B (fide Alt &
Grant).
Baja California to sw. British Columbia, s. Nevada and
sw. :
9b. P. triangularis var. viscosa D. C. Eaton, Ferns N. Am.
2:16, t. 48, f. 5. 1879. Type: usually considered to be: San
Diego, California, “Gymnogramme viscosa”, Nuttall, PH;
but other material cited, or especially that illustrated, by
Eaton might be a better lectotype.
Pityrogramma viscosa (D. C. Eaton) Maxon, Contrib.
U.S. Nat. Herb. 17:173. 1913.
Maritime: Baja California and s. California.
9c. P. triangularis var. pallida Weath. Rhodora 22:119.
1920. Type: California, Heller 8141, GH!
Pityrogramma pallida (Weath.) Alt & Grant, Brittonia
12:168. 1960.
Central California.
. P. triangularis var. Maxonii Weath. Rhodora 22:119.
1920. Type: Arizona, Blumer 3271, US; isotype, GH!
Baja California, s. California e. to Arizona.
10. Pityrogramma sulphurea (Sw.) Maxon, Contrib.
U.S. Nat. Herb. 17:173. 1913
Acrostichum sulphureum Sw. Prod. Veg. Ind. Occ. 129.
1788. Type: Jamaica, Swartz, s-PA (Herb. Swartz) !, frag-
ment us! (the Plumier, Fil. t. 44 cited by Swartz is P. chry-
sophylla) . .
Cuba, Jamaica, Hispaniola, Porto Rico.
11. Pityrogramma “Eggersii”
The commonly accepted name, Pityrogramma Eggersii
(Christ) Maxon" can not be used for this species because
the type specimen (Cuba, Eggers, comm. C. Rensch sub no.
4882a, Herb. Christ-P!) of Gymnogramma Eggersii Christ**
“Contrib. U.S. Nat. Herb. 24: 62. 1922.
® Bull. Soc. Bot. Belg. 337: 92. 1894.
72 ROLLA TRYON
is a species of Asplenium and has been so annotated by C.
V. Morton. Although this species does not have a name, I
am deferring to C. V. Morton’s forthcoming work on the
ferns of Cuba, where it will be described and named.
Cuba and Hispaniola.
Representative specimens: cuBA: Howard 5017 (GH, NY, US); Jack
7124, (GH, NY, US). HISPANIOLA: Ekman (Haiti) 4824 (Ny, US).
12. Pityrogramma schizophylla nape Sapa
Contrib. ie S. Nat. Herb. 24:61. 1
G izophylla Jenm. Jour. = Brit. For.
15:266. 1877. Type: Jamaica, Jenman, “No. 16, Hb. Kew,
1876”, K!, photo GH; isotype, Jamaica, Jenman, NY!
Jamaica and Hispaniola.
13. Pityrogramma argentea (Willd.) Domin, 1928: 6
13a. P. argentea var. argentea
Hemionitis argentea Willd. Sp. Pl. 5:132. 1810.
Tropical and south Africa; Madagascar and Mascarene
slands.
13b. P. argentea var. aurea (Willd.) Domin, 1929: 20.
Hemionitis aurea Willd. Sp. Pl. 5:131. 1810.
Gymnogramma argentea var. aurea (Willa.) Mett. ex
Kuhn, Fil. Afr. 59. 1868.
Pityrogramma aurea (Willd.) C. Chr. Cat. Pl. Madagas.
Pterid. 46. 1932.
d. and M: Islands.
I have made no study of the African species and in this
one and the next I have followed the treatment of Tardieu-
Blot**. Christensen", however, treats var. aurea as a species.
14. Pityrogramma Humbertii C. Chr. Dansk Bot.
Ark. 7:111, t. 43. 1932.
Madagascar (I do not know var. elongata C. Chr., op. cit.
112, from the Belgian Congo).
DUBIOUS TAXA AND NAMES.
The four following names may represent valid taxa but the a
— seen has not been adequate to form a basis for definite co
usions:
Pityre chrysophylla var. euchrysa Ekman ex C, Chr. Kungl.
- Madagas. & & Comores, Fam. 5, 1: 118-120. 1958.
"oe. eit. and Dansk Bot. Ark. 7: 112, t. 48, £. 5. 1932.
TAXONOMIC FERN NOTES. II. 73
Svensk. Vet. Akad. Handl. a 57, t. 14, f. 1-3. 1936. Type: Hispaniola,
Ekman 13831, 8; isotype, U:
This may be a variety = ss chrysophylla, as Christensen proposed,
or savage a variety of P. chrysoconia; or it may represent an endemic
species.
Pityrogramma Pearcei (Moore) Domin, 1928: 9.
Gymnogramma Pearcei Moore, Gard. Chron. 1864: 340. Type:
Cultivated by Veitch from a collection by Pearce in “Chile”. Two speci-
mens at K!, photos GH, may be considered as authentic = beg not find
a proper holotype) : “Pinna of Gymnogramma Peareei, n. sp......-
and Pearce 274, ex Veitch 7/84. The original collection was meine
ly mot made in Chile. Illustration: Schneider, Choice Ferns 2: f. 68.
Baker, Gard. Chron. 1872: 1587. Type:
“Andine side of South America, Hort. Kew ex Veitch”; this is prob-
ably the specimen at K!, photo GH, annotated by Baker as “G. decom-
posita” and with the original label “Gymnogramma Pearcei, H. Kew
10/71, from Veitch”. TU Ie Garden 6: f. on 495.
fi Rosenst. Mém. Soc. Neuchat. Sci. Nat.
5: 45, 54, t. 6. f. 10. 1912. Type: Colombia, E. Mayor 144; isotype, US!,
P! photo GH.
Pityrogramma decomposita (Baker) Domin, 1928:
Pityrogramma fumarioides (Rosenst.) Maxon, Ee aren Soe. Wash.
51: 38. 1938.
The three basionyms listed above all represent plants with highly
dissected leaves. I strongly suspect that this material represents a
single species but it is not sufficient to enable me to present the
synonomy with assurance nor to place the species among the others of
Pityrogramma pulchella (Moore) Domin, 1928: 9.
Gymnogramma pulchella Moore, Gard. Chron. 1856: 597, figure.
Type: “Hort.”, presumably ex Lag ssioaceay- ex Venezuela. I did not
find a proper holotype at Kew; K!, in Moore’s herbarium,
photo GH, “G. pulehella, Hort”. Hort. Vaich, 1860”, is probably
another species.
I have associated the following material with this name on the basis
of the small figure published by Moore. VENEZUELA: (probably Rancho
Grande). Tschudi 176 (VEN); Colonia Tovar, Fendler 357 (YU, the
specimen at kK is figured in Domin, 1929: t. 6); ne. of Bergatin, Est.
Anzoategui, Steyermark 61327 (F, VEN
nel
with the equally uncertain P. Pearcei diesel
Pityrogramma triangulata (Jenm.) Maxon, Contaib. U.S. Nat. Herb.
17: 173. 1913.
Gymnogramma triangulata Jenm., te Bot. Dept. Jamaica n.s. 4:
74 ROLLA TRYON
the margin. It should be mentioned, however, that the character of
the veins and margin in P. argentea is not unique. It is sometimes
+smilar in P. “Eggersii” and its development in P. triangulata may not
show a relation to the African species.
The following names are probably synonyms of some of the American
species but I have not been able to place them with certainty:
Ceropteris monosticha Fée, Mém. Fam. Foug. 7: 44, t. 22, f. 2. 1857.
Type: Mexico, Schaffner 155. Perhaps = P. tartarea var. tartarea.
Ceropteris plicata Fée, Mém. Fam. Foug. 8: 80. 1857. Type: Mexico,
Schaffner 164. Z
Ceropteris serrata Fée, Mém. Fam. Foug. 8: 81. 1857. Type: Mexico,
Schaffner 162. Perhaps = P. calomel var. calomel:
‘ityrogramma subfleruosa Domin, 1928: 7. Type: Montserrat,
Holme in 1879, K!, photo GH. Probably = P. chrysophylla.
Il. THE GENUS ANOGRAMMA
The genus Anogramma includes some five or six species
which are clearly related to Pityrogramma, especially to P.
sulphurea and others of spp. 10-13 (see above). The archi-
tecture of the lamina is similar to some of these, the axes
are grooved in the same manner and the sorus is the same.
Rathe.
soil and the sporophyte evidently lives for only one growing
310. 1904.
TAXONOMIC FERN NOTES. I. 75
season. In the smallest species (A. Lorentzii) the sporophyte
may consist of only a few small fertile leaves (to 1 cm. long),
a minute stem and a few roots. A gametophyte is often a
attached to herbarium i of the sp hyt
prothallus is reported to be perennating, eaeeryinige the fod
season and producing another sporophyte from specially
developed lobes the next growing season’’. This behavior of
the prothallus, however, can not well be used as a generic
character because it has been reported in only two of the
species (A. leptophylla and A. chaerophylla) and because
even in these it is not known to be of regular occurrence in
nature. In other species the gametophyte may not behave
in this way and its frequent occurrence on a sporophyte may
be due to the fact that the growth of the sporophyte is so
rapid that even when it is mature the gametophyte is still
present.
This reduced and specialized luti line can be rec-
ognized as a distinct genus, or as a group within Pityrogram-
ma as Domin® did. Were there truly intermediate species,
I would follow Domin’s classification. However, there are
none and the several species can be separated, among other
less constant characters, on the basis of the rhizome (and
usually the lamina) indument and the annual habit of the
sporophyte.
Rhizome scales atropurpureous to light brown, rigid and sclerotic to
firm and subsclerotic; sporophyte living for a few to many years,
the stem correspondingly well developed; lamina glandular,
strongly pubescent or most often ceraceous — glabrous or
slightly pubescent).
Rhizome scales thin, whitish, sometimes mixed with jaiatiag. trichomes
or only such trichomes present; sporophyte living only one growing
season, the stem correspondingly poorly developed; lamina glab-
rous or rarely sparsely pubescent.
ANOGRAMMA Link, Fil. Sp. Cult. 137. 1841. Type: A. leptophylla
(L.) Link -_ leptophyllum )
The usually accepted speci
1. A. chaerophylla (ean Link (the largest species). ‘Tropical
America.
2. A. guatemalensis (Domin) C. Chr. may be a variant of A. leptophyl-
la. Mexico to Costa Rica.
3. A. leptophylla (L.) Link. Tropical America, Africa and adjacent
islands, southern Europe, and eastward to New Zealand.
See, for example, Goebel, Bot. Zeit. 35: 671-711. 1877; Organographie, ed. 3, 1101.
1102. 1930. Bower, The Ferns 1: 276-278. 1923.
*Domin, 1928: 3, 9-10.
76 ROLLA TRYON
4, A. ascensionis (Hook.) Diels. Ascension Island.
5. A. Osteniana Dutra. Uruguay and Brazil (Rio Grande do Sul).
6. A. Lorentzii (Hieron.) Diels (the smallest species). Argentina,
Uruguay, Brazil. Gymnogramma Regnelliana Lindm. 1903 (not
Christ, 1900) may be a species distinct from A. Lorentzii rather than
a variety of it (Gymnogramma Lorentzii var. megaspora Lindm.
Hedwigia 43: 309. 1904).
Other species that have been placed in Anogramma are
referable to other genera (see C. Chr. Ind. Fil. and Suppls.).
One, however, that has been maintained in Anog by
Christensen (and others) clearly does not belong there.
is is A. microphylla which Domin™ treated in a separate
section, Pityrogramma section Monosorus. He was certainly
correct in removing it from his section Anogramma, but it
is probably not closely related either to that genus or to
Pityrogramma.
Anogramma microphylla (Hook.) Diels (Gymnogramma
microphylla Hook.), of the eastern Himalayas, has a creep-
ing rhizome with blackish, thickened trichomes, one U-
shaped vascular bundle in the petiole, unusual somewhat
elliptical segments, and tetrahedral-globose spores that are
pale yellow and have a perispore with a few prominent
wings and tubercules between them. The leaf, segments
and sorus are well figured in the original publication.
This species may represent a monotypic genus; or perhaps
it is a diminutive Monachosorum; or it may possibly be con-
generic with or related to Cerosora.— GRAY HERBARIUM,
HARVARD UNIVERSITY.
orn arene
*Domin, 1928: 10.
*Hook, Ie. Pl. t. 916. 1854. (Cent. Ferns, t. 16, 1854).
CONTRIBUTIONS FROM THE GRAY HERBARIUM
OF HARVARD UNIVERSITY
Edited by
Reed C, Rollins and Robert C. Foster
NO. CXC
PATTERNED GROUND AND VEGETATION
ON SOUTHERN BYLOT ISLAND,
NORTHWEST TERRITORIES, CANADA
By
Wi1aM H. Drury, Jz.
Published by
THE GRAY HERBARIUM OF HARVARD UNIVERSITY
AMBRIDGE, MASS., U.S. A.
1962
CONTRIBUTIONS FROM THE GRAY HERBARIUM
OF HARVARD UNIVERSITY
Edited by
Reed C. Rollins and Robert C. Foster
NO. CXC
PATTERNED GROUND AND VEGETATION
ON SOUTHERN BYLOT ISLAND,
NORTHWEST TERRITORIES, CANADA
By
Wriutam H. Drury, JR.
Published by
THE GRAY HERBARIUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASS., U.S. A.
Issued July 25, 1962
exe
aes
es
of
e
CONTENTS
INTRODUCTION
Pp. 4
Climatic
Statement BE the Problems ..
PART ONE — The Bylot Island Expedition ccc
The Study Area
PART TWO — Geological Processes. -srvsacsssssssmsneenenenee
I. MECHANICAL WEATHER
Frost-riven Stones .........
Wind-sculptured Stones -crrscccenmescmeneenmneenmennt
Avalanche Gullies in the Sedimentary
Outerops
fel a3
Block Streams
II. SOLIFLUCTION
Frost-heaving
Solifltiction FOr .ecesseeeeeeeeneemeennenseneeetnmnaenetneie
Ma area neeetch NC aiepions sectarian
1S)
aan Surface rae Uplifted Peneplains
in Temperate Regions, --.nsm-nne
Ill. STRUKTURBODEN — THE DEVELOPMENT OF
PATTERN GROUND. ~eccssvssseeseeeemeneenenemeneenenenernectnet
A. Non-sorted stripes — vegetation or
SOU SEYUPCS -eeeveevnereerrrenreneenerne
B. Non-sorted circles — mud-bol frost-
boils, or f1
TOSt-SCATS
Mud-boils and non-sorted poles:
& Contraction eracks on the ue
D. Hummock polygons enone
Origin of hummocks and hummock
OLY LOTS neem
an
on
E. Micropolygonal patterns —
F. Five- and six-sided polygons -ccccsneneom
IV. ORGIN AND MODIFICATION OF MAJOR FOUR-
SIDED PATTERNS OF FROST-CRACKS ccsescsctsnneoen
A. Geographic and climatic distribution
B. Origin of major frost-Cracks ecco
1. What force is strong enough? ...
(a) Contraction due to cold ...
(b) Deformaton of frozen
NG sta
2. Are the cracks forming now?
Cc. Frost-cracks in other dead pn her eueines
D. iewent
barrens
Effect of Vegetation oo ncicccccnun
Effect of Soil Character aCe me rem
vegetated PEROS a ees
1
2.
3.
4.
5.
PART THREE — Vegetation
The effects of vegetation on soil
POUM aN ois 5 Oa
Effect of the presence of organic
material on frost activity ........
Occurrence of isolated contrac-
tion cracks in organic soils ......
How do these two frost-crack
types combine to form peat
1) Gi olen EMU Oe cae
Depths to frost across the peat
ridges
How does one peat ridge develop
from ridges on two sides of a
Eros crack Fo
agoccrage regularly spaced
ponds
ice Bh ees. not associated with
66
GSB
B. Regional and microtopographic differences
68
C. Primary factors controlling vegetation ....... 69
1. Exposure and shelter --e-nnnnnee 70
2. Moisture ilabilit; 71
3B. Sequence of SMOW-Melt ncevccnnnenennnnee TL
4. Effects of the full rays of the sun... 72
D. Tirme Of flO WHI onceceeecseneersneeovreeereeerneeraneornnneonnnrnnce 72
E. Variation in the richness Of Site mm 12
F. Vegetation types and habitat selection by
birds 73
G. Indicators, dominance, succession and frost
action 75
VEGETATION 77
Altitudinal Limit of Vegetation on Mt. Thule... 82
PART FOUR — Plant List 82
Non-vascular Plants 82
Vascular Plants 83
SUMMARY 100
Frost Features 100
Vegetation 101
LITERATURE CITED 102
Map 1. BYLOT ISLAND AND ECLIPSE SOUND ....:+-esss-sssssssrersesseeerenees Cea |
11
69
PRS LU 3
RILL 10
Figure 3. SUBSURFACE WATER-FLOW INTO MUD-BOIL EXCAVATION -. sae
Figure 4, MUD-BOIL HUMMOCKS 22
Figure 5. ARCTIC WILLOW AND MUD-BOIL WITH CRACKS . 24
Figure 6. CONTRACTION CRACKS ON POND SHORE ........ 24
Figure 7. HUMMOCK POLYGONS ON SOUTH SLOPE .......... . 28
Figure 8. mompoioansn® PENTAGONS UNDER LATE SNOWBANK ...... 28
Figure 9. NTO CRACKS, PEAT RIDGE,
AND jee 30
Figure 10. INTO AND
CRACKS 33
Figure 11. CONTRACTION CRACK IN NON-SORTED STRIPE -------e---r--- 40
CONTRACTION CRACK AND PEAT RIDGE ON 16-FOOT BEACH
RECTANGULAR DEFORMATION CRACKS WITHOUT MOUNDS
ON SEA BEACH
RECTANGULAR DEFORMATION CRACKS IN SEA ICE
CRACK
IN SEA BEACH FROM
THE 47
DEFORMATION CRACKS ON PLATEAU TOP FROM THE AIR.... 47
DEFORMATION CRACKS AND PONDS ..... eee 51
FROST 51
ipa INTO DEFORMATION CRACKS AND SOLIFLUC-
LOBE
Heaesensaay CRACKS WITH MOUNDS
CRACK:
ECTING PEAT RIDGES ....
‘TERS!
PEAT RIDGE DIKES
WITH PEAT RIDGE
SHOWING
TION CRACKS
IN DEFORMA-
‘
PATTERNED GROUND AND VEGETATION —
“ON SOUTHERRN BYLOT ISLAND
sate ye Tee Ms
PATTERNED GROUND AND VEGETATION
ON SOUTHERN BYLOT ISLAND,
NORTHWEST TERRITORIES, CANADA
By WILLIAM H. Drury, JR.
INTRODUCTION
The study of frozen ground (tjdle, dauerfrost, eisboden,
permafrost, merzlota, pergelisol, etc.) has always been close-
ly associated with the study of patterns in the vegetation
and in the ground. The features of patterned ground were
recorded in the nineteenth century from arctic regions by
geographical explorers (Adams, 1815; Von Baer, 1837/38;
Beechey, 1831; Figurin, 1823; Léven in Chydenius, 1869;
and Von Middendorf, 1864-67), and by botanists (Kjellman,
1879 ; Martin, 1882). Then Hégbom (1914) laid the founda-
tion of the study which has recently become of broad inter-
national importance. Crampton (1912) and Thoroddsen
(1913) recognized the importance and interrelation of
microtopography and soil processes with vegetation. Frédin
(1918) made a detailed study of the relation between earth
and tati and defined most of the ideas
basic to duditsGuntiig the effects of frost microtopography
on vegetation. The study received a great stimulus with the
excursion to Spitzbergen of the IXth International Geo-
logical Congress led by De Geer (1910) as shown by the
flurry of publications which followed (Meinardus, 1912;
Miethe, 1912; Penck, 1912; Sapper, 1912).
Meinardus (1930), Poser (1931, 1932, 1933), Steche
(1933), and Cailleaux and Taylor (1954) published litera-
ture reviews and th of
producing what Nordenskjéld (1909) first named Struk-
turboden. I agree that this term is well translated (if
necessary) as “patterned ground” (Washburn, 1950).
Washburn’s reviews (1950, 1956) define the problem, and
his discussion of the mechanisms makes another review
did “ay deal with non-patterned
ground including solifl 1906), earth
mounds, turf- and stone-banked pan and block streams.
Steche (1933) reviewed and defined the terms (primarily
German usage) for the various expression of frost action. —
4 WILLIAM H. DRURY, JR.
Troll’s (1944) review included appreciation of the intimate
mauntual associtaion and influence of frost features with their
, and established the climatic correlation of the
major types.
Major frost patterns are not controlled by vegetation,
although minor features may be in some places, such as in
mid-latitude mountains (Denny, 1940). Péwé (1959)
showed that fully-developed frost-crack polygons (Spalten-
boden) are present around McMurdo Sound in the Antarctic
in absence of veg But tation cover and exposure
may suppress eoiinetion (Fri Sah 1918; Sigafoos and
Hopkins, 1952) and strongly influence modification, once
the major Strukturboden patterns are formed.
Antevs (1932) introduced the study of alpine frost fea-
tures and their influence upon vegetation to the United
States, and Polunin (1934) first used patterned ground to
classify vegetation. The study of the geobotanical develop-
ment of bogs in northern Europe has extended the knowl-
edge of soil processes associated with frost action into the
boreal regions (Cajander, 1913; Auer, 1920), although
recent Scandinavian bog studies (Sjérs, 1948, 1950 a &
b, 1959), and alpine vegetation studies (Dahl, 1951;
Nordhagen, 1943, 1954; Gjaerevoll, 1950, 1956) haye con-
centrated on biological ph require-
ments and effects of climate and es plants — — rather than
on soil features.
In America, Raup (1947, 1951) has led the study of the
- of soil movement on the growth of plants and vegeta-
tion patterns. His (1947) study of the McKenzie Moun-
tains used soil forms to classify alpine vegetation, and
similar work has been carried on in the New World chiefly
by his students (see, however, Hanson, 1950; Billings &
Mooney, 1959; Billings & Mark, 1961; Wiggins, 1951).
: and Sigafoos (1950) studied the combined effects
of frost action and plants in the wet or low tundra (Von
Middendorf’s term) region of Seward Peninsula, Alaska.
Sigafoos (1951, 1952) discussed detailed effects of frost
processes on getati
i.
in forested parts of Alaska and the
SOUTHERN BYLOT ISLAND 5
importance of physical factors. Drury (1956) discussed the
interaction of frost action, alluviation and vegetation on a
bog-covered floodplain in interior Alaska. Denny (1951,
1956) and Goodlett (1954) applied these disciplines to the
effects of periglacial climates on vegetation and to interpret-
ing the climate from the frost effects still visible at the
margin of the most recent ice in Potter County, Pennsyl-
vania. Most American studies of periglacial geology have
grown out of the influence of Kirk Bryan (1946, 1949) who,
with his students (Denny, 1936, 1938, 1951; Horberg, 1949;
Sharp, 1942 a and b; Schafer, 1949 ; Smith, 1949; Smith and
Frazer, 1935), joined with students in central Europe in the
study of ground conditions during the maximum Wisconsin-
Wiirm ice advances (Biidel, 1944; Cailleaux, 1942; Diicker,
1933, 1951; Gripp, 1952; Hégbom, 1923; Kessler, 1925;
Poser, 1947; Schenk, 1955; Selzer, 1936; Soergel, 1921,
1939, 1940, 1943; Tavernier, 1944-45). Out of these studies,
and the study of bog pollen and other fossils, they project
climates of the period (Willett, 1953; Zeuner, 1937), and
even map the vegetation (Biidel, 1949; Frenzel & Troll,
1952) and follow its post-glacial changes (Firbas, 1939;
Aario, 1944). Elsewhere (Drury, 1956) I have discussed
why I think most American periglacial geology studies do
not give enough consideration to: (1) the climatic differ-
ences at the margins of the ice which existed between the
Old World and the New; (2) the changes in precipitation
and temperature which caused the ice to stop its advance;
(3) what vegetation would be under climatic conditions
which clearly do not occur anywhere today.
My studies on Bylot Island examine the interrelation of
plants and soil structures in a high arctic region of dry
tundra (Von Middendorf, 1864-67) where land surfaces
are so recently exposed that all forces whose traces can be
found in the soil must be active at present. Thus, we cannot
postulate changing climate as an escape mechanism, when
we are unable to explain land forms using only observable
forces. My interpretations are based upon detailed study of
many, repeated excavations in the thawed surface soil with-
in a mile of our camp, and a limited survey of surface
expressions within ten miles of camp. The study concen-
trates upon: (1) non-sorted circles (Washburn, 1956) (mud-
6 WILLIAM H. DRURY, JR.
boils, frost-boils, frost-scars); and (2) formation and
modification of a four-sided pattern of frost-cracks.
Climatic Requirements.
Troll’s (1944) review showed how very different are frost
patterns in different climatic regimes. As he says, frost
features are not restricted to perennially frozen ground, nor
to humid climates, but only to regions where, periodically,
there is strong and morphologically effective ground frost;
thus patterned ground is not an arctic or a polar, but a
rge forms are characteristic of
areas of perennially frozen ground, extreme cold, and few
annual temperature changes which cross freezing ; miniature
forms are characteristic of high altitudes in low latitudes,
snow-free for only a short period, and then subject to many
shallow temperature fluctuations across freezing. Troll
points out that sorted patterns and hummocks are charac-
teristic of oceanic tundra regions and frost-crack polygons
are typical of continental cold, dry regions.
Hégbom (1914) emphasized that the frequency of freeze-
thaw may not be the only feature of primary importance.
If it were, the Lofoten Islands on the west coast of Norway
should have maximally-developed frost features, as would
most of the so-called temperate climates. He suggested that
the following are important: (1) the number of degrees
the temperature falls below freezing; (2) the number of
months during which the temperature is below freezing;
(3) the depth of frost penetration; (4) the presence of a
sounding board (perennially frozen ground, ice, or bed-
rock) ; and (5) the presence of abundant moisture at the
period of frost activity — whether spring or fall.
Statement of the Problems.
1. How do feels and depositional-slope processes modify
raised these actions expressed in changes visible in
the course of a oe season? Are frost forms sufficiently different
= Sigrcorscnedre older surfaces to give a measure of the progress of
- Pg Some force creates an upwelling of either fine- or coarse-grained
1925; Gripp, 1928, 1929). Are
i : . cryostatic mechanisms suggested to
produce both sorted patterns (Hogbom, 1914; — and Odell, 1924;
913; Frédin, 1918;
SOUTHERN BYLOT ISLAND 7
Washburn, 1950, 1956; Hopkins fe Sigafoos, 1950; Sigafoos, 1951)
strong ge and active locally
3. Many of the forces saps for the development of patterned
ground oe concerned directly with sorting soil fines from coarses,
but many of these forces must be questioned for adequacy of their
strength. Furthermore, because the forms we found were all non-
sorted, can these mechanisms be used, even with modifications, to
explain the patterns we found?
4. (a) Can the mechanisms suggested for the wet ems of
Spitzbergen, where the ground may be wet during autumn freeze-up,
apply where the ground is dry to four feet by mid-July? (b) Do t they
act at the time of year (spring) in which we watched the frost
features in action? (c) What relation does the cheese-like quality
of the drying mud column and the swelling of the tacky surface have
to the now generally discarded hypothesis (Steche, 1933) of colloidal
activity? (d) What part do swelling or upwelling forces in water-
cp Saker soils play in creating or modifying solifluction forms?
Some mammoth force is able to crack open the ground on
bruks geet surfaces, even under the pounding sea surf. Is
Von Bunge (1884, 1887, 1902)-Leffingwell (1915, 1919) ispotheais
of frost-crack growth adequate to explain this large pattern? Taber’s
(1929, 1930) study of the segregation of ice into lenses and wedges
in the pen made Hos tension-crack theory tenable and greatly
advanced o1 of the devel t of ground ice. But
the RES at Bylot did not contain ice. They contained coarse
sediments. On the other hand, is the theory of tension-cracks due
to especially low temperatures adequate to explain the forces creating
the broad uplands they create ghiPeorey moss-colonized frost-cracks?
7. What forces (exposure, moisture, sun, sequence of snow-melt,
soil instability) influence and control the distribution of plants? Are
regional differences greater than microtopographic ones? How do other
organisms respond to Parag associations? part does plant
cover play in soil d id the ification of frost features?
8. Is the vegetation ery different on successively older
saline to suggest progression, development, or regional vegetation
PART ONE
THE BYLOT ISLAND EXPEDITION
We made our studies between 12 June and 29 July 1954,
on the south coast of Bylot Island at the mouth of the
Aktineq River. Maps 1 and 2 show places mentioned.
Scherman (1956) gives a delightful narrative of the expedi-
8 WILLIAM H. DRURY, JR.
tion, emphasizing travel over Eclipse Sound and life with
the community at Pond Inlet. The Drurys (1955) give a
short account of the weeks at Main Camp. The expedition
was paid for by a private grant with help from the New
York Zoological Society. We made arrangements with
Canadian officials through The Arctic Institute of North
America.
Dr. Benjamin Ferris and Edward Ames gave unlimited
help in the field. Ames dug many of the holes, and his
unbiased, critical mind effectively deflated many hypotheses.
Four papers on the biology of the breeding birds of Bylot
Island have been published. The present paper, together
with Miller (1955), Ferris (1958), and Van Tyne and
Drury (1959), completes the reports.
Bylot Island is about 120 miles long and 60 miles across;
the mountainous backbone of the island rises to a height of
600-700 feet, underlain by Paleozoic, igneous and meta-
1oT ISLAND AND ECLIPSE SOUND. Snow fields are represented in black.
Place names referred to in the text are shown.
morphic rocks (chiefly quartzites or granitic, with many
dark minerals). This central mountain area is covered with
a snowfield, from which about one hundred glaciers flow
down to the lowlands on the northeast and southwest sides
(Map 1). Our camp was seven miles from the Aktineq
Glacier’s snout, and on the east edge of the southwest
SOUTHERN BYLOT ISLAND 9
plateau area underlain by poorly consolidated Tertiary sand-
stone containing some low-grade coal seams. These sedi-
ments dip gently north. The plateau extends about 50 miles
west from camp to the shores of Navy Board Inlet; and it
slopes from about 700 feet at the foot of the mountains to
sea level, except near our camp where it ends at the top of
200-foot bluffs. Although frost disturbance and movement
are evident in all soils, the major features of erosion are pro-
duced by a combination of cryoplanations and violent stream
action during the spring thaw. This topography supports
Peltier’s (1950) concept of a cryoplanation-erosional scheme
or cycle as an extension of the Davisian (1909) cycles of
erosion. It agrees with And ’s (1906) discussion of the
relation of solifluction to the removal of detritus by running
water, and the combination of the two as an especially effec-
tive erosion agent. The rolling plateau surface deeply cut
by the shallow stream valleys, and the ankle-high vegetation
make the topography resemble that of the high plains of
the United States, and of the surfaces of most North Ameri-
can “uplifted and dissected peneplains.” Many areas of the
uplands show regular lines of vegetation alternating with
barren soil (Figure 1). Many ponds are regular, both in
outline and distribution, and occur on the tops of the hills
(Figure 17); many slopes are deeply scarred with frost-
cracks (Figure 8), or pitted with mud-boils (Figure 7)
(Washburn 1950, 1956), but frost action on most of the
uplands is inconspicuous, being expressed only as a polygo-
nal pattern of shallow depressions where mosses thrive.
The Study Area.
Map 2 indicates the hic structure of
the square-mile area studied in detail. “We pase the topog-
raphy into five parts: (1) the ridge running north and
south of the west margin of the area — West Ridge; (2)
the beach, sixteen feet above present high tide, at camp —
16-foot Beach; (3) the steep slope and rounded hill —
Kungo Hill (from the Eskimo name for Snow Goose, Anser
coerulescens atlantica) ; (4) the gently-sloping surface —
Plover Plateau; and (5) the shallow mound — Tui-Tui
Tabletop (from the onomatapoetic Eskimo name for Baird’s
Sandpiper, Calidris bairdii). Iceberg Lake is the northern
margin of the study area; the Little River runs north of
10 WILLIAM H. DRURY, JR.
attern suggests that vegetation
lines of developing drainage. Vegetation stripes around the boulder
are parallel. Vegetation is chiefly Avens and Arctic Willow.
east of Ooyarashukj
MELT-WATER RILL. y sand deposits are in the shape of a soli-
fluetion lobe. Vegetation on the lip of the lobe is chiefly sedge and Arctic Willow.
E:
peet.
xeavations (Figure 19) show that many lobes originate in this way. Photo taken on
the south slope of Kung:
o Hill at about 70-foot elevation.
SOUTHERN BYLOT ISLAND 11
Plover Plateau, and Golden Plover Creek runs south of it,
next to Kungo Hill. Both these creeks ran five to six feet
deep, and moved rocks two feet across during the height of
the thaw, but were completely dry by the end of July.
Raised beaches are hatched.
the east side of West Ridge from the 150-foot level.
Areas of sand, damp at low water, are stippled.
Most of Tui-Tui Tabletop is scarred with frost-cracks
(Figures 16, 17, and 18). The prevailing east wind keeps
large areas of it barren of vegetation and the surface is so
churned on the windward side that the major frost patterns
are nearly lost (Figure 18). Plover Plateau is covered with
12 WILLIAM H. DRURY, JR.
a mat of mosses, Avens, Arctic Willow and Bell Heather.*
At the upper end is a marsh, and at the lower end several
ponds (Upper Phalarope Ponds) in another marsh. Kungo
Hill is barren at the top and comparatively heavily vege-
tated on sheltered south slopes. Weedy vegetation covers
disturbed soils (“frostings”), especially on the raised sea
beaches on the south and east slopes of Kungo Hill and
Plover Plateau. The 16-foot beach level at camp slopes
gently up to a 35-foot beach, and is deeply scarred with
frost-cracks modified by the growth of mosses into an order-
ly net of peat ridges, many of which act as dikes and hold
in rectangular ponds or sedgy marshes resembling rice pad-
dies or cranberry bogs (Figures 22 and 23). Loon Pond, the
ponds on the west side of Tui-Tui Tabletop, and both groups
of Phalarope Ponds are rectangular, resulting from thawing
of the frozen ground below such places (“thaw sinks” —
Muller, 1947; Wallace, 1948; Black and Barksdale, 1949;
Hopkins, 1949).
The summer of 1954 was late; the sea ice never left
Eclipse Sound. When we arrived on 12 June, eighty-five per
cent of the land surface was still snow-covered (Figure 1
in Van Tyne & Drury, 1959), but already Opposite-leaved
Saxifrage was in flower. The general thaw started 24 June;
sheets of melt-water flooded the uplands 26 June, and most
of the uplands were free of snow by 28 June. As soon as
the snow left slightly sheltered areas, Bell Heather (con-
spicuous in such places) came into flower; most species
bloomed in the last days of June and the first three weeks
of July. Early flowering plants, such as the Cruciferae, had
set fruit, but the sedges and grasses and most of the plants
of the uplands were still in flower, or their flowers were
just withering on the first of August when we left.
Intense frost churning was in progress in the thawing,
water-saturated, snow-free areas when we arrived, but
under the snow the soil was frozen and undisturbed. The
Most evident frost action occurred on exposed surfaces
lying between snow patches, during the period 15-25 June.
As the surface dried after the first of July, frost activity
_*A taxonomic treatment follows the discussion of the vegetation, and in it seientifie
common names are used in this part of the text.
:
SOUTHERN BYLOT ISLAND 13
PART TWO
GEOLOGICAL PROCESSES
It is convenient to divide frost action into three descrip-
tive categories: (1) those affecting the rapid mechanical
weathering of rocks —frost-riving; (2) those concerned
with down-slope movement — solifluction; (3) those pro-
ducing patterned ground or Strukturboden. The mechanisms
involved grade into, combine with, and influence each other,
as Hégbom (1914), Elton (1927), Washburn (1956) and
others have emphasized.
I. MECHANICAL WEATHERING
Frost-riven stones. We exhumed stones shattered into many
pieces from many excavations, especially those on wet
beaches. Either the sections were still in place or they
were separated by a centimeter or more. (Fig. 18, Hégbom,
1914, p. 317).
Wind-sculptured. stones. Wind-cut stones are numerous on
raised beaches, especially on the pavement of the beach east
of the Aktineq River. These stones have been cut by sand
driven by the prevailing east wind so that nearly flat upper
side slopes down to the east and the anastomosing flutings
point to the east. We found no Drikante. The polish on the
cut surfaces of the stones (chiefly sandstone pebbles) is
dark and shows little glisten.
Avalanche gullies in the sedimentary outcrops. Steep cuts
in the bluffs west of camp have half-rounded, gutter-like
bottoms, from which avalanche scouring has gouged coarse
debris. The debris collects in tongues which often bury
masses of ice at the foot of the gullies. These may develop
when frost-riving products of several years are released in
the spring by sun-melt, or early spring/late winter melt-
water refreezes in the valleys, and gather snow to produce
a glacier-like mass of ice and stones made unstable by later
thawing.
Grenadiers. Hégbom described frost action on mountain
slopes creating stone towers (Felsentiirme). (See Fig. 6 on
p. 282 of his 1914 paper). Similar structures are charac-
teristic of the ridges on the bluffs west of camp. The
castellations — 3-25 feet tall — extend from the shore at sea
level to the top of the bluffs at about 200 feet.
14 WILLIAM H. DRURY, JR.
Block streams. We found block streams 30 yards across and
100 yards long on the depositional-slope-covered margins of
shallow valleys seven miles inland on the west side of the
Aktineq River. They are at the base of the bedrock out-
crops and, because they are heavily lichen-covered, are
presumably no longer active. Because they are at the same
level as the solifiuction products which fill these shallow
valleys, I presume that they are of the same age. The creek
draining the west side of West Ridge runs several feet
below the surface rocks of a block stream.
Il. SoLIFLUCTION
Down-slope movement of soil is everywhere conspicuous.
It has rounded the hills of the plateau region and filled valley
sides with debris so that there is little sharp topographic
contrast. Motion down the valley sides has produced con-
spicuous lobes and low terraces. Does the entire soil mass
move in the formation of slope features? Or is the motion
the result of minor alluviation in which mud bursts out from
under the vegetation mass and flows as a mud river? What
are the effects of these two types of movement upon the
vegetation cover?
Fisher (1866), Kerr (1881), and Davison (1888, 1889)
ted on t of soil on slopes associated with
frost. Wood (1882) showed the importance of frozen
ground as a water source to saturate the soil, and the impor-
tance of this in soil flow. Andersson (1906) first suggested
that frost-riving together with what he named solifiuction
are especially effective erosion agents in arctic regions.
Hégbom (1914) separates Fliesserde from ausfliessungen.
Frédin (1918) recognized the difference in these two move-
ments. Beskow (1930) separated the slow movement Erd-
fliessen at the surface from rapid mud-flow Erdrutsch,
especially at depth. Thus from the beginning, there is
agreement that there are two major types of soil movement
which we may call “creep” and “mud-flow.”
The chief difference seems to be how much movement is
related to the freeze-thaw cycles operating with a down-
slope component with each thaw which Taber (1943)
thought especially important, and how much is related to
__ the flow of waste saturated with water which Andersson
(1906) first emphasized. Clearly, both are active and grade
SOUTHERN BYLOT ISLAND 15
into each other. Frédin (1918) described accurately the
surface expressions of these forces and the effects on vege-
tation in Swedish mountains. Sigafoos and Hopkins (1952)
did the same for Seward Peninsula, Alaska.
Frost-Heaving.
Hamberg (1915) clarified that the thrust is straight
upward unless the force resulting from frost-heaving is con-
fined; and Taber (1929, 1930, 1943) showed that the
mechanism is largely due to growth of ice crystals in the
soil at right angles to the cooling surface. A ubiquitous
expression of this action is the swelling of the topmost layers
of the soil so that they pull away from large rocks (illus-
trated in Hégbom, 1914). This is a common phenomenon
in New England in late winter and spring, and was con-
spicuous on the muddy surfaces on the uplands of Bylot
Island when a sharp freeze occurred during the thaw. As
the thaw penetrates the top few inches of the soil, that soil
becomes saturated with melt-water, and thus ready for
maximum expression of frost-heaving by segregation of ice
crystals on refreezing. Large stones seem to sink when the
ground surface refreezes. It is not clear how this happens
but it may be that: (a) on refreezing, the surface soil forms
a stiff deck over the still unfrozen ground below, and this
deck is pushed up and away from the stone. Being connected
to the thawed layer below, the stone retains warmth during
a short period of freezing, and thus is not frozen into the
deck of soil and heaved up with it; or (b) a large stone
extends down into the still frozen layer and is held there
while the thawed surface layer heaves on refreezing. The
iation of these with a sharp freeze during
spring or mid-winter “thaw is important because of the
implications it suggests as to the conditions under which
related solifiuction features develop.
Solifluction Forms.
(a) As soon as solifluction lobes were visible, we put
white flowerpot sticks into the lobes, perpendicular to the
soil surface and in straight rows along and across them, to
test for any motion. Displacement of just under an inch
occurred between 25 June and 29 July, and amounted to a
gradual sag in the fluid material at the center of a mud-boil
16 WILLIAM H. DRURY, JR.
which contributes a net displacement of the solifluction lobe.
Washburn (1947) surveyed solifluction lobes on Queen
Victoria Island and found movement of one and a half inches
during June —the height of the thaw — and one-quarter
inch during the following four months.
(b) Large boulders on the top of the uplands or terraces
stand out on the surface as if they have either been frost-
heaved or the upland has been moved away from them. On
slopes, the soil overwhelms boulders from above, flows down
the sides, and moves away from below.
(c) Creep. Excavations down the long axis of some soli-
fluction tongues or lobes show organic material and vegeta-
tion mat turned under — the result of a slow movement of
the whole soil-mass. Antevs (1932) and Sharp (1942a) sug-
gested that the vegetation mat acts to slow up solifluction,
hence to produce garlands or turf-banked terraces.
(d) Mud-Flow. In contrast, several excavations (Figure
19F) cutting longitudinally through solifluction lobes showed
the old soil profile was not disturbed, but buried. In many
places on the surface, fresh deposits show a small delta of
mud built out over previous years’ vegetation (Figure 2).
These deposits also take the form of solifluction lobes. Satu-
rated soil has burst out from under the vegetation mat and
flowed over the surface as mud or “slud,” and then when
stopped by low slope, etc., has been subsequently colonized
by vegetation. Frédin (1918) emphasized the force which
pushes soil up through the vegetation mat and out over the
surface to create the lobe and to keep it moving. Hopkins
and Sigafoos (1950) discussed this when dealing with mud-
boils and suggested, as did Frédin, that it is the result of
forces which develop when wet soil freezes. Neither paper
discussed the place of this upwelling in solifluction or the
combined effects of freezing forces and mud-flow forces. I
am convinced that the combination is very important on
Bylot Island.
In small displacements, a general soil-sagging (creep)
moves a large total mass of soil a short distance, but the
usual p surface ion depends upon mud-
flow out from below through the vegetation and over the
surface. Evidence from our excavations supports Andersson
(1906) and Hégbom (1914) that the soil does not move
SOUTHERN BYLOT ISLAND 17
along the surface of the perennially frozen ground as a slip-
ping plane, as Beskow (1930), Steche (1933), and earlier
authors such as De Geer (1910) believed. Instead, the
motion depends upon viscous or fluid movement at all levels
of the soil saturated by rapid snow-melt. Under these cir-
cumstances, runoff is contained above a still-frozen layer,
such as annual frost or perennially frozen ground, which
serves to conserve moisture.
Vegetation Effects. The effects of microtopography of a
solifluction lobe on the vegetation vary according to its loca-
tion on the lobe: (1) on the center of the lobe running
down to the lip, is a relatively barren area scattered with
blackened mosses and lichens ; (2) most of the top of the lobe
is covered with istic of dry uplands.
Frédin (1918) took pains to show that this barrenness is
not the result of wind exposure but of soil instability result-
ing from creep and mud-flow. Moisture held through mid-
summer in organic soil at the margins and in the shelter of
the lobe encourages the growth of mat plants. In such
places we found southern plants otherwise unusual in the
uplands, such as Crowberry, Bilberry, Netleaved Willow,
and Bell Heather.
Sapper (1913) believed that solifluction is more frequent
in arctic regions because the thin vegetation does not
restrain it; Hégbom (1914), on the other hand, believed
that the thinness of the vegetation is caused by disturbance
of the soils associated with solifluction. Sernander (1905)
and Frédin (1918) —and after them Seidenfaden (1931)
and Sorensen (1935) — agree that the two work together.
Solifluction forms, and the forces which produce them,
grade into the forms and forces discussed below under non-
sorted stripes and circles.
Depositional Slopes.
Depositional slopes (Péwé, 1948, 1950) form gentle (2°-
5°) slopes from one raised beach to the next. The slope from
the 55-foot beach to the 35-foot beach is long and smooth,
and almost completely covers the lower beach. We found
peat ridges, mud-boils, small frost-cracks, well-developed
soils, and rich vegetation on the surface of depositional
slopes. The large frost-crack pattern discussed below is
not expressed on the surface of depositional slopes, but
18 WILLIAM H. DRURY, JR.
during June and July 1954 the frost did not retreat deeply
enough for us to dig through the depositional slope material
to find out whether it lies buried underneath. Depositional
slopes are continuous with the stream deltas near Golden
Plover Creek, near Snow Bunting Creek, and the creeks
east of Golden Plover Creek, formed when the beaches
involved were at sea level. Streams have since cut sharp-
sided valleys into these deltas and have re-deposited a delta
on the present beach level.
Excavations through the depositional slope material into
the sediments below will give important information (1) on
dating the formation of frost-cracks and the development
of depositional slopes, (2) on develop t of vegetation on
progressively older surfaces, and (3) on processes of modi-
fication of the major frost-crack pattern with the passage
of time.
The plateau surface and uplifted lains in t te
regions.
The concept of an uplifted, dissected peneplain has long
been used to explain or to illustrate geographical history and
development of land forms. Davis (1889, 1909) introduced
this contr ial ption in the Appalachian Mountains
area. Denny (1956), however, presented cogent arguments
against the existence of peneplains and concordance of
summits in the Appalachian ridges in Pennsylvania where
he studied. His re-creation of the “original” surface showed,
in general, accord of summits with structure, and his evi-
dence indicated effective erosion during several stages of the
Pleistocene.
Tf studies of frost action and cryoplanation processes
observed in the North (Andersson, 1906; Bryan, 1946;
Peltier, 1950) apply validly to uplands in temperate regions
during the periglacial climatic conditions, those uplands
exposed to the most powerful forces of cryoplanation.
SOUTHERN BYLOT ISLAND 19
steep valleys of streams or valley glacier cirques (maps and
photos in Antevs, 1932; MacClintock and Twenhofel, 1940).
Tree line often occurs at the break in slope between the
rolling plateau and the steep valleys. Raup (personal com-
munication) suggests that this line reflects the lower limit
of intense frost action and soil instability.
These mountain tops resemble lowlands of the Arctic
created by cryoplanation, combining solifluction with mas-
sive erosion by snow-melt-water.
When mid-latitude uplands were above the main mass of
an icecap or below a snowfield during an ice readvance, the
uplands were subjected to vigorous arctic frost erosion,
whose local base-level was the top of the surrounding ice
sheet or the lower limit of deep frost penetration and snow-
line (Troll, 1944).
This process may be altiplanation (Eakin, 1916). It sug-
gests that uplands at the tops of the Appalachians and
eastern Canadian mountains may have developed in situ by
rigorous high latitude or altitude erosion and do not require
recourse to the intricacies of regional uplift (Matthes,
1959). The presence of tablelands, then, may indicate a
period of rigorous ice-free climate in the schedule of the
development of the landscape.
III. STRUKTURBODEN — THE DEVELOPMENT
OF PATTERNED GROUND
Non-sorted stripes and non-sorted circles seem to be
formed by similar mechanisms, but we found marked differ-
ences in the type of soils, depth of soils, and relation to
other frost patterns.
TABLE I
Non-sorted Stripes — Vegetation Non-sorted Circles — Mud-Boils
tripes
or Soil St
1. On gentle to steep slopes— 1. On gentle slopes or terracettes
by early July. Although —wet late into season. Melt-
with water during water forms puddles
the thaw, do not “sit” in it. stands on the surface during
2. Soil of angular sand and the thaw.
pebbles, with little cohesion. 2. Soil often dark, usually fine-
Vegetation lines running grained, showing tilth; but
down-slope, but not held in at sometimes centers are of sand.
20 WILLIAM H. DRURY, JR.
Non-sorted Stripes — Vegetation Non-sorted Circles — Mud-Boils
Soil Stripes
the lower end by vegetation 3. If elongated down-slope, lower
or peat. In some places pat- border margined by peat ridge
terns resemble rill-work. which merges into solifluction
4. Occur on rather exposed, high, lobes.
rounded uplands in shallow 4. Occur on south-facing shelter-
soils; covered with snow until ed terraced slopes or valley
main thaw, or blown free of bottoms in deep soils; free of
snow and therefore exposed to snow early in season and thus
deep freezing. exposed to long freeze-thaw
5. Cracks appear parallel to the season.
Stripes. These cracks are 5. Polygonal cracks appear on
wide, deep, and long, as if the the surface as it dries; column
result of a strong force. pulls together as if by local
6. Soil stripe has convex surface. hesion.
7. a) Fines erupt in small 6. Earth column tends to bulge
patches which are elon- upward into convex top —
gated down-slope. Some tacky and cheese-like as it
fines are thrust up through starts to dry.
eracks, 7. On boils on south-sloping up-
b) Micropolygonal pattern of lands, small eruptions appear
eruptions appears where in regular patterns.
stripes are subject to water
seep, such as from a marsh.
A. Non-sorted stripes — vegetation or soil stripes
We found these only outside our study area. They consist
of lines or patches of (1) bare, actively-disturbed tan soil,
(2) less disturbed margins scattered with lichen thalli, tiny
wi mosses, Saxifrage and Draba, and (3) vegetation —
chiefly Avens, mosses, Arctic Willow, Locoweed, and
cushions of Opposite-leaved Saxifrage.
soil during the thaw, and later the hole had nearly filled
while the mounds had slumped, dried and cracked. These
large eruptions are few and occur where surfaces are scat-
tered with flat, angular stones, and disturbed areas are part-
ly colonized by plants; (b) an inch across and three inches
long, where there is fine sand, silt and angular
pebbles, and often associated with cracks running down-
___ Slope; (c) one or two inches across, forming a small mesh
_ in fine, sandy (often cheese-like) silts. These are covered
SOUTHERN BYLOT ISLAND 21
with tiny moss or lichen thalli and resemble miniature
Zellenboden (Hiégbom, 1914, p. 320).
(2) Many vegetation-free soil stripes have cracks one to
two inches wide and five to twenty yards long running down
the center (Figure 11). In some places eruptions of damp,
fine soil are thrust up through these cracks.
(3) Most vegetation lines are underlain by a shallow
humus layer, but many have Locoweed whose deep taproot
suggests deep soil stability.
(4) Some vegetation lines have a moss-filled depression or
crack running down the center.
(5) Some vegetation lines converge where two gentle
slopes come together, which suggests rill-work (Salomon,
1929). (Figure 1).
These observations suggest:
(1) that disturbance of the bare areas may be associated
(a) with soil cracks, (b) with fluid upward flow of water-
saturated soil (Frédin, 1918; Gripp, 1926, 1929; Steche,
1933) ;
(2) that the crack or depression (which may be of inde-
pendent origin) now covered by vegetation is stable. Hog-
bom (1914) suggested that solifluction is closely associated
with the production of soil stripes, and so did Sigafoos
(1952) who found vegetation stripes on obviously unstable
soil. Rill-like patterns where vegetation lines converge sup-
port Huxley and Odell’s (1924) hypothesis of water-flow
through the coarse stones in sorted patterns first established
by frost-heaving. Steche (1933) suggested the combination
of the upwelling force with solifluction (see his diagram
after Gripp, 1926). Most German authors recognize two
separate processes and suggest that modification by solifiuc-
tion must follow establishment of the sorted pattern.
Our observations suggest that (1) modification is con-
temporaneous; (2) shallowness of the soil prevents (a)
formation of solifiuction lobes, and (b) growth of vegetation
which may obliterate the pattern on deep soils.
B. Non-sorted circles — mud-boils, frost-boils, or frost-
sears (Cf. Fleckentundra, Troll, 1944).
As the snow melts, tufts of grass come through first, then
the ground appears next to clumps of grass or Bell Heather;
but the ground in such a place remains frozen to a depth
22 WILLIAM H. DRURY, JR.
+ SUBSURFACE WATER-FLOW INTO MUD-BOIL EXCAVATION. Water flowing
through the soil at least 3 inches below the surface has filled the 18-inch excavation.
ows in the surface foam. Vegetation chiefly sedges and Arctic Willow
ground cover. Site on the south slope of Kungo Hill at about
Field notebook gives si
MOC
size.
" - Mud-boils, showing surface cracks, mineral soil
core, and separated by moss-filled depressions are transitional between mud-boils 2
and hummocks. V izes. Site
slope above the
Vegetation chiefly Avens and
t beach at Ooyarashu:
SOUTHERN BYLOT ISLAND 23
of 2-4 inches while some bare areas quickly thaw to 9-18
inches. These deeply-thawed spots are the nuclei of mud-
boils. I found no way to forecast where they would appear,
except that they are of uniform silts or sands and have few
stones on the surface.
After several days of thaw, mud-boils have a slick, shiny
surface when soaked with water, or a tacky, cheese-like
surface when drying. They are still thawed deeper than
the surrounding soil. How do these soft spots develop?
What happens to the sediments inside from day to day and
through the course of spring’s saturation and summer’s
drying? Our observation showed the following:
(1) Some contain more fines than their surroundings, and
on daily drying pull together into a cheese-like mass. Others
contain coarser materials than the surrounding soil. Some
consist of a column of fines topped by a half-inch layer of
sand. Most boils have a concentration of small, angular
stones near or at the surface, but very few stones within the
earth column; angular, slab-sided stones, 1-5 inches long,
may occur on edge in surface cracks (Figure 5).
(2) Most, when dried out, show disturbance of horizontal
strata on the edge of the column, and soil remnants (partly
organic) turned on edge or floating in the column (Hopkins
and Sigafoos, 1950).
(3) At the height of the thaw they are continuous with
a subsurface flow of water (as Thoroddsen, 1914, observed),
and quickly fill with water while being excavated — but no
water flows out the top. In one case, clear water flowed into
a trench from under the annual frozen layer (Figure 3).
(4) When wet and still active, recent footprints on their
surface disappear, and a trench 8 inches deep is usually
filled and obliterated overnight.
(5) While drying but still damp (water can be squeezed
out), the surface soil draws together and cracks — first on
the up-slope edges of the boil as if sagging, or around stones
as if drawing away. As drying of the soil proceeds, while the
surface is still damp, pentagonal cracks appear usually about
5 inches apart, extending into the soil 1-3 inches (Figures
4 and 5, and Rousseau, 1949). These cracks are damper than
the meshes.
(6) During June, when the thaw has generally reached
24 WILLIAM H. DRURY, JR.
a
tes
ce
Say
IGURE 5. ARCTIC WILLOW AND MUD-BOIL WITH CRACKS. Contraction crack polygons
mud-boil Angular stone on edge in the crack suggests the
garla: ettes.
he stem and some roots. mete ota taken on south slope of The Bluffs at
about 100- fe. elevation
FIGURE 6. CONTRACTION CRACKS ON POND sHore. Algal growth makes scum on mud
SOUTHERN BYLOT ISLAND 25
about 6 inches on sunny south slopes, the surface freezes on
many nights, forming a crust 1/4-2 inches thick; short ice
crystals form on muddy places. Repeated freezing, above a
thawed and water-saturated soil layer, breaks up the soil
tilth and creates a mobile mud. A daily cycle in which the
mass is drawn together in the early morning, filled out and
soaking wet by mid-morning, is repeated over the period of
a week or ten days. Every day the same cracks come and
go, and the soil changes from quicksand to a jelly. When
the jelly forms and the columns swell, or when the surface
freezes, the margins of the vegetation are thrust up, raising
a ridge of black soil and vegetation (cf. Hopkins and Siga-
foos, 1950). The movement of the mass down-slope pushes
up a larger ridge on the down-slope side, upon which moss
growth is lush.
(7) Bleached stems (occasionally roots) of living Arctic
Willows cross the surface, and some roots cross the soil
column two or three inches below the surface. The green
photosynthesizing part of the plant lies beside the boil (Fig-
ure 5) as if the plant grows and produces leaves where there
is less disturbance, while subsurface parts, once rooted, can
persist even when a boil develops. Perhaps the age of such
a plant can be used to measure the age of a boil.
(8) The flowerpot labels which we placed across mud-boils
during the peak of activity, suggested down-slope movement
at the center, but no indication of movement from the center
toward the edge (Washburn, 1947).
(9) Step-like terracettes containing mud-boils show forms
transitional to turf-banked terraces and solifluction lobes
(Figure 5), and suggest functional relation of all of them
via the upward and outward movement of saturated soil
(Andersson, 1906; Hégbom, 1914; Frédin, 1918, Beskow,
1930; Hopkins & Sigafoos, 1950; Hanson, 1950; Warren
Wilson, 1952).
I agree with Washburn (1956) that frost-boils are frost-
sears (Hopkins & Sigafoos, 1950), but I believe that upwell-
ing of fines is necessary for their formation. Ber;
surface where scattered Puccinellia and ee stents — and lichen thalli
Bro sinuous pattern d in
formed by forces pulling meshes apart, not fee isis my art. The present erack
on one race suggests that the site of the crack is perennial but that the
“xact location may vary two inches. The crack is in black organic fine sand with
26 WILLIAM H. DRURY, JR.
(1912) described mud-boils with loamy centers and the
vegetation mat lifted at the margins. He suggested that
saturated soil is thrust up in the center and bursts through
the vegetation cover. Thoroddsen (1913), in explaining
hummock formation in Iceland, suggested the segregation
of crystaline ice into lenses just under the surface of bare
or raised areas both updomes (swells) the surface (Figure
4) and draws up water from damp, unfrozen soil layers to
allow further growth of the ice lenses. Thoroddsen observed
that voleanic ash layers are bowed upward by the forces
thrusting from below. This mechanism specifies Hégbom’s
frost thrust, clarified by Hamberg (1915) as to the direction
in which it acts, i.e., only perpendicular to the cooling sur-
face. It further uses ideas later specified and supported
experimentally by Taber (1929, etc.) relative to aggregation
of ice lenses on crystalization, a process which will draw
water from thawed soil. Thoroddsen (1913) also suggested
that cryostatic forces are produced as frost penetrates the
bare soil between vegetation hummocks and then into hum-
mocks from the sides, creating closed cells (see Sharp,
1942b).
We observed that even when the ground surrounding the
mud-boil is still frozen four inches below the surface, water
and soil can pour through the bottom of a mud-boil and
fill it with silt overnight. This suggests an artesian force.
Our evidence is not sufficient to analyze artesian- or hydro-
static head-force hypotheses as opposed to differential direct
inter-grain pressures (Terzagi, from Washburn, 1956), nor
to analyze Steche’s (1933) suggestion of a colloid sol-gel
alternation.
Although our observation of the movement of the water-
saturated soil indicates some form of upward thrust within
the column of earth, I cannot see any support for a convec-
tion theory suggested by Nordenskjéld (1909) and later
rejected by him (1911) ; then developed as the Brodelboden
hypothesis (Low, 1925; Gripp, 1929: Gripp & Simon, 1933,
1934a & poe If there were convections, there must be
Movement of soil and water down again — and we found no
indication of this. Furth , as Washb (1950, 1956)
__ Pointed out, the mechanisms do not seem to provide adequate
SOUTHERN BYLOT ISLAND 27
Because the action we observed was vigorous in the early
part of the thaw, when daily frost penetrated an inch or less,
it cannot depend upon horizontal thrusting from (a) cryo-
static forces to disturb soil levels (Hopkins & Sigafoos,
1950; and Washburn, 1956), or (b) cellular containments
within which frost thrust creates involutions (Sharp,
1942b). Both these require the pressure to be applied to
unfrozen spots when most of the active layer is freezing
onto the permafrost. If a long, intense freeze occurs follow-
ing a week of thaw, suitable cryostatic forces may develop,
but these did not develop in the summer of 1954.
When plants start to grow rapidly in the spring, the soil
in mud-boils and stripes is deeply thawed. Movement in the
fluid soil, and the results of freezing and thawing, create
stresses in the growing root tips still partly frozen in. The
process is self-stimulating because once the soil is bare, it is
more subject to rapid and deep freezing and thawing than
are the neighboring vegetation-insulated areas. Many
authors, beginning with Frédin (1918), have recognized
these effects.
Mud-boils and non-sorted polygons. On wet, flat places such
as near sedge marshes, on slopes above the bluffs west of
camp, and at Ooyarashukjooeet, we found mud-boils sur-
rounded by a roll of peat (moss-sedge-willow) and separated
by depressions resembling cracks (Figure 4). Transitional
forms found at Ooyarashukjooeet suggest that when moss
growth from the edge of the boil colonizes the bare, raised
center, these boils merge into moss-covered hummocks which
are regularly spaced so that the depression between hum-
mocks resembles a crack. Some of these are raised mud-
boil-hummocks ; some are completely covered with vegetation
and resemble moss or tussock hummocks (Thufur of
Thoroddsen). On the other hand, mud-boils on upland,
south-facing slopes merge into irregular net patterns on
which lines of prostrate Arctic Willow surround bare earth
meshes (Figure 5). The origin of patterns of hummocks
separated by cracks is discussed below, after the develop-
ment of contraction cracks.
C. Contraction cracks on the youngest alluvial surfaces
Ons thes bere qanias pend ces 66 the & nad 16-fovk:
beaches east of camp, there are cracks forming polygons 1-5
28 WILLIAM H. DRURY, JR.
FicuRe 7. HUMMOCK POLYGONS oN s¢
soil overgrown chiefly with sedges and Arctic Willow are outli sense
ice cores nor ice wedges were found in exe
slope above The Bluffs
Ficure 8.
OUTH SLOPE.
Hummocks of black, heavily organie
med by crack:
avations to the frost table. Site is south
ST-CRACK PENTAGONS UNDER LATE SNOWBANK. Photograph across steep
valley side about five miles west of cam
ank whose extent is shown in the vegetation. Charact
shown: (1) ridge-top barrens on top right; (2) Bell Heath
zone; (3) Aretie Willow where snow persis 4)
tan area near deepest cracks is
SOUTHERN BYLOT ISLAND 29
feet in diameter. Excavations show that these cracks (1)
are perennial, (2) contain coarse sediments and organic
material, including leaves of previous years’ vegetation
(Figures 6 & 9A), and (3) often have a zone next to them
where there is much less organic material than in most of the
deposit (Figure 9A). As Washburn (1956) pointed out,
these cracks grade horizontally into polygonal cracks marked
in the vegetation on the low slopes. They also grade into
hummock patterns.
As the snow first melts away from these surfaces, cracks
appear in the bare soil which at that time is thawed only to
a depth of a couple of inches. Cracks without regular pat-
tern, as much as seven inches deep and several feet long,
appear on open slopes (Figure 12), but in wet, muddy areas
they tend to gather into pentagonal or hexagonal patterns
(Figure 6). Excavations across them show no disturbance
of the soil profile (Figures 9A, C, D and E). In time these
cracks become filled with pebbles or organic material (Fig-
ures 10E, H, I, J, K, and 24 B). Our observations are that:
(1) during the thaw they open every night when the surface
freezes, and close during the day; (2) they may be open or
closed as the snow lies on the ground (Hégbom, 1914) and
they close during the late stages of the thaw; (3) they open
late in the season as the whole surface dries out; (4) their
opening is not the result of thrust of materials that collect
in the cracks, as is shown by these excavations (Figures 6,
9A and D, 10H, I, J and K, and 12). One force opens the
erack when the ground is frozen, closing it at the first part
of the thaw; and a second opens the crack as the soil changes
from a saturated to a damp, tacky mass. Both freezing and
drying draw free water out of the wet soil. Many authors
have proposed hypotheses for formation of cracks without
depending upon cold or dry contraction, but all involve sort-
ing of soil particles. Hégbom (1914) and Sapper (1912)
agree that drying alone is not a sufficient mechanism to
explain the formation of polygonal cracks (Zellenboden,
Spaltenboden) for the following reasons: (a) drying cracks
usually have an irregular pattern; (b) they do not cut so
deeply into the soil as the observed cracks; (c) the tops of
where snow lay longest — Aretie Willow and sedges; (5) Aretie Willow, Avens, Bell
Heather mixed vegetation near the figure. Note the small polygons within the larger
WILLIAM H. DRURY, JR.
PEAT RIDGES, AND MUD-BOIL.
Numbers below
CONTRACTION CRACKS,
diagrams refer to vegetation.
FIGURE 9. EXCAVATIONS INTO
Horizontal lines and numbers above
ms indicate depth to frost.
A. Pentagonal pattern of contraction eracks on bed of temporary pond.
Stippled is gra;
Horizontal ohading & is ‘black organie sandy silt.
hatchi
ed by coarses and that crack zigzags down its depth
in the same area, but
not
Soil:
ing is nes =
is not in its “The erack must reappear
exactly me, year after yea
‘The surface is a rough scum, with binding of algal growth.
ised Beach.
es of most boils are st and uniform, without
nts of organic oll. Roots of tiving plants at the edges of the
SOUTHERN BYLOT ISLAND 31
boil suggest that the soil vat is a There are several stones on the surface
of the boil, and such stones e through the soil. These stones usually have
a crack around them when thee ices dries out.
Soil: Black is organic.
Stippled is gray sand.
Vegetation: (1) Hypnaceae,, pes confusa, Sali: shabu is intergrifolia.
(2) Barren with scatter ey Gases thalli and rieat
(3) Seer Luzula confusa, Salix arctica, and Dry latapiiieita:
eat ridge across depositional slope on 16-foot beac!
cacy) to expectations, peat ridges thawed first on the lowland pier and
the frost table was barely raised under them. Vertical t: f red soil suggests
buried contraction Lee and frost-heaving in soil layers sssated with i it.
oil: Black is organi
Horizontal shading is organic silt.
Hori sean hatehing | is gray-brown sandy silt
White is red ochre silt.
Vegetation: (1) Campylium stellatum, Drepanocladus, and Carex aquatilis.
a Polytrichum piliferum and Tortella.
Aulacomnium palustre.
a
,
3
a} Tomenthypnum switens, Carex na and Saxifraga oppositifolia.
o Salix aeetioe & edicularis sense
(6) Di
D. Two parallel ype on 16-foot beach.
Contraction crack was pen to a depth fat heat 9 inches ‘when Ge mow incited
from this surface 25-28 Jun:
This, like other outa cracks in frozen rong? was filled with dead leaves
which must have gathered while the crack was 0)
Leesan oe Tan Tan repent Tomenthypnum nitens, Tortella, Carex aqua-
(2) " Black ipsa Campylium stellatum, Polytrichum piliferum, and
Carex aquatilis
E. Tall, isolated peat ridge on 16-foot beach.
Vertical red silt trace suggests buried contraction crack. Angular stones on edge
occur under the ridge and embedded in frost table. Note that depths to frost show
less suggest.
(6) Carex aquatilis, Salix arctica, and Sarifraga oppositifolia.
drying cracks are on concave, pee convex, with rounded
edges; (d) the arctic forms appear when the earth is wet,
and the cracks do not close even when covered with water;
and (e) drying cracks occur in many regions, whereas the
forms under consideration are restricted to the Arctic.
European authors, following Hégbom (1914) have in gen-
eral rejected the idea of contraction due to drying (Anders-
32 WILLIAM H. DRURY, JR.
son, 1906; De Geer, 1910; Meinardus, 1912) as a possible
mechanism. Steche (1933) retained this hypothesis and
Washburn (1950, 1956) has resurrected it.
My observations show that when the surface gradually
dried in the late evening, the cracks remained damp longest.
The water does not withdraw deeply because each morning
the dry and frozen crust is only an inch or two deep. Below
it there lies 10-12 inches of wet soil, still above the frozen
ground. If hydrostatic pressure from water in the zone
between two frozen layers led to the cracking upward of
the frozen surface, water or silty material would flow up
through the cracks thus created (viz., Frédin’s 1918 exca-
vations) and the cracks should form at the center of the
stress and radiate outward. On the contrary, we found no
indication of flow of water up through the cracks, and did
find that the cracks form on the margins of the jelly-like
mass, as if resulting from contraction.
The formation of cracks by differential segregation of
fines which soak up moisture, gathering ice which thrusts
upward and, when confined, outward, leading to the forma-
tion of cracks, is not shown in these soils in which there
may be, in fact, segregation of coarse materials in a mud-
boil. I have no evidence of thrust at the sides of the cracks,
although I do have evidence of thrust at the side of the mud
column. I agree with Washburn that there are probably
several mechanisms involved in origin and several in subse-
quent modification.
D. Hummock polygons
On the steep slopes of the 16-foot beach, on areas covered
late by persistent snowbanks, on the banks of small creeks
or rivers, and on open slopes which are covered by shadows
in the late evening (Figure 7), a regular pattern of poly-
gons appears which is a bination of vegetation, hum-
mocks, and cracks. In some places, contraction cracks draw
an angular outline around small solifluction lobes (two feet
across) ; on others the mesh is a luxuriant growth of mosses,
Arctic Willow and often Bell Heather ; in others the cracks
are obscured by the growth of moss. Erosion may produce a
slope covered with hemispherical knobs. I agree with
Thoroddsen (1913), Fradin (1918) and Sharp (1942a) that
these several expressions are related.
SOUTHERN BYLOT ISLAND 33
Figure 10, EXCAVATIONS INTO CONTRACTION AND DEFORMATION CRACKS. Horizontal
ines imbers ‘e diagrams refer to vegetation. Numbers below diagram
eensieg devia to come in inches.
A. Down-slope section showing snow patch, melt-water pool, frost-oval mound and
mesh.
The crack is filled with dead leaves and has an ice wedge i in = bottom. The soil
is a coarse, sandy silt with pebbles and stones, and is soakin;
Véewtetiva: (1) Bare: Clumps of Alectoria jubata, promotes alpinum, Stereo-
caulon paschale, Parmelias, small Hypnaceae, and Luzula confusa.
(2) Peaty edge of crack: Hypnaceae, Luzula confusa, Saxifraga oppositifolia,
Cassiope tetragona.
ee Edge of mesh: Alectoria gees Alectoria pian Stereocaulon seni
ccna vermicularis, four Hypnaceae, arctica, and Saxifra:
ee
nm meshes, and contraction cracks.
(1) Cross-hatehing is Stereocaulon paschale, Hypnaceae, Luzula confuse, Sazi-
y
WILLIAM H. DRURY, JR.
fraga oppositifolia, Papaver radicatum, and Cassiope tetra;
(2) A mound: sandy silt with pebbles and some stones ee no tach long. Clum;
Dis shetty jubata, Stereocaulon alpinum, Stereocaulon eng Parmatine
eae, Poa arctica, Luzula confusa, and Babirrape. oppositifolia.
Oe Bnew
(4) Poa arctica, Luzula confusa, and Salix aretica.
(5) Rock.
Cracks are shown black; note their sinuous form. Soil of mounds is churned and
damp with subsoil sated bias it has been heaved up from below.
Diagramatic section a “B” to emphasize that contraction nae occur on the
parts of
Deformation crack on vegetated, lower southeast-facing slope.
Crack is filled wa fe care silt and peat. The section shows lenses of sand and
peat in the gra: coarse silt and a 4-inch layer of red stain on the right
of the erack. me pelanuns runs parallel to the edge of a raised beach across
@ perpendicular crack. At 8-10 inches below the surface, most of the stones were
cracked into many pieces and fell apart as they were lifted out.
totally organic.
Cee ql) Stevens ulon pala Stereocaulon paschale, Parmelias, Alec-
toria jubata, Luzula confusa, Saxifraga vere doi and Cerastium alpinum.
(2) Stereocaulon paschale, Thamnolia
Cetraria nivalis, Luzula
confusa, Salix okies Sazxifraga gonna tary a i Ores integrifolia in a dense
mat of Cassiope tetragona.
(8) Stereocaulon alpinum, Stereocaulon paschale, ieee Alectoria jubata,
Thamnolia vermicularis, Cladonia pyxidata, Hierachloé Poa glauca,
Luzula confusa, Silene acaulis, Cerastium alpinum, Pushing oppositifolia,
among retica.
m pentagonal crack patterns on raised delta of Golden Plover Creek.
Soil at the edges of the crack shows soil-heaving. Heavy, gray-brown soil is
‘ing is gray-brown sandy silt.
Black is totally organic.
i shading is peaty, crue silt.
olla is gray coarse silt.
ochre, coarse si
Vegetation: scien ace a area to the left of center is fibrous organic material,
pees pa paschale, Hypnaceae, Carex, Luzula confusa and Salix arctica.
(2) Bare, with clumps of blackened Hypnaceae.
is orange to yellow ochre, sticky silt.
Vegetation: qa) Stereocaulon paschale in thick moss cover, including Aulacom-
Lassies) and Tomenthypnum nitens; a complete cover
seatte Salix arctica,
seattered lichens, sedges, growing
in folie
SOUTHERN BYLOT ISLAND 35
Note appearance = successive asap layers and what appears to be windblown
fine sand fi exposed area. A fresh contraction erack appears in the
characteristic Bees aoe to the steeper slope.
Black is organic material eat with fine sand.
Diagonal hatching is reddish-tan, fine sand.
Horizontal hatching is strict ‘brown, silty sand.
Stippling is gray silty
White is red ochre pi aines
Depth to frost at 7-10 inch fees is 22 June; at 18-16 inch level is 30 June.
re
ie mound.
Fs
a
his
BRS
Bs
a
7:
Vegetation. e crack is dense growth of Stereoca: um; in
the crack are Parmelias, dead Hypnaceae, Saxifraga ifolia, some Ri
mitrium lanuginosum an ytrichum piliferum, and cushi of Saxifraga
I. Crack on frost-oval mound, Tui-Tui Tabletop.
Crack, filled with pebbles, is 3 feet long.
Soil: A mixture of coarse and fine sand; a tsiteat of ice mixed with dirt and
organic material follows the line of the crack, the trace of the crack showing
soi
ible.
Vegetation: The nearly bare soil of the mound has scattered clumps of candles
oppositifolia and clumps of Cerastium alpinum and Sagina intermedia grow
the
J. Fresh ee crack on deformation crack on boggy slope.
Fresh crack is on the down-slope side of the major crack. Diagram is 3 feet
across,
Soil is chiefly black-brown peat mixed with silty sand, in which are barely
detectable patches of gray econ A lens of pure ice rises an inch above the
frost table and extends
K. sak a ope doodles Se
pone is reddish-tan coarse and fine angular sand with lenses of coarse sand and
Facet hatching is brown fine sand.
Diagonal hatching is red-brown sandy silt.
ial.
materi
Vegetation: on Stereocaulon
paschale, lanuginosum and Polytrichum piliferum and
cqohicre’ ob Geuivasa vara tne
(2) One foot, bare. <
(3) 8 inches, black crustose Parmelias, Stereocaulon alpinum, Saxifraga opposi-
ifolia.
Pee mo ee
Th stfcelpero aie with a avement of pe 1/1 inch in dame i
by coarse sand with lenses of organic material and gravel overlying
polygons
alpinum, black Parmelias on stones, Luzula confusa, Saliz
ifraga f
Excavations such as this suggest that contraction cracks have worked the entire
churning crack has
(1) The best development of mossy hummocks in our
study area is on the 15°-20° slope between Snow Bunting oa
Creek and Lark Gully (Figure 7). The slope is sheltered
36 WILLIAM H. DRURY, JR.
from the east wind, faces south, and is early free of snow.
In early July, a shadow from the bluffs covers this slope and
the hummocks conform to the outline of the shadow at 1600.
The hummocks are ill-developed where the shadow does not
appear until after 1900. These forms resemble the features
described by authors as Biilten (Tanttu, 1915), Hiigel
(Steche, 1933), and Thufur (Thoroddsen, 1913), but the
hummocks I excavated did not contain an ice kernel and are
too small for the Palse of continental tundra, called, Pingos
by Porsild (1938). They consist of moss growth around an
earth center, and stand 5-15 inches high, averaging 18-24
inches across. Mineral soil rises 1-10 inches in the base of
the hummock and in some cases erupts through the top (cf.
Figure 4). Above the mineral soil core, there is a 3-8 inch
transition level of increasingly organic material, and the top
of the hummock is usually capped by lichen growth except
where mineral soil bursts through.
Because the south slopes are free of snow early in the
spring, they are exposed to a long period during which the
temperature rises above and falls below freezing; further-
more, the daily passing of the shadow over the slopes (a)
increases the intensity and frequency of freezing and thaw-
ing, and (b) preserves moisture. The tops of the hummocks
are dry (as shown by growth of lichens), and this may
indicate both that the soil mechanisms are tended and
that the tops are exposed to the wind. The vegetation
changes in these hummocks, reflecting differences in mois-
ture and shelter, are the same as those on the edges of peat
ridges or frost-cracks described below. Billings and Mooney
(1959) discuss evidence that hummocks and soil polygons
can form out of each other by action of the hummock-form-
Ing Processes, or by drying out and degeneration accom-
panied by plant growth between hummocks.
(2) On other steep south- or west-facing depositional
pes, such as below terrace banks, there occur hummocks
which form polygons 18-24 inches across, separated by
cracks 3-10 inches deep. On the edges of persistent snow-
these hummocks are especially high, and a turf may
grow so that when the snowbank has melted away, its
position is marked by a hollow. It must be, then, that a
supply of water and the time of year when rapid growth
SOUTHERN BYLOT ISLAND 37
can take place are both important in the growth of these
hummocks.
(3) Hummocks appear on the lower parts of unstable and
nearly vegetation-free slopes, where the vegetation is first
able to hold on; and as a better cover is formed, the hum-
mocks grow taller and better int ted. These h :
are of mineral soil below, with a humus layer, and are
colonized chiefly by Arctic Willow. They are regularly
spaced in a pattern such as is shown in Figure 7.
Origin of h ks and hummock-polygons. The two forms
are similar in origin, both dependent upon the formation
of contraction cracks and differential growth of plants.
(1) The mechanism for the origin of mossy hummocks,
first suggested by Thoroddsen (1913), does not depend upon
a tussock of sedge or grass as do those described by Hopkins
and Sigafoos (1950). They do not resemble the hummocks
and ridges probably resulting from a combination of tussock
growth, loess deposition and erosion in the New Zealand
Mountains (Billings and Mark, 1961). I presume, however,
that the moss hummock serves the same insulation function
as the sedge or grass tussock to create discrete units of
insulation and water-holding which tend to be isolated from
the next unit by an area exposed to cold and wet. Tanttu
(1915) suggested that moss hummocks in bogs freeze later
in the autumn and are squeezed as the frost penetrates, and
that they freeze deeper in the winter because they have a
shallower snow cover so that ice persists in them in the
spring.
(2) The origin of hummocks on drier surfaces is associ-
ated with the appearance of perennial contraction cracks, as
the depositional slope builds up. Plants colonize the edges
of small solifiuction lobes, tending to retain moisture and
hold the polygon together, as shown by transitional forms
on the snout of talus slopes. The cracks form every year
and plant growth thickens on the top and sides of the poly-
gon. Then the hummock grows as above. On steep slopes,
drainage may erode the cracks and in some places the
hummocks grow together (Thoroddsen, 1913; Sharp,
1942a). This results in some hummocks being 18 inches
across, with a crack or depression 9 inches across between
them. The pattern still keeps an overall regularity (Figure
ay;
38 WILLIAM H. DRURY, JR.
The size, pattern and structure of these cracks and hum-
mocks grade in type and space into the cracks formed on
muddy, flat surfaces, as discussed above (Washburn, 1956).
E. Micropolygonal patterns — Zellenboden
The surface of many bare meshes is marked with hun-
dreds of inch-long eruptions. These change the surface
aspect and create an effect found frequently on the tops of
New England mountains. In its fullest expression, each
lump is surrounded by a polygon of small cracks, but in
some places (with less slope and less exposure) the crack
polygon surrounds every three to five eruptions, and the
pattern is not as regular (Figure 11 bottom right corner).
In other places, a net of the finest silt, or sand and small
pebbles, surrounds a mesh of dark silt eruptions. These
patterns penetrate barely an inch into the soil. They appear
on south-facing gentle slopes, early free of snow, where there
is a long period of shallow freeze and thaw. In this, the sites
have climate similar to that of the temperate zone moun-
tains. Having read Troll (1944), I looked for situations to
mimic alpine conditions, and finding miniature forms in
such places supports his thesis.
These micropolygons resemble those illustrated in Hog-
bom (1914, p. 320). They appear to be the result of two
forces — one which thrusts fine soil out through a more or
less dried surface, and another the result of tension as the
cheese-like surface layers draw together.
Once the eruptions and tension cracks are formed, the
knobs dry; and in the early stages of stabilization, pioneer
plants —lichen thalli and algae — colonize. Drying draws
the lump together, aided by wetting and frost-thrust from
needle ice. The alga-lichen crust holds the knob together
when water runs in the cracks,
Some slopes, down which the overflow from upland ponds
and sedge marshes drains, are covered with knobs three
inches across, encrusted with tiny mosses and surrounded
by one-inch deep cracks. Several knobs together may be
surrounded by a deeper crack. These merge into soil stripes
of higher, darker soil, alternating with depressions filled
with moss growth and having a sinuous crack down
_ the center. Thus they are transitional forms in which micro-
polygons are superimposed upon a pattern of soil stripes.
SOUTHERN BYLOT ISLAND 39
F. Five- and six-sided polygons
A variety of polygonal patterns is expressed on the vege-
tated, dry uplands. These depend upon patterns of frost-
cracks and are divisible into two types: the first are
generally four-sided (Taimyrpolygone — Steche, 1933;
Troll, 1944) ; the second are usually pentagons and hexagons
(Spaltenboden). Frost-cracks are one of the major expres-
sions of frost action in polar regions and were described
almost as early as sorted polygons. Washburn (1956)
referred to Figurin (1823, p. 275-276) and Von Baer (1838,
p. 403), both of whom reported frost-cracks and ice veins,
as did Von Bunge (1884 and 1887) from Siberia. Their
distribution, as far as I can discover, is correlated with
alluvial deposits and intensely cold continental climates
subject to occasional rapid and extreme temperature oscilla-
tions (cf. Troll, 1944). I will discuss the development of
four-sided patterns in the next section.
We found three sizes of pentagonal or hexagonal polygons
expressed in the vegetation: (1) a pattern 10-20 feet in
diameter, within which was (2) a pattern 3-8 feet in dia-
meter, and within this (3) a pattern 5-12 inches in diameter.
If the major pattern is 25 feet across, the next is 10 feet
across and within this is a pattern 3 feet across; or within
a pattern 10 feet across is one 2 feet across, within which is
one 5-8 inches across.
Are all these patterns the result of similar mechanisms?
How do they develop and how are they modified? How do
the combinations of cracks exist together?
Figure 8, taken across a steep valley, shows a large pat-
tern in which the cracks are emphasized by thawing, pre-
sumably as a result of melt-water flow from a persistent
snowbank whose area is evident in the vegetation. On most
of the uplands these patterns appear only as changes in
relative abundance of bare places, mosses and lichens among
the mat plants. In some upland areas the meshes are com-
pletely barren or occupied by low hummocks; in other areas
the whole surface is covered with vegetation — the meshes
covered with Hypnaceae mosses, lichens, grasses and Dryas,
while the patterns of “cracks” are marked by loose mosses
and taller growths of Arctic Willow and Bell Heather. These
variations are discussed below under “Vegetation.” j
ILLIAM H. DRURY, JR.
FicurE 11. CONTRACTION CRACK IN NON-SORTED STRIPE. The soil is damp in the con-
n
traction crack just below the surface. A dep runs down the center of the
ze
Vegetation is chie efly sedges, Rveas. Locoweed and Bell wane on
ivy cee cover of Hypnum and clump mosses. Photo taken on slope
‘ 16-FooT BEACH. This crack is in
™m, e 9D). The soil
ON
deposit: ional Pio material from
is black, wet, and p
foot bea . Figur
aty and is pulled open, not in any way dis cared by the crack.
SOUTHERN BYLOT ISLAND 41
rire (1952), discussing polygons on the arctic slopes of
e Brooks Range in Alaska, suggested that the smaller
paca oli form within the larger meshes as the surface
becomes older. Most authors agree that the cracks are
formed by the growth of frost wedges or ice wedges, as
Von Bunge (1902), Leffingwell (1915, 1919), and Russian
authors (see Muller, 1947) suggested — with modifications
by Taber (1943). Non-sorted patterns and cracks of smaller
dimension have been largely passed by, being of interest
chiefly to botanists, while geologists have concentrated on
sorted rings and nets.
My studies suggest that cracks on bare alluvial surfaces
and the tops of mud-boils are caused by contraction as
excess water present during the height of the thaw leaves
the soil. They suggest that the large hexagonal and poly-
gonal patterns widespread upon the uplands are the result
of some other contraction force. The latter force may be
intense cold and the formation of ice wedges. My studies
did not examine their formation. Southern Bylot Island
appears to be an ideal area for further excavations in this
study.
IV. THE ORIGIN AND MODIFICATION OF
MAJOR FOUR-SIDED PATTERNS OF FROST-CRACKS
Frost-cracks (or frost-gulls, see below) of the second
major type — four-sided — are 1-4 feet across and extend
an unknown depth into the ground; the meshes are 20-100
yards across and 30-500 yards long (Figures 13, 15, 16, 17
and 20). On level or gently-sloping surfaces, these cracks
run parallel or at right angles to the shore (Figures 13, 15
and 20). Characteristically, a well-developed frost-crack
runs just below the crest of a raised beach and intersects
several cracks running at right angles to the beach. On the
vegetated uplands, the traces of these cracks are in places
so suppressed that they are visible only in details of the
vegetation and in excavations. Cracks are most clearly
expressed on south-facing slopes: (1) on flat, well-vegetated
surfaces, the pattern is usually marked by peat ridges and
rectangular ponds (Figures 21, 22 and 23) ; (2) on exposed
surfaces, the pattern often appears only as a line of lichens
or mat plants on an otherwise bare surface (Figure 18). In
42 WILLIAM H. DRURY, JR.
our camp area, the major crack-pattern extends from the
hilltops across the 6-foot beach to the youngest areas of river
gravels and the tidal beaches.
On the beach east of Aktineq, the broad, gently sloping
surface from the mountains to the shore is marked by cracks
that are essentially parallelograms (Figures 13, 15 and 20).
In contrast, on the gently-domed top of Tui-Tui Tabletop,
the cracks run radially from the center of the top and in
concentric circles around the dome (Figures 16, 17 and 18).
Elton (1927) described similar patterns on small hills in
Spitzbergen. Washburn (1947) illustrated them from Vic-
toria Island. In several places, long and especially deep
cracks mark the edge of the floodplain and the upland, and
run far inland for many hundreds of yards from the mouth
of the river.
Frost-cracks are most vivid — often three feet across —
on exposed beaches, where they may be bordered on each
side by a ridge up to two feet high and four feet across
(Figure 20). Cracks may be double and, although topped
with sand, are usually filled with coarser material, up to
middle-sized stones (Figures 19B and 20).
Péwé (1959) found similar structures in the vegetation-
free areas of Antarctica, and his illustrations and discus-
sions show that the cracks are held open “‘by themselves.” I
asked the local Eskimos whether they had seen any of these
cracks open, or whether they had ever appeared different
from the way they do at present. The answer was “No,” but
Tam not sure we understood each other.
A. Geographic and climatic distribution
Troll (1944) pointed out that (1) frost-cracks occur on
the arctic coast of Alaska, the arctic islands, and in north-
central Siberia; (2) their distribution coincides with pres-
ence of ti ially frozen ground. There seem
to the second. Von Baer (1837/38) described both types.
Troll’s discussion (1944) suggested that the patterns tend
to be quadrilateral in dry climates, and hexagonal or penta-
SOUTHERN BYLOT ISLAND 43
gonal in wet climates where there is more peat. This sug-
gests that there may be two types of forces involved.
B. Origin of major frost-cracks
The following factors suggest that the origin of these
cracks is directly related to solidly-frozen ground: (1) their
formation does not disturb the parallel arrangement of
sediments on the two sides of the cracks except at the very
margin; (2) they occur in sediments (coarse sands) in
which the finest material will not stand in a vertical wall
(Figure 19B and 20) ; (3) they appear in a similarly-sized
pattern in sediments of greatly different constitution — peat,
silt, sand or cobbles.
1. What force is strong enough?
(a) Contraction due to cold. Washburn (1956) discussed
the engineering forces and strains developed by super-cool-
ing of the ground. It is a frequently observed phenomenon
(Thoreau, 1854) that in the depth of winter in northern
countries the ground cracks with an explosive sound. Zhu-
kov (1944) had the most direct and specific description of
this event: following a sharp drop in temperature, cracks
filled with ice appeared to a depth of 2.5 meters, especially
on surfaces not protected by snow. The area of study,
Transbaikal, is like Bylot —little snow and temperature
ranges of 25° C. In his study, he found that the cracks
extended into the foundations of buildings.
Cracks of this origin occur on Bylot (Figures 9D, 10A, B,
H, I, J, 11 and 12), but I doubt that they are solely respon-
sible for the large four-sided pattern. How can contraction
be expressed in a pattern of cracks 50-100 yards apart and
at right angles to each other (Figures 13, 15, 16, 17 and 20)?
This requires that 5,000 square yards of surface act as a unit
and that the tension and compression forces release only at
the edges. Even without experimental evidence of the engi-
neering forces necessary for such a cohesion, it seems to me
unlikely that such a large unit of soil or ice should retain
within itself forces of sufficient magnitude to create the
cracks as expressed. A force resulting from super-cooling
is released in lake ice by a number of small cracks 5-10 (not
100) yards anart — and they are not in a regular rectangu-
lar pattern, but rather scattered irregularly or radiating
from centers. Washburn (1947) discussed the distance over
44 WILLIAM H. DRURY, JR.
hele “
ULAR DEFORMATION CRACKS WITHOUT MOUNDS ON SEA BEACH. The
‘ks running away are parallel to the beach east of Aktinea,
rt. The pavement stones are largely wind-cut and show that
rays blows from the east
shovel gives size. The crac!
and are about 20 yards apa: y ,
the wind almost alw; toward the direction shown in the picture.
TAR DEFORMATION CRACKS IN SEA Ice. Cracks about 10 yards
5 rt on the sea ice along the south shore of Bylot Island east of Aktinea.
‘These cracks form during the thaw — not in the cold of winter.
which the stress must extend (about 30 yards) to release a
erack. The cracks which I discuss below (Figures 11 and
12) in the secondary modification of basic frost-crack pat-
SOUTHERN BYLOT ISLAND 45
terns, seem to me to be of the magnitude and proper surface
expression to be the result of the contraction forces suggest-
ed by Von Bunge and Leffingwell.
Washburn (1956) has outlined arguments against the
simple application of rapid cooling contraction as an expla-
nation for all frost-cracks. Black (1952) described the
complexity and variability of polygonal patterns and their
origins on the coastal plain of Alaska. He doubted any uni-
versal correlation with permafrost. Taber (1943) pointed
out that there is no published evidence that contraction
cracks will form regular polygons. He said: “Ice veins
extend to a depth of 100 feet or more, at least three times as
great as the depth of seasonal change in temperature.”
Taber (1943) and Beskow (1947) suggested that ice wedges
(a) form by the crystallization of ice in ground fissures as
moisture from the soil freezes, and (b) their growth wedges
open the ground. If ice wedge growth forces open the
ground, all cracks should have a mound on both sides. Some
do (Figure 20) ; some don’t (Figure 13). Diicker (1951)
showed that (a) the adjacent beds are bent (see illustration
in Péwé, 1957, and Figure 19B), not sheared as they would
be by lateral pressure, and would result from heaving
upward of areas between cracks; (b) the force due to ice
formation should be perpendicular to the cooling surface.
Dylik (1952) was not satisfied with Leffingwell’s hypothesis
and was convinced that Taber’s ice formation better explains
the involutions and other features related to his process—
congelifluction — in Pleistocene deposits of central Poland.
The excavations east of the Aktineq showed double cracks
with a wedge of soil and narrow ridge between (Figures
19B and 20). For such a double crack to form by contrac-
tion or to cause a new crack to open beside the old one would
halve the forces involved, and yet here the pattern size is
the same — 50 yards between cracks. This crack is larger
than its neighbors.
Various materials collect in the cracks: (a) ice —in
cracks in the peat near Iceberg Lake; (b) organic material
—jin many cracks on the vegetation-covered beach levels
west of the Aktineq (Figures 10E and G) ; (c) fine loam to
coarse sand — in cracks on raised beaches near camp (Fig-
pene 9A, 10H and I); and (d) pebbles and cobbles — in
46 WILLIAM H. DRURY, JR.
cracks east of the Aktnieq (Figures 19B, C, and D). In
order for material to collect without dislocation of the adja-
cent sediments, as in these cases, the cracks must be held
open by forces independent of their margins (note the
slight deformation at the sides of the ice wedge illustrated
in Péwé, 1957).
The presence of cobbles is, I believe, explained by a process
similar to that of pavement formation on the beach surface
(deflation). The sediments of the beach consist of layers
one stone thick, alternating with strata of coarse sand
(Figure 19B). The surface of the exposed beach is covered
with a pavement (Figures 13 and 20). The layers of stones,
some of which are wind-etched, suggest that the surface
pavement has existed all through the period of deposition of
the beach materials. Sand rests temporarily in the crack, is
blown out again, and stones edge deeper while the crack
remains open. Persistence and growth of the cracks, com-
bined with the yearly effects of frost and deflation bringing
stones to the surface, concentrate them and allow them to
collect in the cracks. The sorting of cobbles and sand in the
cracks must be an important clue to their history. The
same sorting in nearly vertical wedges occurs in the upland
frost-cracks of all sizes on all soils. They have been illus-
trated by Péwé (1959).
(b) Deformation of frozen ground. On a marsh-peat
deposit northeast of Tui-Tui Tabletop, recently undermined
at its margin by river erosion, we found large cracks lying
open on 16 June. These cracks were parallel to, and perpen-
dicular to the undercut margin and were filled with ice.
Later in the season (mid-July), the cracks were filled with
water, and ficial ination showed no disturbance
of the peat layers at the edges of the cracks. I suggest that
undermining of the edge of the peat deposits by the river has
led to stresses within the frozen peat which are released by
cracking. The cracks were the same dimensions and resem-
bled those in the adjacent coarse sediments of Tui-Tui
Tabletop, which is unlikely if forces dependent upon the
sediments themselves are involved. The dimensions, then,
must depend upon tensile strength of some factor common
_ to both, which suggests ice in the perennially frozen ground.
On this basis, I have sought another force which would be
ISLAND 47
cH FROM THE A
Aktineq lookt
GURE 15. DEFORMATION CRACK RECTA.
ictas picture taken at about 500 feet pa the
Incipient drainage runs from right to left and d mn crac
e right mid¢ ile of ee piet
is peas into th
ity of the cracks in
arc Sr? a:
Heather a
ice the tin
re 17 was taken in the bottom right corner of t
the | lower left-hand edg
48 WILLIAM H. DRURY, JR.
active in a soil behaving like ice, of sufficient magnitude and
acting in such a way as to create cracks of the dimension and
pattern observed.
As the sea ice, approximately eight feet thick, started to
break up in Eclipse Sound, it first broke clear of the shore
and rode up and down daily with the rise and fall of the
tide. With the passage of weeks and the continuation of the
thaw, leads (cracks) formed and extended in a great sweep,
first from the mouth of each river far out into the ice on
Eclipse Sound; then the ice along the shore broke into
rectangular pans because the cracks in the sea ice formed
parallel to the shore and at right angles to it (Figure 14).
We can presume that these cracks resulted from stresses,
both of compression and tension, applied to the edge of the
ice as a result of the rise and fall of the tide. The ice mass
of Eclipse Sound released the stresses by cracking into rec-
tangles of the same dimensions as the cracks on the neigh-
boring beach (Figures 13 and 20). When the sea ice was
let down on a boulder or bar under the water, it cracked in
a radial and circumferential pattern of the same dimension
and type as on the top of Tui-Tui Tabletop (Figure 16).
A similar pattern can be found on frozen lakes when the
water level drops the ice onto a boulder. My wife and I
Separately decided that these similar cracks result from
deformation acting on similar materials,
5 > ese eer tof
di ts as the ice retreats — isostatic
readjustment — will produce tension and compression
forces in the lidated sediments of Bylot Island’s
south shore (obviously a recently glaciated region), whose
release in the solidly frozen ground should form patterns
resembling those resulting from the forces exerted as the
ice presses against the shore of Eclipse Sound. When sedi-
ments formed by the depositing river are covered with water
or ice, they are protected from the development of peren-
nially frozen ground; then as they are exposed, frozen
ground will develop and thicken, and as the shoreline rises
relative to the sea, stresses must develop between the uncon-
solidated sediments and bedrock, since isostatic readjust-
ment indicates differential movement. These forces will be
applied when the ground first emerges and while the depth
of perennially frozen ground is still relatively shallow (less
SOUTHERN BYLOT ISLAND 49
than fifty feet). This origin suggests similarity to the
“gulls” described by Hollingsworth, Taylor and Kellaway
(1944) and Judson (1947) when consolidated sediments are
deformed. Judson suggested that ground ice wedges may be
involved. Such forms should be called frost-gulls, to separate
them from frost-cracks formed by contraction. The cracks
on Tui-Tui Tabletop (Figures 16, 17 and 18) —a gently-
mounded hill — suggest sediments let down on a bedrock
core. The main bulk of saa in Tui-Tui Tabletop con-
sists of river-borne sedi lished
but we have no evidence of a pe core. Any collapse of
the margins of Tui-Tui Tabletop relative to the center would
create stresses and lead to radial and circumferential cracks.
2. Are the cracks forming now?
If the cracks are formed as perennially frozen ground
thickens, and stresses are applied with the uplift of the
ground, cracks should appear on the youngest surfaces in
their fullest development. If, in contrast, cracks are devel-
oped gradually by annual increments of small cracks, accord-
ing to the theory of contraction resulting from cooling ( =
Bunge, 1887; and Leffingwell, 1919), or of ice crystalizatio:
(Taber, 1943; and Beskow, 1947 >. phe cracks should ast
their best p or ty (Black, 1952) on the
oldest surfaces.
Frost-cracks on Bylot reach their fullest expression on
the lowest beaches; that is, the 6-foot and 16-foot beaches.
In contrast, on the oldest surfaces the frost-crack pattern
is extensively modified and, in many situations, obliterated.
In fact, on the 16-foot beaches, many of these cracks have
“already” been extensively modified. Frost-cracks are form-
ing in the beach sediments below high tide and on the second-
ary channels of the Aktineq River. In other words, fully
expressed cracks (6 inches to 2 feet across) are forming on
surfaces still in the process of deposition, and are being
modified as soon as they are formed.
C. Frost-cracks in other regions
The cracking of perennially frozen ground, resulting from
isostatic readjustment, can be of general application in the
origin of frost-cracks and polygonal ground on unconsolidat-
ed and poorly consolidated materials all across the North.
Cracking is probably unlikely from this source where the
50 WILLIAM H. DRURY, JR.
bottom of the perennial frost is more than a few tens of
feet below the surface, but it seems that this is a likely origin
for cracks in those areas recently exposed and where the
depth of the perennial frost is still shallow, i.e., alluvial and
deltaic deposits, and in sediments overlain by peat — all of
which are especially subject to readjustment. Presumably,
contraction cracks will form within the pattern of cracks
caused by the readjustment of sediments, so that this is not
an “either-or” process. The windswept, snow-free surface
of a beach or coastal plain is well suited for development of
persistent frost, for especially effective results of sharp
temperature drops, and for the formation of deeper and
r cracks of both kinds. The deformation process can
have widespread application and can influence the expression
of cracks which result from rapid temperature drops.
D. Sub Aifieat Seb 3
q' on pt barrens
(1) On the most exposed areas of Tui-Tui Tabletop, the
surface is a gently mounded, tan deposit of fines and angular
pebbles, with the major frost-pattern indicated by thin lines
of tufted lichens (Stereocaulon) and Saxifrage (Figure 18).
(2) In areas where there is conti thin vegetation, dis-
soil marks narrow rims next to the rectangular
pattern of major cracks. (3) On the sheltered sides, dense-
ly vegetated areas show no bare soil, and here the major
cracks appear as depressions or small creeks often widened
by thawing and bordered by the moss-mat plant vegetation
on a high mound. The center of the mesh is low and may be
a marsh or a pond (Figure 17). This gradation in vegeta-
tion and exposure, shown from the air in Figure 16, gives
comparative evidence on the processes concerned in the
secondary modification of original patterns.
Figure 18 shows a shallow hollow on the northeast slope
of Tui-Tui Tabletop, and Figures 19A, C and D show dia-
gramatically the sediment structure. The wall of the trench
shows (1) small fingers of organic soil extending down into
the surface, parallel to each other and at an acute angle to
the surface away from the major frost-crack; (2) indica-
tions of cobble-filled, modified wedges ; and (3) undisturbed
flat-lying strata. These concentrations of cobbles occur in
patterns which are of the same size as the frost-cracks on
the lower beaches.
SOUTHERN BYLOT ISLAND 51
Ficure 17. DEFORMATION CRACKS AND PONDS. The deformation crack has m
n ies.
unds with
Pond mar; eta
¥ ponds on be
volgen mosses and Arctic Willow
= peade! and tir
tion is Eriophorum.
ft.
in the mesh of polygons
i margins of the frost mound are auaie
rsii The vege! m in
okt ng northeast
cross
52 WILLIAM H. DRURY, JR.
During the thaw, while snow lines in the hollows (Figure
18), the rounded, barren surfaces are actively affected by
the frost: (1) small cracks lie open and contain ice at their
base (Figures 10A, B and I); (2) the surrounding soil is
swollen and fines appear to break through the pavement of
angular fragments in many places. No lichens or mosses
grow on these disturbed places (Figure 18). The cracks
become filled with leaves or blowing lichen fragments, and
this material grades into organic stain, extending the trace
of the crack into the soil 12-18 inches (Figure 10A, E, G
and J).
An excavation made during the height of the thaw showed
(a) that while snow persists in the mesh between cracks,
water persists between the snowbank and the ridge; and (b)
that on the ridge long, sinuous cracks occur scattered over
the surface, but usually more or less parallel to the major
crack (Figure 10B). The cracks are open and contain an ice
wedge in the top of the frozen ground (Figure 10A and I).
The excavation (Figure 10A) shows that the frost table
drops gradually from the snowbank on the up-slope side of
such a ridge, but drops steeply on the down-slope side. Frost
activity is limited to ridges which are sodden, usually because
they dam melt-water flow from the snowbank in the mesh.
Excavations suggest (a) that these cracks are closed under
the snow, (b) that they open and close daily in the course
of the thaw while the soil is moist, working the crack deeper
and deeper into the soil; and (c) that at the end of the thaw
when the surface is dry, they are closed during the summer.
These cracks are characteristic of places where the snow
will be blown away during the winter so that they are
exposed to penetration of winter cold. They are probably
produced by cold contraction forces and by ice wedges. In
my experience, however, they do not form regular patterns
Excavations show organic traces (identical to those pro-
duced by these cracks) crisscrossing the surface of frost-
disturbed ridges (Figure 19A, C, D and E). This suggests
that these cracks continue to form year after year on the
edge of the disturbed areas, and disturb the soil progressive-
ly farther and farther away from the major crack region
toward the center, because they form on the edge of the
exposed surface next to the snow patch where soil is damp-
est. As the formation of cracks moves toward the center
SOUTHERN BYLOT ISLAND 53
of the mesh, the sediments in the center are disturbed,
churning up the soil and creating an oval of disturbed soil
surrounding a more or less undisturbed central area (Figure
18). Figures 19A, C and D show excavations across the
mounded border of a frost oval where this process is about
one-third completed. The centers gradually become smaller
and tually are obliterated. When the entire area has
been churned up by minor cracks (Figure 10L), the mound-
ed area of tan, almost barren gravel is oval-shaped with the
long axis pointed down-slope and with dimensions controlled
by the original pattern of frost-cracks (Figure 18).
All transitional stages, from obliteration of the center to
ponds in the mesh (Figure 17) occur on Tui-Tui Tabletop
(Figure 16). Thus, the expression of the frost-crack pattern
on the surface is not measured in time but according to
degree of exposure, since all of Tui-Tui Tabletop can be
considered the same geological age.
Effect of Vegetation. The survey of exposure, vegetation,
and type of frost features on Tui-Tui Tabletop suggests that
vegetation suppresses the opening of this secondary or
annual crack, which in turn controls the modification of the
soil next to the major crack patterns. Many authors agree
on the relation of thin snow and thin vegetation mat to deep
penetration of sudden temperature drops, and Zhukov
(1944) relates it to the formation of cracks. Exposure,
through its control of the vegetation cover, affects frost
action which modifies the major frost-crack patterns.
Effect of Soil Character. Why do frost-ovals form on Tui-
Tui Tabletop and not on the beaches? The cross-section and
exposure of the major cracks are similar, but a marked
difference in the size and nature of the sediments exists.
The finest material east of the Aktineq was a coarse sand
—stream- and wind-rounded — while much of the soil of
Tui-Tui Tabletop was fine sand, perhaps coarse silt — angu-
lar, possibly from continued frost-riving. There are coarse
pebbles and cobbles, many of them cracked by frost, in the
sediments of the raised beach deposits, but few cobbles in
the sediments on Tui-Tui Tabletop. The presence of finer
material which (1) when wet, held together in clumps, (2)
stood easily in a vertical wall in an excavation, and (3) pro-
duced eruptions in the sides of frost-cracks, allows the
frozen soil to behave as described.
54 WILLIAM H. DRURY, JR.
FIGURE 19. EXCAVATIONS INTO DEFORMATION CRACKS AND SOLIFLUCTION Lose. Hori-
diagrams refer to vegetation. Numbers below diagrams
zontal lines and numbers above
depth in in
exeavation is on the sparsely vegetated edge of the exposed top of Tui-Tui
Tabletop (Figures 16 and 18). Exeavation is 12 feet long.
Soil: Black is organic material. Note that these traces dip steeply into the soil
io]
fs
SOUTHERN BYLOT ISLAND 55
‘re contraction cracks are to be expected on the mound.
Dieeais hatched is brown sandy silt.
Diagonally hatched and cross-hatched is a peat layer.
Sploteched tone is coarse sand.
getation: (1) Isolated clumps of Stereocaulon paschale, small Hypnaceae,
Polytrichum piliferum, Luzula confusa, and Salix arctica.
2) Bare sand.
(3) Chunks of peat blown out of op sandy soil, with clumps of Luzula
confusa, Cerastium alpinum, a Sostirere oppositifola.
(4) Organic soil at the surface, protected from the wind and covered bios
naieaonts paschale, Alectoria jubata, Micstseks ochroleuca, and Cassio;
Deformation bei with Mound:
‘cavation crack Aone up the beach east of Aktineq hriegaes 15 and
ae Prevailing wind, coat, is from the right. Finest material is a coarse sand
01
Excavation on Tui-Tui Tabletop (Figures 16 and 18).
Soil: Black shading is peaty, sandy silt.
iagonal hatching is dark brown, sandy silt, grading into sand in many places.
Stippling with sic k is allies and cobbles in a matrix of sand.
Hucavation fi is 20 feet long.
that organic streaks suggest lines of contraction cracks on both sides of
the deformation crack which is marked by the deposit of sand and cobbles.
Vegetation: (1) Complete cover of mats of lichens, Hypnaceae, Salix arctica
and Ce
(2) Patehwork cover of Stereocaulon paschale, Stereocaulon alpinum, Alectoria
jubata, Cetraria nivalis, Cladonia thalli, blackened Hypnaceae, Pol
piliferum, clumps of Poo alpina, Luzula confusa, Saxifraga oppositifolia,
Silene acaulis, and Cassiope tetragona.
(3) Bare surface with patches of Stereocaulon paschale, Stereocaulon alpinum,
and clumps of Saxifraga oppositifolia.
(4) Stereocaulon paschale, black Parmelias, blackened Hypnaceae, clumps of
Luzula confusa and Saxifraga
Frost Oval Mound.
Exeavation on the exposed top of Tui-Tui Tabletop (Figures 16 and 18).
Soil: Black is organic ie material.
Diagonal hatching is damp, sandy silt.
Slipping sed slivien ieee With tacky wei
Uae enses Cine Bees Se eee ‘Under the mound thefr traces surest
heaving and contraction cracks. G
is 25 feet long.
Vegetation: (1) Seattered clumps of Stereocaulon paschale, Salix arctica, Papaver
, Sazxifraga oppositifolia, and Cassiope tetragona.
(2) Barren.
(3) Clumps of Stereocaulon alpinum and Sazifraga oppositifolia.
(4) Mat of Hypnaceae, Salix arctica, and Cassiope tetragona.
Pe Mound, and
Excavation on central Tui-Tui Tabletop (Figures 16 and 17).
is
56 WILLIAM H. DRURY, JR.
ee is 15 feet long from pond margin to crack.
Vegetation: (1) isananasond ei feeeser ‘Sabine
(2) ee ahoren Scheuchz
(3) Tomenthypnum aber A nium palustre, and Carex aquatilis.
(4) Hypnaceae, Bryum, and seh herbacea.
(5) —— paschale, Hypnaceae, Salix arctica, Papaver radicatum, and
Cassiope tetra;
(6) Shateoeeslen | paschale, dead Hypnaceae, Luzula confusa, Papaver radicatum,
and Saxifraga oppositifolia.
(7) Stereocaulon paschale, Hypnaceae, Carex aquatilis, and Cassiope tetragona.
(8) Calliergon, D: hus.
F. Longitudinal Section of Soliftuetion
be.
a0 — slope of Kungo Hill.
Stippling is gray
Exeavation suggests that the es results from deposition over a soil profile.
Soil spss and peat layer is not turned under.
Roots cluster at the border of ie frost-boil and a root of Salix arctica is shown
extending across it.
ics tation: (1) Hypnaceae, Salix reticulata, Vaccinium uliginosum, Cassiope
e Steecawen paschale, Poa arctica, Papaver radicatum, Saxifraga oppositi-
(3) Hypnaceae, Carex Bigelowii, Luzula confusa, Salix arctica, Dryas intergri-
folia,
(4) Hypnaceae, Luzula fusa, Salix aretica.
nd
with edia.
(6) Hypnaceae, Arctagrostis latifolia, Salix arctica, sen ee oppositifolia.
(7) Blackened Hypnaceae, Luzula confusa, Salix a
E. Subseq dification in moss-vegetated areas
On the 16-foot beach there are a number of areas where
single peat ridges, as orderly as if man-made, surround
rectangular marshes or ponds (Figure 22). These patterns
are especially well developed on the 1° to 3° slopes (by
Brunton Compass), on the 55-foot beach, on Tui-Tui Table-
top, on Plover Plateau, and on the lower edge of the deposi-
tional slope, from the 35-foot beach to the 16-foot beach.
(Figure 23), turning nearly at right angles as it meets the
next crack, following an angular course between the peat
ridges as if in a canal. The frost-crack is often deepened and
__“idened by thawing and the places where two cracks meet
SOUTHERN BYLOT ISLAND 57
perpendicularly is often widened into a pond with sloping
banks.
Over most of the uplands the ridges are relatively dry and
are occupied by dry upland vegetation so that a spectrum
of vegetation, from the most barren to the richest, is found
in a transect across one frost-pattern (Figures 17 and 19E).
(See below under vegetation types). In the marshes, the
ridges are marked chiefly by more healthy plant growth of
the same species found in the meshes. In the wettest places,
the vegetation on the ridges is of emergent mosses, marsh
sedges, and Arctic Willow, in contrast to aquatic mosses and
sedges of the centers. During the thaw these areas appear
as a number of rectangular, ridge-bordered ponds (resem-
biing rice paddies) which correspond in size and shape to
permanent rectangular ponds two to four feet deep, such as
(1) Loon Pond, (2) Phalarope Ponds, (3) Upper Phalarope
Ponds, (4) ponds on the southeast corner of Tui-Tui Table-
top, and (5) ponds on the upland surfaces at Oonakuktooyuk
and Ooyarashukjooeet. These forms have been described
from other regions as early as Conway (1897) for Spitz-
bergen, Von Middendorf (1864-1867) for Siberia, and Kjell-
man (1879) for Siberia. In addition to these, there are many
small isolated ridges and these grade into longer and longer
ones, many running across the slope, but some running
down the direction of the prevailing east wind. These peat
ridges are probably the welts (Wiilste) of Hégbom (1914).
Because the ridges accompany cracks on most surfaces and
their pattern is identical to the major crack-patterns, I
presume that they are secondary modifications associated
with the crack.
Will differential frost-heaving and plant growth in vege-
tated areas explain these peat ridges? How can there be two
ridges in some places and one in others? How are the ridges
developed from cracks and how do they influence the develop-
ment of bogs? How are the ponds formed? What is the
origin of isolated, irregular peat ridges? Even superficial
examination shows that these phenomena are confined to
surfaces that are vegetated. What is the effect of vegetation
upon soil formation, and what effects do vegetation cover
and humus have upon the expression of frost-cracks and
frost-heaving?
58 WILLIAM H. DRURY, JR.
1. The effects of vegetation on soil formation. The most
“mature” soils are found on the youngest beach surfaces on
southern Bylot Island (the 6-foot beach), and many of the
oldest upland surfaces where there is a little vegetation (the
55- and 85-foot beaches) have no soil-profile, but traces of
organic soil are found in contraction cracks of previous
years, Excavations show that the more luxuriant the present
vegetation, the richer and deeper the soil-profile. In this
region of the Arctic, soil-profile thus does not measure
passage of time but the amount of vegetation present. Wind
exposure seems to have an overriding influence because it
governs the distribution of vegetation cover.
2. Effect of the presence of organic material on frost activi-
ty. Figures 10, 19 and 24 show sections across major frost-
cracks. In those in Figure 19C and D, the regular sediments
of the meshes are not disturbed at the sides of the crack, and
there are coarse materials at the top of the crack, grading
below into increased concentration of organic materials.
Where the organic material increases, convolutions and
churnings of organic soil are apparent. In other areas (Fig-
ures 24A, B and C) where the cracks are deeply filled with
silt and organic soil, complex convolutions appear. It may
be that we can see the effect of breakage and convolutions
only where the organic material shows it, but certainly such
activity, if present, would appear in excavations on Tui-Tui
Tabletop. The of lutions only in the defor-
mation cracks suggests either that the meshes are firm or
that the cracks are subject to different forces. By digging
across these ridges, and by tracing the patterns into areas of
getation, I found transitions which show that the origin of
the major peat-ridge pattern is the same, and that the differ-
ence between the ovals of the mounded regions of Tui-Tui
Tabletop and the rectangular peat ridges on the bog surfaces
must be explained in terms of modifications subsequent to the
formation of the major frost-crack patterns.
3. O e of isolated contraction cracks in organic soils.
On depositional slopes and in low places, during the early
part of the thaw, straight or sinuous cracks 1 inch across, 2-
6 feet long and 1-7 inches deep appear with no overall pattern
that I could find. They, like others, open with freezing at
_ hight, and close with the increase of moisture in the thaw
SOUTHERN BYLOT ISLAND 59
during the day. These minor cracks, as well as the large
ones, are characteristically filled with fine- or coarse-grained
material in exposed places, and with plant debris or organic
soil in vegetated places. The small-scale cracks shown in
Figure 12 occur without disturbing the soil-profile (Figure
9C). In many places these cracks run next to a sinuous peat
ridge which also runs diagonally down-slope or without
evident relation to soil or other frost patterns.
Where deep, wide, major cracks between peat ridges are
not filled with water during the thaw, narrow cracks may
appear in them, 2 inches across and 1-15 feet long, sinuous
in form and asymmetrically placed in the major crack.
Often these tiny cracks run diagonally across the bottom of
the large crack which has collected a mixed organic and
mineral soil (Figure 10G).
The smaller dimensions of the annual crack and its daily
opening and closing show its relation to the annual cracks I
have suggested to be operative in exposed places. Péwé
(1959) illustrated this type of crack appearing in the major
pattern of cracks in the Antarctic.
The difference in occurrence and expression of the two
sizes of cracks, in this case as well as in the upland case,
indicates that there are two forces involved, which I suggest
are deformation and contraction.
4. How do these two frost-crack types combine to form peat
ridges?
(a) Excavations to the frost table across isolated peat
ridges consistently showed frost-heaving: (i) organic layers
are disrupted upward and (ii) a ridge of frozen soil rises
above the general soil-frost level underneath the peat ridges
and is bordered by slight depressions in the frost table.
Broken stones and fragments of stones occur on edge on the
frozen ridges. The frozen ridge is consistently above the
level of the water table.
(b) The cross-sections of frost-cracks east of the Aktinea,
on Tui-Tui Tabletop, and on the vegetated surfaces, show
shelter where plants, especially mosses, grow luxuriantly,
producing an organic soil which holds moisture. Frost-
heaving occurs and is more evident in the organic
60 WILLIAM H. DRURY, JR.
than in the coarse soil next to it. This frost-heaving raises
ridges on one or both sides of the crack according to local
conditions (see below) but leaves the sediments in the
meshes undisturbed (Figures 10E and 24C).
On the uplands, where dryness and wind limit plant
growth, ridges are barren and secondary cracks form on
them (Figures 18 and 10A). In transitional areas, where
ridges have little vegetation the ridge stops growing because
lack of vegetation means little humus (Figure 17). In
sheltered, level areas where water-soaking inhibits plant
growth, and when a ridge is raised by heaving, clump mos-
ses, marsh sedges, and Arctic Willow grow more luxuriantly
than in the wetter places (Figure 9C and E). Differential
plant growth encourages humus collection and raises the
ridge farther, thus encouraging additional plant growth.
This disturbance is limited to the margins of the cracks
(Figure 21).
Once a ridge is formed, it acts as a dike during the spring
thaw (Figure 23) and holds moisture during the summer.
Growth of aquatic mosses and sedges tends to fill the depres-
sion to water level, and this smooths out the irregularities
inherent in sediment deposits, producing a peat bog, the
mesh of which is flat, crossed by rectangularly-shaped peat
ridges (Figure 22).
5. Depths to frost across the peat ridges. During the thaw,
a series of temperat ts and depths to frost
across the bogs showed that the depth to frost below the
vegetation of a ridge (Figure 9E) — 12 inches on 8 July —
Was greater than that below the vegetation between the
ridges — 8 1/2 to 9 inches on 8 July. The frost level was
deeper under the surface of the plants on the ridge because
of the looser growth, in contrast to the stunted, dense plants
in the centers. The more open growth allows better penetra-
tion of heat. Even so, the frost table was higher under the
peat ridge because the plants on the ridge were much thicker
and the frost table is higher under the ridge. This suggests
that it is not actually the frost level that influences the lux-
uriance of growth, but that the plants on the ridge are raised
during the thaw and can start to grow sooner.
: In rice-paddy-like peat ridge areas, the frost is consistently
_ deeper in the mesh on the up-slope side of the ridge (10
SOUTHERN BYLOT ISLAND 61
20. DEFORMATION CRACK WITH MOUN!
s fe d ttern. Cracks are 6 feet ac:
fia hi
ite of 3
DEFORMATION CRACK
ie fore;
showing low un
creep peat 7
zh
course
and between two parallel ridges r rn
i Photo looking west across Loon Pond on
the conspicuous one. An ice ax gives
aa
62 WILLIAM H. DRURY, JR.
inches extending 2 to 3 feet away from the ridge up-slope)
and consistently shallower (8 inches within a foot of the
ridge) below. This is related to the sun’s heating the stand-
ing water held in by the ridge. The heated water stands on
the up-slope side, then flows through or over the peat ridge,
and is cooled by passage through the ridge. Auer (1920)
pointed out this phenomenon in Strangmoor and discussed
differential plant growth on peat ridges.
6. How does one peat ridge develop from ridges on two sides
of a frost-crack? The crack excavated east of the Aktineq
and a survey of the cracks on Tui-Tui Tabletop show that a
crack may be simple, with a ridge on both sides; or com-
pound, with a larger hillock on one side than on the other; or
a hillock may form in the middle between two small cracks
(Figure 20). Figure 21 shows a crack with low ridges on
both sides which can be traced on the photograph into a
single and then double peat ridge. On low slopes (1°-3°),
water flow will emphasize one ridge at the expense of the
other, since each ridge tends to act as a dam and the whole
tends to form step-like terraces, such as on the slopes from
the 55- and the 35-foot beaches.
7. Rectangular, regularly spaced ponds. Standing water in
the low places hemmed in by the regular pattern of peat
ridges is heated by the sun and this leads it to thaw the
frozen ground below, producing a thaw sink, as described
from western and northern Alaska by Wallace (1948),
Black and Barksdale (1948), and Hopkins (1949).
8. Peat ridges not associated with frost-cracks.
(a) On depositional slopes, minor mud-flows burst out
from under the vegetation and flow over the surface of the
age (Figure 2). A ridge of vegetation usually forms at
(b) On the edges of the delta of the ephemerous streams
off Plover Plateau, peat ridges run cireumferentially around
the snout of the delta, tracing the margins of the garlands
of soil deposited by daily melt-water floods.
(c) The water flows over these deltas as a sheet, within
which riffles (tiny terraces about an inch high) appear in
the same pattern as the lines of peat ridges. The sinuous
form of these rifles, coinciding in size, form and pattern
_ With the peat ridges, suggests that one is developed from the
SOUTHERN BYLOT ISLAND
64 WILLIAM H. DRURY, JR.
other. As the delta deposits thicken, and the surface is no
longer flooded daily, these areas are raised slightly above
their surroundings by the collection of silt or by drying out
of the step, and vegetation of Arctic Willow and clump
mosses grows more luxuriantly, producing peat ridges.
Similar effects control the development of ridges on peat
bogs farther south (the Strangmoor) (Drury, 1956). The
Ficure 24. EXCAVATIONS SHOWING CRYOTURBATION IN DEFORMATION CRACKS, Hori-
zontal lines and m above the dia; ji
" pie grams refer to vegetation. Numbers below the
diagrams indicate depth to frost in inches.
A. Frost-oval mor
jortheast slope of Plover Plat. r i ivi ight
Peng hee ieareg eau toward the Little River (top right
Soil: Horizontal shading is peaty.
Splotehed tone is angular sand and pebbles.
Diagonal hatching is red-brown, partly organic, sandy silt:
mes. 3 i ovals are
ine irregular
~
SOUTHERN BYLOT ISLAND 65
‘ical and convoluted organie traces at the base show the beginnings of
frost heaving. Excavation is 10 feet long.
Vegetation: (1) Bare, eee ae of vhenirsie Seer eat ore ‘myosur-
ic an
ides, Salix arctica, C
(2 mat of Pai eee aa ellicaria, ‘Stren paschale, Stereocaulon
alpinum, pedis a brachyphylla, Kobresia m: Salix arctica mats and
roe itifolia cushions grow on the two ma:
Bare pean with helps stones partly eter with crustose lichens.
crak - rectangular patte
On edge of Tui-Tui eaney on the south edge of the exposed plateau
(Figure 6 at the top of the pict
mecca 22 and 30 apn ides me of 30 June shown.
Bender ee is pea iuoee fine sand.
ochre, coarse and fine sand containing angular pebbles.
Satie Katoh is nacre frigas ee
Stippled with ovals is
Vertical and patti organic oe show that frost heaving reaches nearly
ast edge of the bebop crack is a contraction crack 1/2
On the right south
inch across on 22 Fine which a crack into the still-frozen subsoil.
It was full ix arctica leaves. cay the I 01 ic trace extending into
the sand below this oval on the surface near the center of de-
formation crack is pale gray sand. At 18 inches in the center of i
wel shat ito plat to crack.
excavation is 7 1/2 feet long.
ith Parmelias, Stereocaulon
black
a and racuaa tt of Saxifraga oppositifolia.
(2) Mat of Salix arctica with black Parmelias and Stereocaulon paschale in the
its:
confusa, and Cassiope te
Ovals gravel lenses.
Vertical hatching is wet sand.
The low place on the edge of the crack is filled with a rotting mass of Salix
arctica leaves.
thrust up and to the right from the ridge in the frost
edge of
Convolutions suggest frost
oe en ee ee ee ee of the
formation sediments.
and Pedicularis lanata.
Redlpsaes with individuals of Saliz arctica, Silene acaulis, Papaver radicatum,
‘raga oppositifolia.
a Margin of the crack a mat of Lucile confuse, Salis arctica, and Sazifrage
ifolia.
(4) On the matted willow leaves Luzula confusa, Stellaria
hw Sa, anes nti Poot pari Salix arctica branches
into this from the m
Luzula confusa, Salix arctica,
extend
@) Terk. Chit chase ot Ulercascuise peschel
Polygonum viviparum, Cerastium alpinum, Papaver radicatum, and Sazxifraga
oppositifolia. ?
(6) On the gravel, Stereocaulon paschale, blackened Hypnaceae, Salix arctica,
t 4 Saxif ‘tifolic
66 WILLIAM H. DRURY, JR.
step-like riffles occur wherever water runs over a surface
in a sheet. They are especially well expressed where daily
melt-water floods cross frozen ground.
(d) Ina number of places, minor frost-cracks appear on
depositional slopes. Peat ridges, usually rather short but of
the same height and width as other peat ridges, often accom-
pany them. We found ridges of this sort 15-35 feet long and
parallel to the prevailing wind. Perhaps Arctic Willow
grows faster downwind in its own shelter and these ridges
form by heaving of organic material collecting under the
willows; then the cracks form later. Figures 9D and 12
show a crack, and the soil structure under such isolated
peat ridges.
PART THREE
VEGETATION
INTRODUCTION
The small number of plant species which grow in the area
limits the number of “plant associations,” but the habitats
of the tundra are highly varied and therefore the variations
find expression as different combinations of these few plants.
It is generally recognized, but often forgotten, that each
plant species grows over a spectrum of conditions to which
it is well-enough adapted to exclude others, and to prevent
its own exclusion by competition (Darwin, 1859). Even in
this extreme habitat, many species coincide or complement
each other in their i ts and tol , and thus
occur together. Mason and Langenheim (1959) have recent-
ly proposed the term “aggregates” to get away from the
term “association,” but why add a new word to the mess
that already exists?
: The early students of the relation of vegetation to patterns
in the ground in the North were not concerned with the
nature of the relations between the plants that grow togeth-
er. Frédin (1918) simply described the plants that grew in
certain sites and their relation to solifluction, Polunin
(1934), and Seidenfaden and Sorensen (1937) included the
European concept of characteristic or indicator species, and
numerical abundance of a species in a site sample. Polunin
(1934) was the first after Frédin to classify vegetation on
basis of site and habitat, and discussed why — coming to
SOUTHERN BYLOT ISLAND 67
the conclusion that the surface is kept young by the extremes
of climate and soil disturbance that the vegetation has not
time to develop a humus layer and thus achieve any maturity
which is necessary for the development of an organized
vegetation type. Seidenfaden and Sorensen (1937) devel-
oped the site concept further for the botany of East Green-
land. Raup (1942) has urged the inductive method to
describe what is found and to associate what is found with
the habitat conditions found. But we must choose which
details to stress. Polunin’s (1948) descriptions emphasized
the recognisable, but only slightly different, associations
which proliferate over the North, perhaps because of the
homogeneity of the subsoil by cryoturbation (Raup, 1951)
combined with many site differences created by micro-
topography. My field notes repeat these minutely separable
differences and the main intent of a field study must be to
bring some simplicity out of the evident complexity in
nature. To do this, we may try to answer some questions:
A. Ecological amplitude and species success
Do plants occur together in a limited variety of associa-
tions, or do they occur in all combinations? Of the more than
100 species collected in southern Bylot, only 34 formed the
vegetation of 85 per cent of the land surface; the rest were
scattered plants in specialized parts of bogs or sunny, weedy
sites — the “good collecting” places. The primary species
(those that give the aspect of vegetation to the human
observer) consisted of ten species of higher plants: Hier-
ochloé alpina, Arctagrostis latifolia, Eriophorum angusti-
folium, Carex aquatilis, Luzula confusa, Salix arctica,
Papaver radicatum, Saxifraga oppositifolia, Dryas integri-
folia, Cassiope tetragona; and the following lower plants:
Stereocaulon paschale, Cetraria cucullata, small Hypnaceae,
Aulacomnium palustre, Campylium stellatum, Drepanocla-
dus sp., Tomenthypnum nitens, Polytrichum piliferum.
Plants like Salix arctica, Saxifraga er bic
small, dense Hypnum mosses, and Cetraria
essentially everywhere. These plants are of no use in sans
ing site. Others, such as Dryas integrifolia and Luzula con-
fusa, occur widely in the open areas of the uplands, while
Cassiope tetragona and Salix reticulata, Cetraria islandica
and Ri grow in sheltered spots and
68 WILLIAM H. DRURY, JR.
give indication of site character. Some species occur in
dense stands over many habitats, such as Salix arctica, while
others occur over a wide variation in habitat, but are every-
where rare, such as Saxifraga cernua; still others occur only
as one specimen in one locality (Androsace septentrionalis).
Certain species are restricted, although abundant where they
occur, such as Vaccinium uliginosum in sheltered nooks
below solifiuction lobes ; Salix herbacea in late snow patches ;
Plewropogon Sabinii in the centers of boggy ponds; Merten-
sia maritima and Arenaria peploides on sea beaches. In
some places there is profusion of weedy flowers, such as
the screes below the bluffs: Taraxacum lacerum, Arnica
angustifolia, Erigeron uniflorus, E. positus, Epilobi:
angustifolium, Draba, Papaver radicatum, Oxytropis May-
delliana, and Astragalus alpinus.
Thus certain species occur essentially everywhere,
whether pi or i pi , rare or common; but
other species also rare, common, conspicuous, or inconspicu-
ous may be valuable as site indicators.
B. Regional and microtopographic differences
Do major regional vegetation types exist, or do local
t phic diff. create all the differences in vegeta-
tion that one may find? Are there hilltop vegetations, in
contrast to valley-bottom vegetations?
The vegetation of the heavily frost-cracked area on the
west side of Tui-Tui Tabletop shows, in a transect from the
center of one polygon to the ridge next to the frost-crack
(Figure 19E), the following types: (1) a pond in the center
of the mesh with Plewropogon and Drepanocladus; (2) a
border of T thyp , Auli ium and Carex aqua-
tilis; (3) a thick ground cover of Hypnum mosses and
Cassiope; (4) an area of mixed Dryas integrifolia, Luzula
confusa on the slope of the ridge, and (5) scattered clumps
of Luzula, Stereocaulon paschale, and cushions of Saxifraga
all but the wettest sites, and Cassiope tetragona grows in
scattered clumps on the ridge and in mats on the sheltered
SOUTHERN BYLOT ISLAND 69
(1947, 1951), showed that the physical features of the habi-
tat are of over-riding influence. There are no special vegeta-
tion types that occur on slopes, or on hilltops, or on valley
bottoms.
ES ory maT PLANTS
pees oe VEGETATION
Map 3. VEGETATION OF THE STUDY AREA. Rivers and pond
i be:
are primarily behind raised
isolated barrens are usually raised beach
au tops, or on Louk
pl
cm ridgetops.
C. Primary factors controlling vegetation
All factors act together and two may be directly dependent
upon each other, or both upon another governing force. For
example (as above), the depth and clarity of the soil-profile
are not correlated with the age of the surface, but with the
exposure to the east wind, and moisture factors which con-
trol the growth of the vegetation mat. This in turn controls
growth of organic material in the soil, affecting the subse-
quent frost action.
Raup (1951) and his students, Benninghoff (1952) and
70 WILLIAM H. DRURY, JR.
Sigafoos (1951), at the same time as they point to the uni-
formity of the soil because of the homogenizing effects of
eryoturbation, hasize that the physical conditions of the
environment, especially expressed in frost action, are more
important than the age of the surface. Sigafoos (1952) with
Hopkins, could not use vegetation as any measure of the age
of a deposit, of its materials, or slope, because frost action
has, by disturbance and by mi lief, sup ded all.
These interpretations may offer a dilemma but the observa-
tions are repeatable anywhere in the Arctic. In this area
many factors are limiting, and microtopographic details
change the specific limiting factor which applies and what
plants grow there. Wiggins (1951) said that for the growth
of plants, uniform soil conditions are generally lacking even
within a few square meters (humus, muck and clay, sand
and gravel, all occur within a meter.)
1. Exposure and shelter. (a) In the most exposed places
are scattered tufts (Luzula), cushions (Saxifraga oppositi-
folia), or mat plants (Salix arctica), clumps of lichens such
as Stereocaulon, and large areas of blackened, dead mosses.
Most of the surface is barren. One of the first signs of
increasing shelter is the appearance of Cassiope tetragona,
_ (b) Most of the uplands are covered with a patchwork of
lichen-predomi d mos: domi , over which a
general net of Dryas, Cassiope, and Salix arctica (in which
are scattered grasses, sedges and Luzula confusa) , runs as if
ignoring the ground cover.
(c) As moisture and shelter further increase in sheltered,
and. locally, with Salix reticulata appear Vaccinium uligino-
nigrum, and a complete moss under-
cover. This is the maximum expression of “matured”
vegetation on drained sites,
_ Wiggins (1951) showed that the densest and most con-
vegetation occurs where snow lies deepest and most
SOUTHERN BYLOT ISLAND 71
continuously during the winter, near Point Barrow, Alaska.
Flat surfaces and standing water for part of the season are
necessary for the next step in the thickness of the vegetation
mat.
2. Moisture availability. The first effect of standing mois-
ture is the sudden appearance of a thick growth of sedges,
Eriophorum es nasi etre and Carex aquatilis, and mosses,
and Tomenthypnum. Salix
arctica and Camas coppouttitells are still abundant in
these marshy places, especially on depositional slopes and
behind raised beaches.
3. Sequence of snow-melt. (a) Areas exposed to the pre-
vailing east wind which blows away the snow cover, are
subject to bitter cold in winter and to deep freezing and
thawing during a long spring thaw. The effect is to produce
a barren, frost-churned surface. These areas cover about 10
per cent of the surface we studied.
(b) There are areas which are covered by thin snow in
winter, and melt before the main thaw — about 15 per cent
of the surface studied.
(c) Most of the uplands lie under a few inches of snow,
which is probably seldom more than a foot deep during the
winter, and this cover disappears during a week or ten days
of rapid thaw. We put out several hundred flowerpot stakes
on the broad slopes and the general uplands, marking the
day-by-day change in the outline of snow during the main
thaw — 20-30 June, 1954 — and found no correlation with
vegetation patterns, except that sheltered and moist places
are grown to Cassiope and mosses. These areas cover about
50 per cent of the surface studied.
(d) Areas on the shady side of hills or on the stream
bottoms keep snow cover until two weeks after the general
uplands have thawed. These places are occupied by thick
patches of moss, verdant growth of sedges, and dense
growth of Salix arctica. They show a characteristic lack of
Dryas and Cassiope. Mosses form cushions or hummocks
surrounded by a crack and topped by lichens. These
sheltered, shady areas cover about 20 per cent of the surface
studied.
(e) In some places snow lies in persistent drifts in valley
bottoms and northwest slopes until the third week of July,
72 WILLIAM H. DRURY, JR.
and in these places the general ground cover is almost entire-
ly Salix herbacea, a few tiny Equisetum arvense, Juncus and
Ranunculus, and weedy Bryum mosses, This specialized hab-
itat exists uniformly in site and vegetation across Siberia,
North America, and in the mountains of Norway and
Sweden. Gjaerevoll (1956) has discussed it in detail. These
cover about 5 per cent of the surface studied.
4. Effects of the full rays of the sun. There was no vegeta-
tion shelter from the sun’s rays except for mosses, but cer-
tain slopes were exposed to full sunshine for the maximum
period of time during the heat of the day, while others slop-
ing to the east, north, or northwest were in shadow most of
the “heat” of the day. We found no general differences in
vegetation between that of the gentle upland slopes and that
on south-facing gentle slopes. On gentle northeast, north,
and northwest slopes, there is an increase of Cassiope. On
disturbed soils of south-facing frostings, scree, or alluvium
in the ravines, weedy masses of flowers grow.
D. Time of flowering
When we arrived, the temperature was below freezing
most of the time, and regularly went below freezing every
Draba nivalis, D. glabrata, Potentilla Vahliana, Cerastium
alpinum, Salix arctica, Erysimum Pallasii, and Oxyria digy-
na@ were all in flower.
After the first of July, when there was no regular occur-
rence of frost, the full flood of flowering appeared ; and on 2
pes, Papaver radicatum, Draba alpina, D. nivalis,
Braya purpurascens, Lesquerella arctica, Saxifraga nivalis,
S. cernua, Dryas integrifolia, Astragalus alpinus, Oxytropis
E. Variation in the richness of site
: Our study area shows the familiar phenomenon of some
sites being rich in species and others poor. Disturbed soil
SOUTHERN BYLOT ISLAND 73
and fine-grained alluvium of steep, sandy slopes in the
ravines west of camp — out of the wind and south-facing —
were covered with a weedy growth of many species of limit-
ed distribution. Yet some of the species in these places (such
as the Drabas, Epilobium, Astragalus, Saxifraga oppositi-
folia, Salix arctica, and Oxytropis) also grew on the uplands
or on river gravels. In contrast, on “poor” sites such as
the persistent snow patches in valley bottoms, only one or
perhaps three species occurred together. There are all
gradations imaginable between these two extremes, while
the average site — 75 per cent of the land surface — sup-
ports the fifteen common species growing everywhere, mixed
into which occur a variable number of less conspicuous, less
common species,
Seidenfaden (1931) states one unanswered problem : “The
difference in types of vegetation, therefore, can hardly be
said to consist in a more abundant representation of species
in the gneiss localities, for nearly all the species represented
there may, with more or less success, be met within the areas
of sediments. When, therefore, the difference is so pro-
nounced as is practically the case, this is principally due to
e | i —a ph which cannot easily be
expressed numerically.” He mentions, further, plants which
grow as a few scattered plants nearly everywhere without
anywhere becoming abundant.
I agree with Gjaerevoll ( 1956) in his discussion of the
ani getation “value” of Salix herbacea. He
makes the point that acidophilous species of plants have a
general adaptation and lack of specificity, while calciphiles
are much more specific as indicators. The relative inhospi-
tality of limestone prevents rich, closed societies from devel-
oping, and by the openness of the society, the poverty of
limestone allows the plants which would otherwise be less
able to compete, to hold on in small numbers (Polunin,
1934). On this basis only, acidic barrens should also be rich
in species. I presume that isolation, which is to be expected
by the geological nature of the limestone deposits, has
encouraged formation of species by geographic isolation.
The flora of southern Bylot is rich in calciphiles.
F, Vegetation types and habitat selection by birds
To test the validity of the vegetation types, I compared
74 WILLIAM H. DRURY, JR.
them with habitats selected by birds. Many experiments
show that birds respond to the same sort of environmental
stimuli as men. If they select the same vegetation types as
we do, we can assume it is for a different purpose, and that
our assay indicates something real, or expresses factors of
general influence in the environment.
I found a large degree of coincidence ; for example, Baird’s
Sandpiper (Calidris bairdii), Snow Goose (Anser coerules-
cens atlantica), Black-bellied Plover (Pluvialis squatarola),
and Horned Lark (Eremophila alpestris hoytii) nested on
the exposed areas. I am convinced, however, that Baird's
Sandpipers and Horned Larks selected the only surface areas
free of snow when they “had” to nest — not necessarily
because of the vegetation. Black-bellied Plovers and Snow
Geese arrived later and, being wary, selected areas with a
wide prospect — terrace margins or hilltops — and perhaps
the barrenness is coincidental in the case of the Snow Goose;
but in the case of the Black-bellied Plover, the nests are
specifically placed in areas of nearly barren soil. These four
species are characteristic of the High Arctic dry tundra.
Other species of birds — White-rumped Sandpiper (Heter-
opygia fuscicollis), Oldsquaw (Clangula hyemalis), and
Golden Plover (Pluvialis dominica) — selected areas of more
or less complete vegetation cover. Their geographical dis-
tribution is more southern, coinciding with the wet tundra
vegetation. Thus there is a general phi incid
in the distribution of vegetation and habitat selection in
these species.
Lapland Longspurs (Calcarius lapponicus) nest in a recess
under a hummock, usually a sedge tussock, over most of its
range. On southern Bylot Island where there was little
development of tussock growth, it nested on the edges of
frost-cracks, under overhanging vegetation of Salix arctica
and Cassiope, or in the edges of peat ridges (Drury, 1961).
Each of these situations, most conspicuously Cassiope, sup-
_ plies a recess with an overhang, and, I presume, the recess,
not the plants, is selected.
Raunkiaer’s (1934) systcm of form categories has been
used by Bécher (1933a and b) in contributions to under-
standing the ecology of arctic plants in Greenland. Orni-
thologists studying habitat and nesting-site selection have
SOUTHERN BYLOT ISLAND 75
come to the conclusion that there are many features to be
considered: (1) form; (2) aggregation of the individual
plants to create topography (relation of clumps to open-
ings) ; (3) internal structure of the individual plants; and
(4) details of the ground cover. For example, Dwarf Wil-
lows and Dwarf Birches and Alders fill the function by
being bushes, but, having different internal branching, they
are separated as nesting sites.
Further pursuit of this topic gets us into Gestalt or total
impression psychology and the Umwelt or selective reactivity
of non-human organisms. It is a treacherous morass.
conclude that birds respond to landscape and to the form
of the vegetation cover, and that associations are important
only as they supply required forms.
G. Indicat Domi Ss ion and Frost Action
There are fundamentally different bases for the systems
of classifying vegetation as developed in the United States
and in Europe. One difference is the idea of succession and
development described by Cowles (1899, 1901) and Clements
(1916), and expanded by Tansley (1935, 1939) to include
the concept of eco-systems. On the other hand, the initially
more inductive Continental systems are divided into two
parts: The Braun-Blanquet Montpelier school considers the
concept of indicator species; and the Scandinavian system
includes the idea of dominant species. Separate from these
diff is the bift tion between Clements’ super-
organismic concept and Gleason’s (1917, 1926) individual-
istic concept of plant association.
A further dichotomy is between those who seek similar
and uniform details of vegetation and follow the details over
many sites where the similar vegetation appears; and those
who seek similar sites and deal in summarizing and explain-
ing variation in vegetation found on the sites. The concept
of the indicator, or characteristic, species tends toward site
characteristics. On the other hand, pure plant sociology
discovers layers or units which repeat themselves regularly,
while other species may come and go, independent of that
layer. Bog students (Sjérs, 1948) have been working in
detail on the physiology of “site” requirements. Goodlett
(1960) discussed in detail the development of the concept of
site and the tremendous importance that soil, microclimate,
76 WILLIAM H. DRURY, JR.
slope, and so on, have in fi ting stand develop: and
productivity. This is the conviction of students of habitat,
ie., given similar site and flora, similar associations or
communities appear. Stout (1952) illustrated studies of site
in the deciduous forests by showing how deep soil structure
directly affects the composition of a stand of trees.
One difficulty is extricating ourselves from the all-pervad-
ing Clementsian dogma; for example, Whittaker (1960), in
his able discussion of Serpentine Vegetation, feels he must
justify a permanent “successional” stage. If the vegetation
retains characteristic elements, of what importance is it
whether it fits into some overall climax scheme? Each spe-
cies of plant is out for itself and where two or more grow
together, it is the result of coincid of requi t
The differences in “systems of classification” lie not in the
conclusions drawn from observable facts, but in the develop-
ment of these ideas following deductive lines of logic beyond
the facts.
Understanding will come from detailed studies of the
biology of individual species and of geographical forces
producing ground structures and soils. The plant association
is useful to describe landscape and, to an esoteric few who
are familiar with the plant species, habitat, but it is of little
ecological importance per se to the plants concerned.
I use the association concept developed by Raup (1951),
but include the subjective concept of site. This association
may be equivalent to the alliance of the Zurich-Montpelier
school. But within it, many detailed variants may appear,
some of striking uniformity. These variants would conform
to the Scandinavians’ sociations. My descriptive categories,
however (primary vs. secondary) are equivalent to the
Hult-Sernander scale in that primary is equivalent to the
value 5 in the case of mat plants, and to 4 in the case of
plants growing in clumps. The other concept, secondary,
includes 1, 2 and 3; that is, a cover equivalent from 1/4 to
less than 1/16 of the total area. Primary species used here
include a combination of frequency and degree of cover,
, regard for dominance. The judgment which is
involved in this weighs frequency more than degree of cover-
i Thus, a grass, sedge, or clump
itifolia) is called primary, while
SOUTHERN BYLOT ISLAND Te
according to the Braun-Blanquet system’s degree of cover-
ing, its habitat form gives it 2 or less. Botanists are familiar
with the observation that in a certain stand, using any classi-
fication, certain plants are much more conspicuous than the
second-level plants. These are of frequency over 80 in
Raunkiaer’s (1934) system, or over 3 in the Braun-Blanquet
system. The next level of frequency in Raunkiaer might well
be 50 or 60, and Braun-Blanquet, less than 2. There are
similarities of association between my studies and those in
East G ] i and Sorensen, 1937), in the
Canadian Arctic (Polunin, 1984, 1948) and in Scandinavia
(Nordhagen, 1943). This shows that botanists choose simi-
lar patterns.
1. Exposed Sites: Map 3, Barren
Primary: Blackened and dead clumps of mosses (Polytri-
chum and Hypnaceae — including Dicranum), Stereocaulon
paschale, Luzula confusa, Salix arctica, Papaver radicatum
and Saxifraga oppositifolia.
Secondary: Cetraria nivalis, Stereocaulon alpinum, Poa
arctica, Cerastium alpinum.
2. Uplands: (a) Exposed: MAP 3, Barren
Primary: Stereocaulon paschale, Cetraria migra Par-
melia saxatalis, Alectoria jubata, A.
piliferum and some Hypnaceae.
Secondary: Cetraria islandica, C. nivalis, Thamnolia ver-
micularis, Cladonia rangiferina, C. sylvatica, Stereocaulon
alpinum, Hypnaceae (including Rhacomitrium lanugino-
ee, Hierochloé alpina, Poa arctica, P. glauca, Festuca
latifolia, Kobresia myosuroides,
Luzula ¢ confuse, Cerastium alpinum, Silene acaulis, Papaver
radicatum, Draba nivalis, Saxifraga oppositifolia, Dryas
integrifolia, Cassiope tetragona.
(b) Sheltered: Map 3, Dry Mat Plants
Primary: Stereocaulon paschale, Dicranum ‘Sp., Polytri-
chum piliferum, Tortella sp., Did Sp.,
lanuginosum, Aulacomnium acuminatum, Poa glauca, Luzu-
la confusa, Salix arctica, Dryas inteerifolin, Cassiope tetra-
gona.
Secondary: Alectoria ochroleuca, Cetraria cucullata, Equi-
setum arvense, Arctagrostis latifolia, Alopecurus alpinus,
78 WILLIAM H. DRURY, JR.
Hierochloé alpina, Salix arctica, Oxyria digyna, Polygonum
viviparum, Salix herbacea, S. reticulata, Stellaria longipes,
Silene acaulis, Papaver radicatum, Eutrema Edwardsii,
Draba alpina, D. glabella, Saxifraga oppositifolia, Pedicu-
laris lanata.
These two — (a) and (b) —alternate, merge and com-
bine in Map 3, Dry Mat Plants. In places, streams of one
run through a general cover of the other, or meshes of (a)
are surrounded by lines of (b) in non-sorted polygons.
Gentle north and east slopes are covered with (b) but with
Cassiope tetragona forming almost a complete cover on their
upper slopes and almost completely lacking on the lower
slopes.
On many upland surfaces, polygons of several sizes are
expressed in the vegetation. The pattern expressed in the
meshes five feet across is typically: (a) the barren areas
have scattered, blackened mosses, Stereocaulon paschale,
Luzula confusa, Salix arctica, and Saxifraga oppositifolia;
(b) the margins of the meshes have low Hypnaceae mosses,
Luzula confusa, or Carex Bigelowii, Salix arctica and Dryas
integrifolia; (c) growing out onto these margins from the
cracks which form the polygons are Aulacomnium palustre
and Tomenthypnum nitens. These mosses fill the cracks and
in them grow Arctagrostis latifolia, Alopecurus alpinus,
Eriophorum angustifolium, and Cassiope tetragona.
On more exposed sites the cracks have only Hypnum
mosses; the marsh mosses and the marsh grass and sedge
do not appear. The patterns are built of relatively exposed
Sites alternating with relatively sheltered sites, rather than
of any specific and regular vegetation differences. On all
sites, the impression is that the plants from the cracks are
constantly advancing into the centers of the mesh and is
prevented from col by disturb of the soil. As
soon as the disturbing f diminish, the vegetation of both
centers and cracks becomes “more mesophytic.”
3. Damp Slopes: map 3, Wet, Mossy
i : Tomenthypnum nitens, Dicranum sp., Didymo-
don sp. and other Hypnaceae, Carex aquatilis, Luzula con-
fusa, Salix arctica,
Ss , Cetraria islandica, C. cucullata, Cladonia
tagr latifolia, Carex misandra, Salix
SOUTHERN BYLOT ISLAND 79
reticulata, S. herbacea, Oxyria digyna, Cerastium alpinum,
Ranunculus nivalis, Eutrema Edwardsii, Saxifraga oppositi-
folia, S. cernua, S. nivalis, Potentilla hyparctica, Dryas
integrifolia, Cassiope tetragona, Pedicularis lanata, P.
hirsuta.
4. Late Snow Areas:
Primary: Bryum sp., Tortella sp., Salix herbacea.
Secondary: Cetraria islandica, Stereocaulon paschale,
Alopecurus alpinus, Carex aquatilis, Luzula confusa, Juncus
biglumis, Salix arctica, Oxyria digyna, Polygonum vivipar-
um, Ranunculus nivalis, Saxifraga oppositifolia, S. cernua.
5. Sunny South Slopes: MAP 3, Dry Mat Plants
(a) Fine-grained slump:
Primary: Stereocaulon paschale, Cetraria cucullata, Poa
glauca, Trisetum spicatum, Hierochloé alpina, Kobresia
myosuroides, Luzula confusa, Salix arctica, S. reticulata,
Dryas integrifolia.
Secondary: Thamnolia vermicularis, Alectoria ochroleuca,
A. jubata, Equisetum arvense, Poa arctica, Festuca brachy-
phylla, Alopecurus alpinus, Oxyria digyna, Polygonum
viviparum, Cerastium alpinum, Silene acaulis, Papaver radi-
catum, Draba nivalis, | Samir raee: oppositifolia, S. nivalis,
Potentilla hyp lus alpinus, Oxytropis May-
delliana, Vaccinium sifenutan Antennaria Ekmaniana.
(b) Sandy alluvial fan:
Primary: Poa glauca, Hierochloé alpina, Salix arctica, S.
reticulata, Oxytropis Maydelliana, Arnica alpina, Taraxa-
cum lacceurt Conspicuous flowers: Cerastium alpinum,
Papaver radicatum, Erysimum Pallasii, Astragalus alpinus,
Epilobium latifolium.
Secondary: Poa arctica, Festuca brachyphylla, Trisetum
spicatum, Elymus arenarius, Kobresia myosuroides, Carex
maritima, Oxyria digyna, Polygonum viviparum, Lesquerel-
la arctica, Draba subcapitata, D. lactea, D. nivalis, D. glabel-
la, D cinerea, Arabis arenicola, Braya purpurascens,
Saxifraga oppositifolia, S. nivalis, S. hieracifolia, Potentilla
hyparctica, P. rubricaulis, P. Vahliana, Pedicularis hirsuta,
Erigeron uniflorus, E. compositus.
(c) Stable raised beaches or ridges, lemming fertilized:
Primary: Poa arctica, Poa glauca, Trisetum spicatum,
Hierochloé alpina, Luzula confusa.
80 WILLIAM H. DRURY, JR.
Secondary: Equisetum arvense, Festuca brachyphylla,
Stellaria longipes, Papaver radicatum, Saxifraga oppositi-
folia, S. cernua, S. nivalis, S. hieraciifolia, Potentilla hypare-
tica.
6. Marshes: MAP 3, Wet, Mossy.
(a) Raised ridges and margins:
Primary: Hypnaceae, Aulacomnium palustre, Tomenthyp-
num nitens, Carex aquatilis, Salix arctica.
Secondary: Poa arctica, Arctagrostis latifolia, Eriophor-
um angustifolium, Salix reticulata, Stellaria longipes, Silene
acaulis, Ranunculus nivalis, Saxifraga oppositifolia.
(b) Wet centers:
Primary: Drepanocladus sp., Tomenthypnum nitens,
Campylium stellatum, Arctagrostis latifolia, Hierochloé
pauciflora, Eriophorum angustifolium, E. Scheuchzeri,
Carex aquatilis.
Secondary: Stereocaulon paschale, Salix arctica, S. reticu-
lata, S. herbacea, Cerastium alpinum, Silene acaulis, Poten-
tilla hyparctica, Pedicularis lanata, P. Langsdorfii.
(c) Ponds:
Primary: Drepanoclados sp., Pleuropogon Sabinei.
alpinum, Papaver radicatum, Saxifraga oppositifolia, Astra-
galus alpinus, Oxytropis Maydelliana, Epilobium latifolium.
8. Sea Beaches:
Phippsia algida, Puccinellia Langeana, Deschampsia
Elymus arenarius, Sagina intermedia, Arenaria pep-
loides, A. humifusa, Stellaria humifusa, Cochlearia offici-
nalis, Arabis arenicola, Mertensia maritima.
To illustrate how these site types are built together into
the vegetation which covers a certain slope, I include two
samples:
A. T>.
solifluction lobe on a south-facing slope: (Cf. Fig-
ure 19F)
(a) Top:
Primary: Blackened = Arctagrostis latifolia, Luzula con-
fusa, Salix arcti
Secondary: Stereocaulon paschale, Cetraria cucullata, Alectoria
ochroleuca, Poa Carex Bigelowii, Polygonum vivi-
alpinum, Papaver radicatum, Saxifraga
oppositifolia, Dryas integrifolia, Cassiope tetragona.
SOUTHERN BYLOT ISLAND 81
(b) Mud-boil—the center of the top:
Carex misandra, Luzula confusa, Polygonum viviparum, Sagina
intermedia, Saxifraga oppositifolia.
Mud-boil — the edges:
Arctagrostis latifolia, Alopecurus alpinus, Luzula confusa, Salix
arctica, Saxifraga oppositifolia, S. nivalis.
(c) Crescent-shaped, gentle slope forming nite of the surface of
the lobe:
Primary: Carex Bigelowii, Luzula confusa. Dryas integrifolia.
Secondary: Dead mosses, Stereocaulon paschale, Poa arctica,
Hierochloé Pies Oxyria digyna, Eolyponum Uae Silene
ees vate Oxytropis Vac-
ligi
(d) ‘Sheltered Tullow} telow the lobe, and on the riser of the lobe.
Primary: Hypnaceae mosses, Cassiope tetragona.
Secondary: Stereocaulon paces Luzula confusa, Salix arc-
tica, S. ances Papaver radicatum, Dryas integrifolia, Vac-
cinium uliginosum.
B. Transect across frost-crack polygon: (Cf. Figure 19C and D)
(a) Exposed area:
ee ger clumps of Stereocaulon paschale, Cassiope tetra-
2. Ridges: bare or clumps of Stereocaulon paschale.
3. Centers: Stone pavement, Stereocaulon paschale, Salix
arctica, Papaver radicatum, Saxifraga oppositifolia,
Cassi: na.
(b) Out of the direct wind: (Cf. Figure 19E)
1. Crack: water, Calliergon sp., Drepanocladus sp.;
Bank: Aulacomnium palustre, Salix herbacea;
Sheltered slope of ridge, in crack: Primary: Hypnaceae,
cularis, arctica, Stellaria longipes, Saxifraga
oppositifolia, Pedicularis lanata.
to]
Primary: Blackened mosses, Lichen thalli, Stereocaulon
paschale, Luzula confusa, Saxifraga oppositifolia, Cas-
siope tetragona.
Secondary: Cetraria cucullata, Alectoria jubata, Tham-
nolia vermicularis, Polytrichum piliferum, Poa aretica,
Salix arctica, Papaver radicatum.
3. Sheltered sides of the polygon center:
Primary: Stereocaulon paschale, Cassiope tetragona.
Secondary: Cetraria cucullata, Cladonia _Sylvatica, Alec-
toria jubata, Thamnolia
piliferum, Poa arctica, Hierochloé alpina, Reriigtoctio
latifolia, —- confusa, Salix arctica.
4. Wet margins :
Primary: Tomenthypnum nitens and Aulacomnium Sate :
82 WILLIAM H. DRURY, JR.
tre or Polytrichum piliferum, Carex aquatilis or Salix
herbacea.
Secondary: Cetraria cucullata, Stereocaulon paschale,
Hierochloé pauciflora, Arct. tie latifolia, Eriopl
angustifolium, Luzula confusa, Salix arctica.
5. Pools:
Moss: D.: 1; sp., Pl pogon Sabinei.
Mud: Eriophorum Scheuchzeri.
The Altitudinal Limit of Vegetation on Mt. Thule:
Ferris and Ames reported that Cetrarias, Alectoria
ochroleuca, Stereocaulon paschale, Thamnolia vermicularis,
Hierochloé alpina and other grasses, Luzula confusa, Salix
arctica and Draba cinerea grew in sheltered places up to
2,600 feet. At the highest level of vascular plants (3,000
feet) grew Cerastium alpinum, Papaver radicatum, and
Saxifraga oppositifolia. Black Umbellicaria, crustose Par-
melias and Alectoria jubata grew to 4,00 feet. Crustose
Parmelias and unidentified mosses ( collected) grew at 4,800
feet.
PART FOUR
PLANT LIST
In identifying m: Iections, I have used the Gray
Herbarium and Arnold Arboret llections, and chiefly
the works of Polunin (1940, 1959), and Porsild (1957), as
well as recent phs. My lusi follow closely
those of Polunin even in preference to the work of recent
ialists. My ti trate on species about
which I differ or which I consider of especial interest. The
short list of non-vascular plants includes those which I felt
IT could recognize in the field. I have not compared these col-
lections with herbarium specimens.
All collections were made in 1954; all localities are at the
_ mouth of the Aktineq River on southern Bylot Island, North-
West Territories, Canada, unless otherwise noted.
| NON-VASCULAR PLANTS
Common Name
Alpine Reindeer Lichen
Reindeer Lichen
Scarlet Wand Lichen
Goblet Lichen
Gray Lichen
SOUTHERN BYLOT ISLAND 83
Lichens ‘ommon Name
Stereocaulon denudatum FIk. Smooth Gray Lichen
Parmelia sp. Shield Lichen
Cetraria islandica (L.) eae Iceland Moss Lichen
Cetraria nivalis (L.) A Snow Lichen
Cetraria cucullata (Bell. ; Ach. White Lichen
Alectoria nigricans (Ach.) Nyl. Black Mane Lichen
Alectoria ochroleuca (Ehrh.) Nyl. Roan Mane Lichen
Caloplaca sp. Orange Star Lichen
Thamnolia vermicularis (Sw.) Ach. Worm Lichen
Mosses
Sphagnum sp. Sphagnum
Mnium sp. Leafy Moss
Dicranum sp. Broom Moss
Tortella sp. Twisted Moss
Didymodon sp. spo
Rh itrium | i (Hedw.) Brid. Gray Moss
Bryum sp.
Aulacomnium palustre (Hedw.) Schwaegr. Marsh Moss
Campylium sp. Star Moss
Drepanocladus sp. Curly Moss
Calliergon sp. Golden Moss
Serna nitens (Schreb.) Loeske Shining Moss
Polytrich Haireap Moss
Poipieleioan ae Hedw. Awned Haircap Moss
VASCULAR PLANTS
Conspicuous species are in boldface type. Species which are numer-
cus but do not attain as great “importance” in the vegetation are
marked with an asterisk.
Equisetum arvense L. (Horsetail) 54204 — 27 July — Muddy shore of
the Little River.
Widespread but scattered; on sparsely vegetated, sheltered
places, Len late snow patches.
Equisetum variegatum Schleich 54114—14 July — Mossy sedge-bog
on the floodplain at Ooyarashukjooeet.
Hierochloé alpina (Sw.) R.&S. (Sweetgrass) 54201 — 27 July — Dry
frost-heaved, raised beach on the south slope of Kungo Hill.
Widespread, and especially common on dry, sunny slopes.
Hierochloé pauciflora R.Br. (Marsh Sweetgrass) 54178 — 26 July —
Mossy, wet marsh 3
Bie narrow senha with the spreading awned heads
H. alpina. This was common in “rice-paddy” areas.
pa i aigeeis Sm. (Foxtail) 5: 54134—13 oT at
Ooyarashukjooeet; 54156—20 July — Blackened moss
of non-active mud-boils on the 35-foot beach; 54218 — 26 duly
— Lower part of sandy scree below the bluffs one-half mi
west of camp.
et eS
on south slopes. It grows as scattered individuals. ie
84 WILLIAM H. DRURY, JR.
Arctagrostis latifolia (R.Br.)Griseb. (Redtop) 54174 24 July —In
sedgy, wet seep at the head of Golden Plover Creek.
Most upland habitats, but is commonest in closed vegeta-
tion on undisturbed soils, usually with a und cover of
mosses. It occurs in wet seeps and on solifluction lobes, but
is uncommon in dry, exposed sites and in marshes covered
with water.
Deschampsia pumila (Ledeb.) Ostenfeld. (Hairgrass) 54184 — 26 July
—— On mud and sand shore of temporary fresh-water pond
behind the 6-foot beach ridge.
The inflorescences of these specimens were just starting to
open.
Trisetum spicatum (L.)Richt.* (Spikegrass) 5419526 July —In
clumps on steep slope of sandy scree below the bluffs, one-half
mile west of camp.
Abundant on steep, south-facing, well-drained slopes, on
the edges of beaches, on solifiuction lobes, and especially on
lemming mounds; scattered in the upland vegetation. The
spikes of the local population are dense, purplish and not
interrupted.
Phippsia algida (Soland.) R.Br. 54182 — 26 July — On mud-sand shore
of fresh-water pond behind the 6-foot beach.
Pleuropogon Sabinei R.Br.* (Pond Grass) 54120—14 July — On
muddy shores of an oxbow at Ooyarashukjooeet; 54166 — 24
July —In standing water with Drepanocladus in the Upper
‘ope Ponds.
All ponds with a mossy bottom.
Poa abbreviata R.Br. 54110, 54112 — 14 July —In a sheltered frost-
crack on the sandy seashore beach at Ooyarashukjooeet.
A low, dense tuft with dense, purple inflorescences. 54110
has just started to flower; 54112 is past full flowering.
Poa glauca M. Vahl* (Blue Grass) 54190, 54192, 54225, 54229 — 26
July — On well-drained, sandy scree, south-facing ravines in
the bluffs west of camp.
Common on the uplands, but most conspicuous on disturbed
soil and widespread on solifluction forms and frostings. It
are
is a large, coarse, tufted plant with coarse inflorescence.
r areas on terrace and ridge-tops in our study area.
Poa Hartzii Gandoger, var. vivipara Polunin (Viviparous Bluegrass
_ 94211 25 July—With Ozyria on wind-swept low dune
sand, of the Little River.
: D to Polunin’s material, this has longer, more
_ Pointed ligules and whitish leaf-sheaths. It grows in tufts.
is It does not match well, but fits best here.
SOUTHERN BYLO1T ISLAND 85
Poa arctica R.Br.* (Arctic Bluegrass) 54216 — 26 July — Bog in mesh
of frost-crack pattern near the Upper Phalarope Ponds;
54219 — 26 July—On steep, sandy scree at the base of
ravines in the bluffs west of camp.
ried habitats: well-drained sandy scree, frost-churned
Pasi beaches, barren ridge-tops, dry upland slopes, and low
places with peaty, sandy soil; widespread but nowhere in
dense stands.
var. vivipara Hook. 54213 —25 July —In peaty, sandy
soil in hollows in low dune area northeast of the Little River.
My specimens have rather broad, angularly divergent
leaves. The other grass at this site was vivi
Fernald used ae rigens Hartm. which Porsild id Sara
list in synonym
Dupontia Fisheri we var. aristata Malte ex Polunin (Marsh Grass)
168 — 24 July — On wet margins of the Upper Phalarope
Ponds; 54217 — 25 July — On muddy shore of pond in frost-
crack polygon mesh on Tui-Tui Tabletop.
This material is hard to key out, because as Polunin says
“The genus lacks an awn.” The lemmas in my material have
long awns.
Puccinellia Langeana (Berl.) Sorensen
(syn. P. paupercula (Holm) Fernald). (Goose Grass) 54183 — 26
July— On muddy, sandy shore of temporary pond on the
6-foot beach,
tiny, dense tufts are less than 5 em. tall, panicle is
more than one-half the length of the culm, branches are not
re 0.8 mm. long. Serensen’s key in Porsild (1957) is un-
paren and Swallen’s key in Hitcheock (1950) does not
work either. My material fits well with Polunin’s (1940)
discussion, in which he calls this material P. paupercula.
In Polunin (1959), he treats it as P. Langeana, and my
material matches his description.
Festuca brachyphylla Schultes (Fescue Grass) 54111 —14 July —In
heap, a previous year’s nest of Red-throated Loon (Gavia
stellata), the site being covered by water until mid-July.
Plants low and dense; infi Hl and narrow.
Festuca baffinensis Polunin (Fescue Grass) 54194, 54222, 54223 — 26
July — On steep, sandy scree at the foot of ravine in the
bluffs west of camp.
Habitat different from that of F. brachyphylla, but in the
field I considered all Festuca material tobe the same.
Taller than 10 em., has a longer, broader inflorescence than
86 WILLIAM H. DRURY, JR.
brachyphylla and the upper half of the culm has a short,
dense tomentum.
Elymus arenarius L., var. villosissimus (Scribner) Polunin (Strand
Wheat) 5422426 July — On a sandbank at high limit of
ice-shove at the foot of the bluffs west of camp.
On sandbanks and dunes along most of the shore, common
in dune areas on the uplands, and at all three Eskimo camps
which we visited.
Culms of this material are about 25 cm. tall; glumes and
lemmas are soft villous.
Eriophorum Scheuchzeri Hoppe (Mud Cotton-grass) 54121 — 14 July;
54135 —13 July. On mud shores of low wet areas and old
stream channels at Ooyarashukjooeet; 54164 — 24 July — Old
tream channel] at Aktineqjuak; 54177 —24 July — Sandy
delta of temporary stream at the head of Golden Plover Creek.
nm muddy sand bottoms of shallow ponds. Its
spherical heads are less than 2.5 cm. tall; anthers 1 mm.
long, but, as Polunin points out, many scales are more or
less attenuate and pale-margined.
Eriophorum vaginatum L. (Tussock Cotton-grass) 54122 —14 July
— In wet swamps, on river flats, low slopes and hilltops at
Ooyarashukjooeet.
Locally distributed: near Upper Phalarope Ponds and in
stony sedge marshes on the West Ridge. Tussocks 10-15 em.
from the roots to the tops of the heads.
My ee of E. vaginatum in Alaska convinced me that
spis a geographical variety of it. My eastern Arctic
material sau the same, and Polunin (1959) now includes
+ Spi Fernald in vaginatum, reversing his earlier treat-
ment (1940).
angustifolium Roth (Cotton-grass) 54173 — 24 July —In
muddy delta of temporary stream at the head of Golden
Plover Creek.
, Widespread and conspicuous in nearly all places where
in aaa It had
and gre
just started to flower in the hen week of Jul
The Eskimos i
ue weet
peste ee Suds lace weenie
area on top of the bluffs west of camp.
Kobresia is not conspicuous because of its size, but it
is widespread on unstable soils, exposed places, and dry
: (ii marta hte Uieass Metals, Wash scaleg dnl Breit lowe
| gate-ovate, — eign at the base, i.e., pyriform.
SOUTHERN BYLOT ISLAND 87
Carex maritima Gunn 54221 — 26 July — On steep, vegetated, sandy
scree below the bluffs west of camp
To all appearances, the heads are e single, until dissected,
which makes keys to Carex (Fernald, 1950) misleading.
Carex misandra R.Br.* (Brown Sedge) 54125 —14 July —In moss
on uplands at Ooyarashukjooeet; 54159 —21 July —In thin
moss on a barren place on the 100-foot beach on Kungo Hill.
mmon where there is a thin cover of Hypnaceae, on
margins or stable areas in disturbed soil on the dry uplands.
Carex Bigelowii Torrey ex Schwein.* 54203 — 27 July — In wet moss
on depositional slope on the 35-foot beach.
h of my material appears to be transitional between
this species and the next; but my field notes show that the
Bigelowii-type was associated with nutans ex areas and
‘assiope, rather than with wet, mossy marshes.
Carex aquatilis Wahleng.,var. stans (Drej.) Boott ( swale Sedge) 54167
— 24 July — Sterile on pool margins, fruiting on moss ridges
in the Upper Phalarope Ponds; 54172 —24 July —In sedgy
seep at the head of Golden Plover Creek; 54208 — 25 July —
On wet meadow on the 35-foot beach; 54215 — 25 July —
Bog ridges in the Upper Phalarope Ponds.
A most conspicuous and widespread species. It grows in
damp places everywhere, in wet seeps on steep slopes, on
sheltered, mossy slopes; and is the main vegetation of mossy
marshes and river flats. It is an important element in the
diet of Snow Geese.
I agree with Polunin (1940) that much of this material is
not readily assigned either to aquatilis or to —— ss
numbers 54208 and 54215. My material is low; the
tend to aggregate; most of the leaves are below the om
of the culm; they are glaucous and not scabrous; the lowest
bract equals the inflorescence; the terminal spike is 1 em.
long. The problem of intergradation between these two is
conspicuous in the eastern Arctic, bat I have found no con-
fusion between the two in Alaska and the Yukon Territory.
Carex membranacea Hook 54124—14 July —In a wet marsh on a
above the 55foot beach.
yecatored on the egos of tudo, sled
88 WILLIAM H. DRURY, JR.
beaches, frostings, and barren ridges; identified by its flat
leaves and solitary head.
Juncus biglumis L. 54206 — 27 July — On muddy shores of the Little
River, late snow patches, and mud-boils on the 35-foot beach
and Tui-Tui Tabletop.
Juncus castaneus Smith Not represented in my collections, but
recorded on wet mud behind raised beaches and on an
abandoned river channel.
Toffieldia coccinea Richardson Not represented in my collections, but
n several times in barren areas such as terrace edge above
Little River.
Salix reticulata L.* (Net-leaved Willow) 5467 (male) and 5468
(female)—11 July — Under soliffuction lobe on the 35-foot
beach at camp.
Especially important on south slopes, sites with more
shelter and moisture than pure Cassiope areas, also on mossy
ridges in marshes,
The Eskimos call the leaves of this plant Okowyuk, the
same as those of S. arctica, but they do not eat them.
Salix herbacea L. (Herb Willow) 5487 (male) and 5488 (female) —
uly — On west-facing slope in Lark Gully — a late snow
patch.
cteristic of late snow patches, where it forms a golf-
green-like carpet. Occurs also on shady, Hypnum-covered,
shallow frost-cracks, between hummocks of Cassiope, or
under solifluction lobes.
Salix arctica Pall. (Arctic Willow) 5405 (male), 5408 (female) — 17
June — Prostrate mats on low, sandy, well-vegetated, sunny
__ to Salix arctophila Cockerell on the basis of catkin scales
or leaves, but seldom both. My #54180 matches many her-
SOUTHERN BYLOT ISLAND 89
barium sheets of S. arctophila and agrees with descriptions
of several authors. The leaves of all my material are glab-
vary from soft hairy gray-green to bright shining
dark green and glabrous according to site and exposure.
Some caangeigre have —. with denticulate margins.
ause n find no characters in the discussions in
Schneider i: 1921), Fernald (1950), Hultén (1943),
Porsild (1957), and Polunin (1940, 1959) that are not inter
a .
belonged to a population separate from arctica either in
Alaska or the eastern Arctic, I am convinced that S. arcto-
phila is not distinguishable in the areas where I have worked.
The collections I have which might be included in S. arctophila
are parts of populations continuous with S. arctica; they
appear as ecotypes both in the field and when compared
with danmeryees specimens.
is materials is referable to var. kophophylla
(Semeider Polunin, although 5423 has broad-based, glab-
ite lea Further, I hesitate to suggest that two geo-
sma races occur in the same area.
Idlouk, one of our Eskimo companions (Drurys, 1955),
called the young leaves of S. arctica Okowyuk and said
that the local people eat them in the spring. He called the
stems Ookpiyuk and said that people make baskets out of
them, but I could not establish whether he meant his people
or other people. He called willow cotton Shuput and said
they use it for wicks in their seal-oil lamps. Occasionally
cpa the roots, which ny eall Eyeroi
Salix Richardson Hook., var. McKeandii Polunin® (Shrub Willow)
shukjooeet; on low es of , and in the
most sheltered places at Akti
Ooyarashukjooeet.
My material has short, hairy capsule pedicels as Polunin
(1940) pointed out. Porsild (1957) says that the eastern
Arctic material is “incorrectly said to have shorter and
hairy-pedicelled capsules.”
Oxyria digyna (L.)Hill Speer Sorrel) 5421 — 25 June — Sunny,
“ the foot of the bluffs west of camp,
growing in silty sats 54212 — 25 July — Wind-swept area
of low dunes northeast of the Little River.
ee
ravine slopes, frostings, raised beaches, and frost-heaves.
The plants are a rich purple and the Eskimos, who call
it Kungalik, eat it in the spring as a salad, savoring the acid
te. They also use it as emergency
Polygonum viviparum L. (Alpine Knotweed) 54145 —13 July —On
90 WILLIAM H. DRURY, JR.
vegetated lower scree, below slate outcrop two miles from
the mouth of the river at Ooyarashukjooeet.
Disturbed soil in ravines, frostings, raised beaches, sunny,
grassy slopes, and in open spots above and below solifluction
obes.
Silene — (L.)L., var., excapa (All.) DC.* (Moss Pink) 5478 —11
y — On stabilized, sandy landslide below the ravines in
ie bluffs west of camp.
Widespread but scarce on sunny, well-drained sites such
as frost-churned soils of raised beaches and tops of solifluc-
tion lobes.
Lychnis affinis Fries (syn. furcata Raf.) 54109 —14 July — On edge
of frost-crack on the lowest sea beach at Ooyarashukjooeet ;
on stabilized, sandy scree and a fans below the
bluffs west of camp; and on terrac ges.
Stems are thin and glabrous; ‘ecsea mostly basal, and
flowers nodding.
Cerastium alpinum L.* (Mouse-ear Chickweed) 5406—17 June—
On sandy, sunny, well-vegetated slopes west of camp; 5459
—5 July — On slide rock at 2600 feet on Mt. Thule; 54100
—1l1 July — On slaty alluvial fan at the foot of ravines
west of camp; 54197 — 26 July —On sandy scree, in open
vegetation at the foot of the bluffs west of camp.
Cha: i
It is densely tangled hairy on all surfaces.
Stellaria longipes Goldie* (Star Flower) 5480—11 July —In open
place on vegetated steep slope below the bluffs west of camp;
54103 —14 July —In a frost-crack on the lowest beach at
Ooyarashukjooeet.
Inconspicuous but widespread in open and closed vegeta-
tion on sunny, dry, grassy slopes, and especially common on
lush, Pies ctbgsr gl aeoeks of lemming mounds; scarce on dry ridges in
Bisa aad Miia ae Rottb. 54226 —26 July — On muddy sand by tem-
porary fresh-water pond on the 6-foot beach; scattered but
locally common on shores of temporary ds.
ao Wahlenb. 54103a—14 July —In frost-crack on
at Ooyarashukjooeet.
Scattered but locally common; prostrate on damp, sandy
Arenaria sincera L. (Sea Beach Sandwort) 5475—11 July —On
vat at — mouth of Lark Gully.
sea beaches just above high tide.
ona i (Wahlent Sm 54105a — 14 July — In a frost-crack
J kj
jooeet.
Sagina intermedia Fenzl.* thine June — (previous year’s fruit)
SOUTHERN BYLOT ISLAND 91
— Sandy beach on the delta of stream out of Lark Gully;
54105 —14 July —In a frost-crack on the lowest beach at
Ooyarashukjooeet; 54210 — 25 July — On low sand dunes in
wind-swept place northeast of the Little River.
A tiny wiry plant in damp sand, the edges of mud-boils
and on old river deposits.
Ranunculus hyperboreus Rottb. (Water Buttercup) 54232 — 26 July —
On sandy mud on the edge of a temporary fresh-water pond
on the 6-foot beach; on many sand- or mud-bottomed shallow
ponds.
supe — L. (Snow Buttercup) 5435 — 22 June — On peat
's beside frost-cracks on the 16-foot beach; on deposi-
raat slope below the 35-foot beach; and in frost-cracks in
the 16-foot ee crest; 54113 — 14 July — In Drepanocladus-
Tomenthypnum. bog in an abandoned river channel at Ooyara-
shukjooeet; 54115 — 14 July — On moss and mud in late snow
area next to creek draining into the river at Ooyarashuk-
jooeet; 54176 — 24 July — In sedgy, wet seep at the head of
Golden Plover Creek; 54181—25 July —On lower slope,
beside Golden Plover Creek.
Common in wet, mossy places where snow persists after
the main paca on peat ridges, in frost cracks and in moss
hummock a: a
Numbers "BALI6 and 54181, both of which grew in creek
banks, are small and delicate; = match specimens of R.
Sabinei in the Gray Herbarium, except that they have brown
eurly hairs on the sepals score of pale brown-tan sparse
hairs of R. Sabinei.
Ranunculus sulphureus Solander apud Phipps 54131—13 July —
In wet hummocks area, with Salix arctica, on the floodplain
of the river at Gupcraskakjooee: 54175 — 24 July—In
sedgy seep at the head of Golden Plover Creek.
Papaver radicatum Rottb. (Poppy) 5414— 25 June — Sunny, sandy
scree on the edge of the bluffs; 5452 — 5 July — On dry slope
seree at the foot of the bluffs west of camp; 54145 — 13 July
—On vegetated sandy scree fan below slate rocks along the
Common and widespread on (1) disturbed soils of frostings,
alluvial fans, sandy slopes below the bluffs, and frost-churned
soil next to frost-cracks, (2) & areas: raised beaches
and ridge-tops, and aii scattered in dry Dryas, Salix arctica-
Cassiope areas of the
In a dozen places sedis cna clumps with white flowers, e.g.,
5452a, var. albiflorum Lange. This is certainly not equivalent
to a geographical variety and Polunin’s quotation from 1 Abbe
describes the genetic activity of a sport at best.
WILLIAM H. DRURY, JR.
Hae peak of flowering was about July 10. Flowers were
and young fruit developing by July 22.
Cochin omni L., var. groenlandica (L. i Gelert apud Anders-
sselman
—17 June—On sand in a our
pile sneaky a bluffs west of camp; 5465 — 2 July — On san
and cobbles in front of an Eskimo house at Sermilik; ror
— 14 July — In a frost-crack on the lowest beach at Ooyara-
shukjooeet.
Damp soil of raised beaches or alluvial fans; often asso-
ciated with Eskimo settlements.
The siliques are ovate and acute at the tips; plants de-
pressed rosettes with the siliques barely raised above the tuft.
Eutrema Edwardsii R.Br.* 54148 —13 July —In mossy ridges be-
een mud-boils on hilltops about three miles inland at
Ooyarashukjooeet.
Widely scattered as individuals, on mossy or muddy places
on uplands.
comers bellidifolia L. 54214 — 25 July — On mossy ridges in a bog
east of the Little River.
kogiovns arctica (Wormskj.)S. Watson 5451—29 June — and
5
Draba alpina
493 —11 July —both on sandy alluvial fans below slaty
sediments in the ravines in the bluffs west of camp; con-
spicuous among the weedy flowers.
L., var. nana Hook.* (Yellow Mountain Mustard) 5446
— 29 June — Almost buried in other — (Salix arctica,
Potentilla hyparctica) on sunny slopes of alluvial slope
below the bluffs west of camp; 5497 — it July — Slaty scree
at the foot of the bluffs west of camp; 54144a — 13 July —In
stems, d fruit.
Idlouk called this, and all little yellow Gee nuk nyu-
Draba subeapitata Simmons (Mountain Mustard) 54107 — 14 July —
In sand in a frost-crack on the lowest beach at Ooyarashuk-
~
A large series — all typ‘cal.
Draba lactea Adams (Mountain Mustard) 5412314 July—In
ay Eriophorum
y
mossy areas, especially frost-cracks, on the uplands of Plover
Plateau; 54170 — 24 July — On moss ridges at Upper Phala-
Tope Ponds.
eee: in upland vegetation.
Polunin (1940) includes this i in D. = Wulfen, but
T am not following his fine treatment tt because my material
SOUTHERN BYLOT ISLAND 93
matches the D. lactea material in the Gray Herbarium
and not D. fladnizensis. Fernald (1950) treats this as D.
icha.
fl
Draba nivalis Liljebl.* (Common Mountain Mustard) 5413 and 5416
—25 June — (opening flowers fixed) 5445 and 5449 —
June — (opening flowers fixed) 5479, 5481 and 5482—11
July; 54185—26 July. All these numbers are from the
sunny, south-facing slopes of fine to coarse sand alluvial fans
and screes at the base of the bluffs west of camp; from nearly
closed grassy Salix arctica vegetation to bare places with
it the
isolated plants, on all bu most open and active soil.
54108 —14 July —In frost-cracks on the lowest beach at
Ooyarashukjooeet.
This is one of the two really common Drabas a is
readily recognized in the field. It prefers bare pa‘
Draba glabella Pursh, var. brachycarpa (Rupr.) Fernald teem
Mustard) 5414413 July—In tussocks and around rock
remnants on hilltops three miles inland at Ooyarashukjooeet.
Resembles D. cinerea closely, but habitat is distinct.
Draba cinerea Adams (Ashy Mountain Mustard) 5403 and 5407 —17
June; 5415 — 25 June (opening flowers fixed) ; 5447 and 5448
— 29 June (opening flowers fixed) ; 5484 and 5498 — 11 July;
54186 — 26 July— All on sandy screes and alluvial fans
below slaty sediments in bluffs west of camp; 5459a — 5 July
— On slide rock at 2600 feet on Mt. Thule; 54142, 54143 and
5415113 July —On vegetated, sandy alluvial fan below
slaty rocks on the river bank above Ooyarshukjooeet.
A large Draba, a in vegetated alluvial or scree fans.
It grows in Poggorsie igs Salix arctica and other mat plants
more than D. nivalis and avoids the open sandy patches.
The flowers vary from rich cream as they open to white
in some several days later. The siliques are hairy. Porsild’s
Lanna key, which otherwise works well, does not include the
ream flower color; the scapose infloresence which he uses
be — cinerea from groenlandica is not useable in my
rial.
Arabis pots (Richardson) Gelert, var. pubescens (Wats.) Gelert
5491 — 11 July — On sandy alluvial fan eee a ravine cut
into slaty rocks on the bluffs west of cam
On fine-grained, pcsGe God. ek ct en cache
soils.
Sparsely hairy, the younger leaves being much more
hairy than the older.
Erysimum Pallasii (Pursh)Fernald 5409—17 June, and 5496 —11
July — Both on open, unstable alluvial fans at the foot of
slaty bluffs west of camp.
1 umerous. :
The flowers are magenta and siliques radiate from the
94 WILLIAM H. DRURY, JR.
rosette of leaves. The magenta flowers raise a question of
how they are pollinated (Epilobium and Pedicularis as well),
because insects are said not to see this color. Presumably
they respond to the blue, but if so, why the red at all?
Braya purpurascens (R.Br.) Bunge apud Ledeb. 5450—29 June
(flowers fixed); 5486—11 July— Both on mud, sand and
angular pebbles of scree and alluvial fan at the base of
ravines in the bluffs west of camp; scarce.
Flowers and fruits are weakly erect, 5 em. above the ros-
ettes; the siliques are long compared to the material in the
Gray Herbarium, and they taper to their tips.
Saxifraga rivularis L. 54102—14 July —On sand in a frost-crack
on the lowest beach at Ooyarashukjooeet; 54119 — 14 July —
Tn a late snow patch on the bank of a creek tributary to the
river above Ooyarashukjooeet; 54228 — 26 July —In damp,
muddy sand by loon’s nest on the temporary fresh-water pond
on the 6-foot beach.
Sites are muddy, cold, and wet late in the season.
Saxifraga cernua L.* (Nodding Saxifrage) 5456 —5 July — On west
slopes with seasonally advanced vegetation above the 56-foot
beach on Kungo Hill; 54104 — 14 July — Im a frost-crack on
the lowest beach at Ooyarashukjooeet; 54149 —13 July — On
the edges of mud-boils, among hummocks, on hill-tops two
miles inland from Ooyarashukjooeet; 5418726 July —
Steep vegetated scree below the bluffs west of camp.
Scattered individuals in grassy, closed vegetation or damp,
fine-grained open soils.
5456a —5 July, and 5420527 July — Muddy shores of
the Little River are slender, short, glabrate, and late-flowering.
They match var. exilioides Polunin, and differ from the typi
Population but grow mixed in with typical plants. Their
growth may be affected by local conditions, their genetic basis
simple, or there may be apomixis.
Sazifraga caespitosa L. 5490 —11 July — Low, wet place in front of
alluvial fan below Lark Gully; 54136 —13 July — On Dryas-
area at the 10-foot beach at Ooyarashukjooeet; 54188
—26 July— Sunny, vegetated scree below the bluffs west
of camp.
All material is low and has one or rarely two flowers: forma
f uniflora (R.Br.) Engler & Irmsch.
Sarifraga stellaris L., var. comosa Retzius* (Star Saxifrage) 54116
‘ Matches S. foliolosa R.Br., because most of the flowering
SOUTHERN BYLOT ISLAND 95
head is bulbils, which raises the question whether this popu-
lation should be a species or a Vv: a
Saxifraga hieraciifolia Waldst. & Kit.* (Hawkweed Saxifrage) 5455
—5 July—On sunny frosting above the 56-foot beach on
Kungo Hill; 54117—14 July—In a late snow area on a
creek bank above pensalenere ec 541302 — 14 July —On
dry Dryas-Salix raised beach
ooeet.
54117 is young with clustered terminal inflorescence; its
coarse stem and leaves indicate sai sees but the speci-
mens are hard to separate from Saxifraga nivalis.
Saxifraga nivalis L.* (Snow Saxifrage) 54 ee June — Sunny,
vegetated slump below bluffs just west of camp; 5454 —
5 July —On dry slope of Kungo Hill at about the 135-foot
beach; 54118 — 14 July — Im a late snow patch on creek bank
above Ooyarashukjooeet ; 54130 — 14 July — On a dry raised
beach with Dryas, Salix arctica and Sazifraga tricuspidata
at Ooyarashukjooeet; 54198 —26 July —In vegetated scree
below bluffs west of camp.
Widespread on sunny slopes soon free of snow and soon dry.
54198 is large and coarse, also old; the rest of the material
is slender and low, but I cannot separate out any population
such as var. tenuis Wahlenb.
Saxifraga tricuspidata Rottb. (Spiny Saxifrage) 5473—11 July —
On sunny scree below the bluffs; 54138 —13 July — On dry
raised beach about 25 feet above sea level at Ooyarashuk-
jooeet; 5420227 July—On dry frosting on the side of
Kungo Hill.
Scattered but abundant where it appears on dry, sunny,
sandy soil.
54202 has the lateral teeth and spines on the leaves almost
completely lacking: f. subintegrifolia (Abrom.) Polunin.
The Eskimos make a brew out of the whole plant, and call
it kakilangnakotee. Kakil refers specifically to this species.
Idlouk allowed that the brew is pretty grim.
—e — Willd. 54150 —13 July — On aie of mud-boils
a hummocky area on the hilltops two miles north of Ooyara-
proses
Widely-scattered individual plants.
Sazifraga Hirculus L., var. propinqua (R.Br.)Simmons 54137 —13
July —In wet sedge area on the floodplain at Ooyarashuk-
jooeet.
Saxifraga oppcsitifolia L. 5412—17 June—On sandy scree with
angular fragments below the bluffs west of camp; 5460 —5
July — On slide rock at 2600 feet on Mt. Thule.
the
little shelter, (b) in the densely vegetated uplands, (c) on
sunny slopes, (d) on unstable ravine serees, (e) on the drift-
96 WILLIAM H. DRURY, JR.
ing sands of floodplain washes, and (£) in wet Tomenthypnum
ridges in the water-soaked bogs. It grows everywhere we
found plants, other than the salt-tolerant strand plants, and
is a conspicuous part of the vegetation almost everywhere
it grows because of its cushions and purple flowers which
continue to bloom through July. During June many clumps
were in full bloom while the plant’s base held snow and ice,
and while the roots were embedded in frozen ground.
The Eskimos call it Owbiletungwak and eat the flowers
oat —— occasionally — calling the root Eyerok.
L. 54127 —14 July —In eS
area on the floodplain of the river at Ooyarashukjooee'
These specimens have 5 and 6 stamens, and cannot piri to
var. tetrandrum Lund, which is the only form Polunin (1940)
found in our area.
Potentilla hyparetica Malte* (Cinquefoil) 5402—17 June — 0:
sandy, steep, sunny banks in clumps with Salix arctica,
grasses and many flowers below the bluffs.
We iene on steep upland slopes, on sheltered places next
0 barrens, and on the edges of lemming mounds.
attire (1940) calls this P. emarginata Pursh; Fernald
(1950) and Polunin (1959) call it hyparctica.
Potentilla rubricaulis Lehm. (Cinquefoil) 5411 — 17 June — On sandy
uvial fan at the foct of ravines in the bluffs west of camp.
e denseness of tufts and number of flowers seem to be
unreliable characters in Potentilla, but my material matches
the specimens of this species in the Gray Herbarium, and it
differs from other species.
Potentilla Vahliana Lehm. (Lemon Cinquefoil) 5477—11 July —In
open areas on = sunny slopes of ravines cut into the
bluffs west of camy
Its long hairs ens ‘all surfaces, and beards at the tips of
the teeth on the leaves make it easy to recognize in the field.
Found only at the bluffs.
Dryas integrifolia M. Vahl. (Avens) 5419 — 25 _. sunny,
vegetated slopes below the bluffs west of c:
Primary on sunny, well-drained sites on ele to steep
vegetated slopes, and over most of the uplands. It forms
the cl characteristic closed closed vegetation of many places between
mud-boils, ele ligne, hs tone ob setae then, or the
sheltered sides of ridges next to frost-cracks.
The peak of flowering was aoa 10 July, and the fruits
- were just appearing, still twisted, when we left.
Examination in the field and at the Gray Herbarium shows
this i. has napie or dentate leaves, and glabrous or
one tuft. I can find no support for
ing the os into intermedia, canescens and
en agetioass alpinus L. (Vetch) 5420—25 June, and 5492a—11 July
SOUTHERN BYLOT ISLAND 97
— Sunny, vegetated screes below the bluffs west of camp.
Abundant on sunny, sandy, well-drained slopes all over
the area, on sandy alluvial fans at the mouth of ravines
below the bluffs west of camp, on nt sandy soil on
the river floodplains, and on sand dunes.
This is a conspicuous weed in oratoad but not actively
churned =o and thus it is absent from the bare soil of
frost patt
Its panache peak was 20-25 July.
Oxytropis Maydelliana Trautv. (Locoweed) 5457 — 5 July — On edges
of mud-boils and frostings on the 135-foot beach on Kungo
Hill; 54196 — 26 July — On steep vegetated scree below the
bluffs west of camp.
A conspicuous flower on sunny, well-drained slopes from
the sides of be Hill to vegetated screes in the ravines,
and common with Dryas in vegetation stripes on the uplands.
The hence: teak came 20-25 July.
The Eskimos call this Eyerok and dig and eat its thick root
which, when fried, tastes like parsnip. The name probably
means root because they call the root of Salix arctica and
Sazifraga oppositifolia the same, and eat them also. They
prefer this one.
dit
Empetrum nigrum L., var. ~p ee (Crow-
berry) p17 26 gpebrciats dry Cassinpe warn below the
55-foot beach on Kungo Hill.
This is the only station, but Leah (Idlouk’s net said
that it is common at their main camp on the south shore
of Eclipse Sound. They call it paonga and eat berries.
Epilobium latifolium L. (Willow-herb) 5495—11 July—On slaty
ree at the foot of the bluffs west of camp.
Abundant on sand and gravel alluvium at the mouth of
creeks and ravines in the bluffs, on the abandoned channels
and gravel bars of rivers, and on wet shores of fresh-water
ponds behind raised beaches.
— — Samuelss. pes Oe July — On sedge-moss, wet
t the mouth of a tributary creek above Ooyarashuk-
ea
Polunin (1940, 1959) keeps this as a variety of E. davuri-
cum Fischer ex Hornem.
Cassiope tetragona (L.) D. Don (Bell Heather) 5458 —5 July — On
edge of mud-boils and solifiuction lobes on sunny sunny slope near
the 135-foot beach on Kungo Hill; 5459 —5 July — On slide
rock sil ioc Mt. Thule.
98 WILLIAM H. DRURY, JR.
cracks, the vegetated pattern of soil stripes and mud-boils,
polygon meshes, and the edges of solifluction lobes,
The complete lack of Cassiope is an outstanding feature
of stream banks or places so sheltered from the sun that the
snow lies until the first of July.
The flowering peak was 15-20 July.
The Eskimos call this a gutteral chyuktak and fill bags
with it to make mattresses. They know the widespread use
of its waxy branches to make an emergency fire.
Vaccinium uliginosum L., var. alpinum Bigel. (Bilberry) 5469 —11
uly — Under the slope of a solifiuction lobe on the side of
the 35-foot beach at camp.
Scarce and restricted to sheltered hollows in the ravines
in the bluffs, and the lips of raised beaches or solifluction
lobes; rare in frost-cracks where most of its associates are
Polunin (1959) includes this in A. maritima (Miller) Willd.
I did not look into the question.
Martensia maritima (L.)S.F. Gray, var., tenella Fries 5489 — 11 July
— Sandy sea beach at Ooyarashukjooeet.
Characteristic of sea beaches above high tide.
Pedicularis sudetica Willd. (Fernweed) 54128 —14 July —In sedgy
. Their
flowers are pale purple with a magenta end on the long-
toothed helmet and magen' ij
Pedicularis lanata Cham. & Schl.* (Woolly Fernweed) 5437 — 22 June
— On exposed, mossy Luzula area above the 135-foot beach.
Common and widespread, growing singly in the Salix
arctica-Drayas-Cassiope uplands. It is one of the first plants
: to flower outside the sunny south slopes. Flowers wholly pink.
Pedicularis Langsdorfii Fisch., var. arctica (R.Br.) Polunin (Marsh
Fernweed) 54155 — 20 July — In wet moss, sedges, and Salix
est Ridge.
Pedicularis hirsuta L. 54139 — 13 July — On Dryas-Saliz arctica low
alluvial fan slope below the terrace, one mile above Ooyara-
shukjooeet, in a weedy stand.
Flowers small, lavender with a paler lip and purple
is slender, with flowers sticking out to
SOUTHERN BYLOT ISLAND 99
In Dryas-Salix arctica on wet, hummocky area on a deposi-
tional fan about a mile above Ooyarashukjooeet.
The only station for the species in flower, but it is widely
scattered as individual very fern-like leaves in hummocky,
mossy areas on the uplands. The silver spots on the leaves
add to the resemblance to a fern because they suggest
sporangia. If this were an animal, it surely would be credited
with mimicry.
Its pale puff flowers have a pale purple tip to the helmet.
Erigeron uniflorus L., var. eriocephalus (J. Vahl) Abromeit (Flea-
bane) 5474—11 July, and 54189a2—26 July—On sunny
screes and steep ravine sides in the bluffs west of camp;
54231a— 26 July— Mud-sand shore of temporary fresh-
water eee on the 6-foot beach.
The ray flowers are pale lavender; the young material
(5474) — heads 1.5 em. across and the older heads are 2-2.5
ross. Cronquist (1947) and Polunin (1959) include
this in E. uniflorus.
Erigeron compositus Pursh. (Daisy) 5492—11 July —
sandy alluvial fans at the bottom of the ravines in the blaf bluffs
west of cam
Antennaria hnanane A.E. Porsild (Pussytoes) 5418 — 25 June, and
72 —11 July — On sunny, well-drained, sandy slopes above
the depositional fans at the base of the bluffs west of camp;
54200 — 27 July — On dry frostings on raised beaches on the
side of Kungo Hill.
5418 and 5472 match herbarium material; 54200 includes
labeled A. angustata in the Gray Herbarium, but seem to me
to belong with other specimens. Polunin (1959) includes
labradorica (where he placed this in 1940) in Ekmaniana.
Arnica alpina (L.) Olin, var. angustifolia (Vahl) Fernald 5464 — 29
June, and 5476 —11 July.
A conspicuous flower on the sunny, oer 4 slopes of the
bluffs west of camp where the vegetation of grasses, Salix
arctica and Oxytropis has stabilized the slopes.
Senecio congestus (R.Br.)DC., var. booqatnn (L.) —— 54129 —
14 July — Im sedgy, mossy old stream channel near Ooyara-
shukjooeet.
Does not match Polunin’s f. polycricos.
Taraxacum lacerum Greene (Dandelion) 5410 —17 June, 5499 — 11
July, 54191 —26 July, and 54220 —26 July — All from un-
stable, open fans and vegetated ravine sides on sandy screes
aty bluffs west of camp.
5499 and 54220 differ in that many plants lack the horns
on the inner involucral bracts, but all belong to one variable
population. i
100 WILLIAM H. DRURY, JR.
SUMMARY
Frost Features
Washburn (1956) has divided minor topographic features
produced by frost action into solifluction forms and struc-
tural soils. On Bylot Island, solifluction forms (crescents,
tongues, and transverse terraces) modify nearly all slopes
below the bedrock mountains into microtopography to which
the vegetation responds in detail. Soil creep and mud
avalanches, covered by open nets or closed mats of plants,
nearly obliterate some old raised beaches, while mincr
alluviation creates forms a foot across and an inch high.
Frost-riving and solifluction have sculptured a gently-sloped
plateau cut by steep stream valley sides on southwestern
Bylot. These strongly resemble the so-called peneplain rem-
nants of eastern parts of Canada and the United States, and
suggest that cryoplanation whose base level is the lower
altitudinal limit of periglacial arctic erosion processes may
be the source of the raised surfaces, not a previous stream
erosion cycle.
Patterned ground (Strukturboden) is expressed as non-
sorted circles, polygons and stripes, or as frost-crack
patterns. Circular, bare areas, usually of fine material, mud-
boils, are active during spring thaw and may appear in
aggregates as non-sorted polygons (bare patches alternating
with mat vegetation, Arctic Willow, Avens, mosses, and a
few scattered flowers). On gentle slopes, they become
elongated into non-sorted stripes, tan angular fines and
pebbles, alternating with stripes of vegetation. In some
these areas are hundreds of yards long and several
Square miles in area,
During the thaw the soil is saturated with water down to
the top of persistent frost. Certain bare spots thaw rapidly
to a depth many times that of the vegetation-covered sur-
roundings, and islands of frozen ground may be underlain
by thawed ground. Water flowing over, through, out of,
and into the soil, or some related force, thrusts fines up from
depths and creates mud-boils, solifluction lobes and mud
_ avalanches which appear related in origin. The sediments
mud-boils (usually fines at the surface ‘but often coarse
at draw together into a cheese-like surface and
crack progressively from the margins and around stones —
SOUTHERN BYLOT ISLAND 101
thus suggesting colloidal action (Steche, 1933). Cracks on
these surfaces (Zellenboden) are formed both by stresses
and strains of alternating freeze-thaw and wet-dry. The
instability which results prevents plant growth.
A master pattern of frost-cracks (ice-wedge polygons,
Spaltenboden, Taimyrpolygone) exists on all surfaces of
unconsolidated material, most clearly on the youngest
beaches. They run down or across the slope and (a) meet
at right angles on flat beaches, or (b) form concentric cracks
following the contours crossed by cracks radiating from the
center on hilltops.
(1) The close similarity of this pattern of frost-cracks to
cracks formed when the sea ice yields to pressures and
strains at the tidal zone, (2) the lack of deformation of
sediments, and (3) the uniformity of cracks disregarding
sediment sizes, suggest that the pattern results from the
fracturing of perennially frozen ground under strains, such
as differential isostatic readjustment. Expansion of ice in
the cracks by addition of annual increments and yearly
formation of new contraction cracks caused by violent drops
in temperature raise a ridge beside the cracks of the master
pattern.
Sub it hic modification on exposed _hill-
slopes results from formation of secondary contraction
cracks parallel to and on the edges of the ridges of the large
patterns. Year by year these advance toward the center of
the mesh where snow persists. As centers become smaller
and are obliterated, the former master pattern persists as
traces in lichens, or a new high-centerd polygon results,
resembling those of northern Alaska (Black, 1952). In
areas heavily grown to mossy and sedgy vegetation, modifi-
cation results from peat formation associated with differen-
tial plant growth on the microtopographic features according
to differences in moisture. Peat ridges form on the rectang-
ular patterns beside frost-cracks and act as dams and create
a rice-paddy effect. The ponds trapped this way lead to
deeper thawing of the ground; and regularly spaced, rec-
tangular thaw-ponds (Muller, 1947; Wallace, 1948) are
produced.
Vegetation
The vegetation of southern Bylot Island is the result of
102 WILLIAM H. DRURY, JR.
responses of plants as species and individuals to dati
in wind and sun exposure, soil stability, moisture availabil-
ity, and time of year when sites become free of snow. The
variations in the vegetation from ridge-top to valley-bottom
are often the same as that from the center to the mounds on
the margins of the polygons formed by frost action. Sedge
marshes occur around ponds on the tops of hills, and vege-
tation-free frostings are found where the east wind can blow
through the bottom of a valley.
The minor elevations created by frost-heaving or slump
lobes produce drainage and exposure effects which greatly
influence the vegetation. On dry tops of such forms, dry
sites are produced ; the steep slopes are well supplied with
moisture and are covered with a rich mat, usually of Bell
Heather. Minor hollows collect moisture and hold it above
the perennially frozen ground, allowing sedge and moss
vegetation to colonize,
Most of the vegetation is made up of ten species; 85 per
cent is made up of thirty-four. As is universally true, some
Species grow almost everywhere on Bylot, others are rare
and local; some have site significance, others seem to have
none ; some sites are rich in species, most are poor. The flora
of the area is rich in calciphiles,
Vegetation types selected by site and aspect agree with
habitats selected by birds, and suggest that birds’ selection
is based on life form and the t pography of the vegetati
Cryoturbation makes a homogeneous subsoil but the
resulting microtopography creates local physical conditions
reflected in great variability between and within plant asso-
ciations over a short distance. Dominance, site indication
and frequency are all useful, but I find the concept of succes-
Sion of little value in d ibing this vegetati
e recorded 101 species of vascular plants.
Contribution No. 38 from the Hatheway School of Conservation Education, Massachu-
setts Audubon Society, South Lineoin, Massachusetts.
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CONTRIBUTIONS FROM THE GRAY HERBARIUM
OF HARVARD UNIVERSITY
Edited by
Reed G. Rollins and Robert C. Foster
NO. CXCI
THE SOUTH AMERICAN SPECIES OF ERIGERON
Orro T. Sousric
THE GENUS ERIOSORUS IN COSTA RICA
By
EpirH ScAMMAN
TAXONOMIC FERN NOTES, III
RoLLa Tryon
A MONOGRAPH OF THE FERN GENUS JAMESONIA
By
Altice F. Trron
Published by
THE GRAY HERBARIUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASS., U.S. A.
1962
CONTRIBUTIONS FROM THE GRAY HERBARIUM
OF HARVARD UNIVERSITY
Edited by
Reed C. Rollins and Robert C. Foster
NO. CXCI
THE SOUTH AMERICAN SPECIES OF ERIGERON
By
Otto T. SOLBRIG
THE GENUS ERIOSORUS IN COSTA RICA
By
EpITH SCAMMAN
TAXONOMIC FERN NOTES. Ill
BY
ROLLA TRYON
A MONOGRAPH OF THE FERN GENUS JAMESONIA
By
ALICE F. TRYON
Published by
THE GRAY HERBARIUM OF HARVARD UNIVERSITY
CAMBRIDGE, MASS., U.S. A.
Issued November 23rd, 1962
THE SOUTH AMERICAN SPECIES OF ERIGERON
Otto T. SOLBRIG
The genus Erigeron is one of the largest and taxonomi-
cally most difficult genera of the tribe Astereae of the family
Compositae. Primarily a montane group, it is common in
eastern and western North America, Central and South
America, Europe and Asia.
In spite of its abund and wid d distribution (or
possibly because of it) the generic boundaries are hard to
define. Technically, Erigeron is usually characterized by its
triangular-tipped stigmas with their surfaces covered by
short and stubby collecting-hairs ; by the possession of stout
imbricated involucral bracts, arranged in approximately
two series; and by its heterochromous heads, that is with
the ligulate flowers of a different color from the tubular
ones. If these criteria were applied rigorously, many species
hitherto referred to Erigeron would have to be excluded.
Such a procedure would lead to the creation of a series of
small, often monotypic, genera, not adding to our knowledge
of the evolution and phylogeny of the group, but probably
increasing the taxonomic confusion. It is therefore best to
maintain the genus as now established, until more and better
information relating to these species is available.
Erigeron is very closely related to two other large genera,
Aster and Conyza, and the separation of these three genera
is not always an easy task. On the basis of characters of the
style and the involucral bracts it is usually possible to decide
if a species belongs to Aster or Erigeron: the styles being
elongate or subulate and the involucral bracts more rigid
in Aster. The separation between Erigeron and Conyza is
more difficult. Classically, these two genera were separated
according to the presence or narra of ligules a ere ray
flowers. Although this is app tl litati
careful observation revealed the seid of minute ligules
in the ray flowers of species usually considered as Conyza,
and in some cases the presence or absence of ligules can
be only determined after careful microscopic study. The
absolute presence or absence of a ligule is not correlated
with any other character of the genus. This led Cronquist
(1943) to redefine the generic boundary between Erigeron
and Conyza, according to the criteria of Bentham and Hooker
4 OTTO T. SOLBRIG
(1873). Rather than by the absolute presence of a ligule,
Erigeron is characterized by the possession of ray flowers
with ligules larger than the tubular flowers, usually arranged
in one series. This new arrangement includes in Conyza, all
the species previously treated as belonging to section
Caenotus of Erigeron, including such well known weedy
species as Conyza(Erigeron) canadensis and C. bonariensis.
This new arrangement seems to be a more natural one,
although reports of hybrids between Erigeron acris (sect.
Trimorphaea) and Conyza canadensis (sect. Caenotus) cast
a certain shadow of doubt on the “naturalness” of this new
generic boundary.
In any case, it is clear that more study, especially of an
experimental nature, is needed to solve this problem. The
generic boundaries in the whole tribe (in the whole family
unfortunately, with notable exceptions) are in need of much
study, and the problem will not be solved from morphology
alone. Cronquist’s arrangement so far seems the most prac-
tical division of these two very closely related genera. This
arrangement tends to separate the plants into two ecologi-
cally different groups: the montane, largely perennial
Erigeron, and the more tropical, largely annual or biennial
Conyza.
Erigeron has its center of distribution in western North
America (Cronquist, 1947 ). The South American species
occupy a similar habitat with the exception of those belong-
ing to section Leptostel : i , E. tweediei, E.
meyeri, E. t is and E. posportoi). This sec-
tion has diverged into a completely new habitat, the humid,
often swampy, openings of the tropical and subtropical
South American forest. Nevertheless it is not found in the
Amazonian basin, but only from Goids in Brazil south to
Uruguay and west to the eastern part of Bolivia, usually
on higher elevations (around 1000 m.). The new ecological
requirement has produced taller plants (up to 4 m.) with
large leaves, while the floral characteristics have changed
relatively little. Nevertheless, certain doubts as to the
= serene cpa of Leptostelma have been raised (see
_ Another niche occupied by Erigeron in South America
is in the oceanic islands: Galépagos with one species, Juan
Fernandez with four and, in the Atlantic, Falkland (Mal-
THE SOUTH AMERICAN SPECIES OF ERIGERON 5
vinas) with one species. The morphological uniqueness of
some of the species indicates that we are dealing with ancient
invasions. Especially remarkable is E. tenuifolius of the
Galapagos Islands, a small tree, with stiff lanceolate leaves,
unlike any other species found in the genus; also E. fernan-
dezianus of Juan Fernandez, another shrub or small tree,
sometimes considered a link between Erigeron and Tetra-
molopium of Hawaii (Bentham, 1873).
Finally it should be mentioned that Erigeron successfully
occupied the Patagonian territories after the last Quater-
nary glaciation. The taxonomic confusion reigning in those
species might reflect an early stage in the evolution of
the group (see under E. andicola complex).
Considering the number of species and wide distribution
of the genus Erigeron, relatively little is known concerning
its cytology. This is rather surprising, since the available
information indicates a high incidence of interesting and
anomalous situations.
Montgomery and Yang (1960) have recently reviewed
the literature and found only 35 species for which chromo-
some numbers are known. A few additional species have
been counted by Turner (1959), Raven et al. (1960),
Turner & Irwin (1960), Turner and Ellison (1960) and
Turner et al. (1961). The basic chromosome number for
the genus is x = 9, and most of the species counted are
diploid, although tetraploids are also known. A surprising
feature is that nine of the 40 odd species so far counted
are triploids, while one species, E. compositus, appears to
be a hexaploid. If the species so far counted are a random
sample of the genus, the proportion of triploids is over
20%, a remarkably high percentage of species which, unless
they possess a special mechanism, must produce a high per-
centage of sterile pollen and ovules. When investigated,
some of these species proved to be at least partially apomic-
tic, a mechanism which can successfully circumvent meiotic
sterility.
No South American material has been investigated cyto-
logically with the exception of E. maximus (Turner & Irwin,
1960), for which an inconclusive count of 40+4 was
reported, and of E. karwinskianus, which is a common weed
of Mexican origin in many tropical and temperate areas.
This has been counted from Europe by Larsen (1954),
or domi:
6 OTTO T. SOLBRIG
who reports a tetraploid count of 2n = 36 for the variety
mucronatus (sometimes considered a distinct species), and
by Turner et al. (1961) who reported a triploid plant with
27 univalents at meiosis and a diploid plant with n = 9 II,
both from Mexico. Turner et al. indicate that the triploid
plant is probably apomictic, which has also been reported
for European material. It might be interesting to know if
two fertile chromosome lines, a diploid and a tetraploid,
exist in EF. karwinskianus, the triploid being the result of
hybridization. However, in such an event a higher degree
of pairing at meiosis would be expected. Apomixis in the
European E. karwinskianus var. mucronatus has been
reported by Carano (1919) and Fagerlind (1948), although
Gustafsson (1947), feels that reéxamination is necessary.
Our knowledge of the embryology of Erigeron is relatively
satisfactory, due in great part to Harling’s (1951) careful
studies. Nevertheless, so far only some 25 species have been
investigated in detail, no South American species among
them, and certainly more work is necessary in order to con-
firm some of the interesting data obtained.
All genera of Astereae so far investigated, with the excep-
tion of Erigeron, possess a normal, monosporic type of
embryo sac (Harling 1951, 1954). Among the species of
Erigeron studied, E. philadelphicus is unique in possessing
an lusively poric, normal type of embryo sac
(McDonald, 1927). All other species have either a tetra-
sporic (the most common situation) or bisporic embryo sac;
or possess an unstable condition with monosporic, bisporic
and tetrasporic embryo sacs present in different plants of
the same species or sometimes even in the same plant
(Harling, 1951). It is of interest to note that the species
considered advanced by Cronquist ( 1947), in his revision
of the North American species of Erigeron, are exclusively
Erigeron derived from the genus Aster and embryological
data seem to back this assumption, or at least do not dis-
prove it. Aster peregrinus (considered by Cronquist as an
Erigeron on the basis of its affinities) which is related closely
a to both Erigeron and Aster has a monosporic type of embryo
__ Sae (Harling, 1951), as have all other species of Aster.
THE SOUTH AMERICAN SPECIES OF ERIGERON i
vat
ee:
eee
ES.
es
—
=
Toro aes
le
oe
epee OR
1
So
<
a
LZ
eectge
<<
a
Je
ce
=,
6. E. lanceolatus Wedd. (Lorentz 627, parw). 7. E. leptorhizon (
8. E. karwinskianus (Garaventa 941, GH). 9. E. pratensis (Hicken, S.I. 20798, DARW).
)-
10. E. ecuadoriensis (Andre 3.n., GH)
8 OTTO T. SOLBRIG
Another very interesting result pertains to the relation-
ships between Erigeron and Conyza. Conyza bonariensis
and C. canadensis, the two species of Conyza section Caeno-
tus (formerly ascribed to the genus Erigeron) so far studied,
have monosporic, normal embryo sacs, and agree in this
respect with the only other species of Conyza investigated.
If these results were to be confirmed for more species, it
would provide additional evidence for the inclusion of sec-
tion Caenotus in Conyza.
From the results so far obtained it is evident that Erig-
eron presents a unique and highly interesting type of embryo
sac development, and further study along these lines should
be very rewarding for the embryologist and the taxonomist.
Five species of Erigeron have been reported to be apomic-
tic: Erigeron annuus (Tahara 1915, 1921; McDonald 1927;
Fagerlind 1948) ; EZ. karwinskianus var. mucronatus (Carano
1919, 1920, 1921, 1924; Fagerlind 1944, 1948) ; EZ. strigosus
(Holmgren 1919; McDonald 1927) ; E. divergens (Harling
1951) ; and E. compositus (Beaman, unpublished). Of these,
only the last species has been studied in relation to the tax-
onomic implication of apomixis. No South American species
has been investigated so far, but the data obtained in this
study (see under E. andicola complex) point to the possible
presence of apomixis, at least in some degree. Embryological
study should be very rewarding in this group.
The South American species of Erigeron have never been
monographed. Over one hundred species have been de-
scribed, largely by R. A. Philippi from Chile. The present
investigation was started in 1953 as a revision of the
tine species of the genus. However, it was soon
apparent that in order to understand the group, all of the
South American species would have to be included. Even so,
the difficulties encountered have been very great, especially
in trying to understand the patterns of variation, and the
author is not sure that he has achieved a true comprehension
of the problems and their solution. Apomixis and hybrid-
ization are suspected in certain groups, but the confirmation
or disproof of these assumptions will have to await embryo-
logical and genetical studies. Lack of adequate material was
also a big handicap. However, it is felt that this monograph,
incomplete as it is in these respects, might serve the useful
purpose of pointing out avenues of research, and of indi-
THE SOUTH AMERICAN SPECIES OF ERIGERON 9
cating some of the problems to be solved. Not more than
20 per cent of the herbarium material studied in the course
of this investigation had been identified as to species,
dramatically pointing out the need of a revision of the South
American representatives of the genus.
This work was initiated while I was associated with the Museo de
Ciencias Naturales of La Plata, Argentina, at the suggestion and
under the guidance of Dr. Angel L. Cabrera, to whom I want to
express my thanks for his help and encouragement. I was fortunate in
being able to study the material of the following herbaria, to whose
directors and curators I want to express my appreciation: Arnold
Arboretum (A) and Gray Herbarium of Harvard University (GH);
British Museum, Natural History (BM) ; Instituto Botanico Darwinion,
Argentina (paRw); Chicago Natural History Museum (F); Jardim
Botanico, Rio de Janeiro (RB); Royal Botanic Gardens, Kew (K);
Instituto Miguel Lillo, Tucum4n (LIL); Museo Botanico, Cérdoba
(corp); Museo Javier Prado, Lima (UsM); Museo de Ciencias Nat-
urales, La Plata (LP); Museo Nacional, Santiago (sco); Museo
Nacional, Rio de Janeiro (Rk); Missouri Botanical Garden (Mo); New
York Botanical Garden (Ny); U. S. National Herbarium (us); and
Botanisches Museum der Universitat, Wien (wU).
ERIGERON L. Sp. Pl. : 863. 1753"
Annual, biennial or perennial herbs, subshrubs or shrubs,
rarely small trees up to 3 m. tall. Leaves alternate or rosu-
late, entire or lobed, rarely all basal or all cauline, the basal
rosette leaves often quite different from the cauline ones.
Heads borne on peduncles, solitary or in groups of less
than ten, rarely in large numbers. Involucre hemispheric
or campanulate; involucral bracts narrow, [varying from]
herbaceous and subequal [to scarcely herbaceous] and imbri-
cate; [loss of herbaceousness either uniform throughout or
more prominent towards the tip] ; receptacle flat or slightly
convex, naked or with very short projections in section
Leptostelma; ray flowers ligulate, pistillate, usually in one
series, ligules narrow of variable length but always larger
than the tubular flowers [or rarely absent], generally white,
pink, [blue] or yellow; disk flowers yellow, tubular, numer-
ous, [some species with eligulate pistillate flowers between
the hermaphrodite flowers and the ligulate pistillate
flowers], tube with five triangular lobes; style appendages
varying from lanceolate and acute to broadly triangular
Generic synonyms, now rarely used, are omitted. Parts of the deseription in brackets
do not apply to South American material.
10 OTTO T. SOLBRIG
and obtuse, [rarely obsolete]; pappus of a few to rather
numerous capillary and often very fragile bristles, [com-
monly but not always with at least a few outer short setae
or squamellae, or rarely the pistillate flowers without bristles
and bearing only short squamellae], in one or two series;
achenes terete, often compressed, commonly 2-nerved [some-
i as much as 14-nerved].
KEY TO THE SPECIES
A. Large, non-woody herbs with hollow stems; plants 30 em. or more
an Hes
B. Ligules 10-15 mm. long; inflorescences open; heads long-pedun-
c. sane at least the basal ones, petiolate, 1-10 cm. wide; plants
0.7-4 m. high.
C. Leaves all sessile, 1-2 em. wide; plants ap > new
. Ligules 25 mm. long; inflorescences tight, rong peseile or short-
3. E. tweediei.
A. Plants woody, ne least at the base, with solid stems; if herbaceous,
then usually less than 30 em. high.
D. Plants pulvinate; leaves rosulat serereesereereeee 19. EB. rosulatum.
D. Plants not pulvinate, with mlbeehat aes if pulvinate, then
never with rosulate leaves,
E. Small woody tree, 1.50 m. or more in heiphts ce ie bees
15. E. tenuifolius.
E. eo subshrubs or shrubs not more than 1 m. tall.
- Basal rosette not present.
= Annual herb; leaves pubescent, lobed. ...cscssssssssssssssssssssseesseese
17. E. leptorhizon.
G. Gheube: subshrubs or perennial herbs with at least a
woody base.
H. Leaves, at least some, divided or lobed.
I. Ligules less than 3 mm. long. ...... 13. E. ecuadoriensis.
as Ligules more than 5 mm. lon;
+ Leaves glabrous or with occasional hairs; ligules 5-7
mm. long.
J. — always pubescent; ligules 2 mm. long. ......
cee she pe meen
many heads, leafy throughout. ... camposportoi.
K. Plants less than 50 cm. in pate z gr memes a
panicle of few heads, or a single-headed scape, naked
or with very few leaves.
L. Ligules less than 3 mm. long, in 2-3 series.
M. Leaves 10-15. mm. wide, margins —_. toothed
or I srevevctaoveureconsueveveseneseseceesae incaicus.
THE SOUTH AMERICAN SPECIES OF ERIGERON 11
M. Leaves 1-4 mm. wide, margins entire. .............
12. E. pazensis.
L. Ligules more than 5 mm. long, in one series.
N. Prostrate shrubs, 5-25 cm. high; heads solitary
at the end of the branches. ..
8. E. oth
N. Erect shrubs, 30-70 cm. high; heads borne in
ener at the end of the branches or occasion-
ally solitary.
OQ. Involucre 10-18 mm. wide, plants densely
ibe: 1. EB, laxuri
pubescent. ........:c0 ou rians.
O. Involucre 5-12 mm. de pienin glabrous or
— nr apssouereocsd: 6. E. fasciculatus.
F. Basal rosette
P. Leaves at least ae em. long.
P. Leaves less than 10 cm. lon;
Q. Leaves spathulate, at least in part. ...... 18. E. rupicola.
Q. Leaves not spathulate, ac
R. Leaves stiff, often Laser
S. Leaf margin entire. ........sc-ccese 20. E. lanceolatus.
S. Leaf margin toothed or serrate.
T. Leaves and achenes pubescent, er 10-30 em.
high. . EB. ingae.
T. Leaves and achenes glabrous, joa 25-35 cm.
high. 11. E. luteoviridis.
R. Leaves neither stiff nor heavily glutinous.
21. E. pratensis.
4, E. meyeri.
long, 5-10 mm. wide, obovate. ......
V. Leaves without a clearly marked petiole, more
Se Jess than 5 mm.
wide and not obovate. .... 23. E. andicola complex.
1. Erigeron maximus (D. Don) DC., Prodr. 5: 284. 1836
(Fig. 11)
Leptostelma maximum D. Don, in Sweet, Brit. Flow. Gard. II, 1: 38.
1831; D. Don, Sweet, Hort. Brit. edit. 2: 299, 1830, nomen nudum ;
Aster maximum Less., Syn. 182. 1832.
Erigeron suleatum DC., Prodr. 5: 284. 1836; based on Lund “in
+P
Eri econ alpestre Gardner, Journ. Bot. tomion, 4: 123. 1845, based
on Gardner nites “moist places on the Organ Mts. . . . 6000’” (Holotype
geron maximus
62): 28. 1882; based on Gardner 507 “In marshes, common in the
Organ Mts., . . . 3000’” (Holotype BM!, Isotype BM!, GH!, K!)
Erigeron seaberrimum Gardner, Journ. Bot. Eondon: 7: 80. 1848,
based on Gardner 4923, “in marshy campos near Villa do Principe,
12 OTTO T. SOLBRIG
Province of Minas Gerais, Aug. 1840” (Holotype BM! Isotype K!).
Erigeron schuchtii var. minor Baker, in Martius, Fl. Bras. 6(3) : 28.
1882, based on “prope Villa do Principe, Gardner 4923” (BM!)
Erigeron catarinensis Cabrera, Arq. Jard. Bot. Rio de Janeiro 15:
75. 1957, based on Reitz 2341 “Santa Catarina, Campo dos Padres, Bom
Retiro, 2000 m.s.m, 15-II-1948” (LP!)
Giant herb, (0.5)0.7-4 m. tall. Root fibrous, slightly woody, some-
times tuber-like. Stems herbaceous, slightly woody at the base, espe-
cially in tall plants, 0.5-5 cm. in diameter, sulcate-striate, glabrous or
slightly to heavily pubescent, often scabrous, hollow, green, green-gray
or brown, leafy throughout and branching towards the top. Leaves
large, lanceolate, oblong-lanceolate to oval, 5-90 cm. long and 1-10
em. broad, margins entire, toothed or more commonly variously and
irregularly lobed or serrate, acuminate, petiolate, sessile or more often
clasping (especially in smaller and upper leaves), glabrous or pubes-
cent, especially along the veins, scabrous, midvein protruding on lower
‘ace, petiole, when present, up to 10 em. long, often somewhat
winged. Heads numerous, cymosely arranged at the end of the branch-
es, peduncles 2-15 cm. long, glabrous, or more often pubescent. Involu-
eres large, 10-20 mm. wide and 5-15 mm. high; involucral bracts
triangular-lanceolate, 5-10 mm, long, arranged in one loose series,
borders entire or somewhat toothed towards the apex, glabrous or
pubescent, often resinous-punctate, center portion when dry usually
of darker color than the borders; ligulate flowers numerous, in two
series, white, showy, tube 1-3 mm. long, ligule 10-15 mm. long, 1-2 mm.
wide; tubular flowers numerous, yellow, 4-6 mm. long; pappus copious,
white, bristles ca. 5 mm. long; achenes terete, 1-2 mm. long.
TYPE: “A native of Mexico, a plant which was obtained by Mr.
(not seen). The reference to Mexico is incorrect, and the specimen
may not be now in existence.
COMMON NAME: “Margarida da banhado” (Brazil).
SELECTED SPECIMENS: Brazil. Minas Gerais, Ouro Preto, Saramemba,
Macedo e Barroso 2793 (Mo, NY) ; Distrito Carangola, Summit of Serra
da Gramma, 1700 m., Mexia 4273 (Mo, NY, us); Vicosa, Agricultural
College lands, 670 m., Mexia 4341 (Mo, NY); Bello Horizonte, Serra do
Jaguaril, Barreto 2937 (Mo); Vicosa, A. Chase 9436 (mo); Santa
Catarina, Campo Alegre, Smith & Klein 10527 (yy, Us). Paraguay.
In viciniis Caaguazt, Hassler 8950 (BM, DARW, GH, F, LIL, MO, RB, US).
This is a very common species of south and central Brazil,
extending from the state of Minas Gerais along the coastal
mountain chain to Santa Catarina and Rio Grande do Sul.
One collection is from Caaguazi in Paraguay (fig. 12).
Erigeron maximus is most frequent in low marshy areas
itself to disturbed conditions where sufficient moisture is
—— Specimens from the higher altitudes are usually
THE SOUTH AMERICAN SPECIES OF ERIGERON 13
Fic. 11. Erigeron maximus, general aspect, X 4% (Mexia 4351, GH). Drawing by
Ruth Hsy.
14 OTTO T. SOLBRIG
smaller and more pubescent. The leaves vary greatly in
size and shape. A true basal rosette, such as that found in
E. tweediei, is usually absent, although many plants, espe-
cially large ones, have larger, petiolate leaves toward the
base of the plant (which can be up to 90 cm. long), while
the upper leaves tend to be smaller and basally clasping.
Equally variable is the foliar surface, which in a few plants
is smooth and glabrous, in others scabrous-pubescent and in
still others (especially those from the State of Minas Gerais)
remarkably scabrous, the scabrousness being produced by
a large number of minute crystalline projections. A certain
amount of variation is also found within the same plant:
young leaves as well as first leaves are much less scabrous
than old and later leaves. Although most plants have sessile
leaves, some specimens from Santa Catarina (described as
E. catarinensis by Cabrera) show basal leaves which are
petiolate and somewhat sagittate or auriculate at the base.
Some intermediate conditions are also found.
A collection from the state of Espirito Santo should also
be mentioned. This material has much smaller and narrower
leaves, the plants are smaller and very reminiscent of E.
tu is from Argentina. It is the only collection made
in that state.
Although some of the variations mentioned, especially
those which seem to have a geographical or ecological asso-
ciation might deserve subspecific recognition, it is felt that
in view of the tremendous variation present (which becomes
obvious when one studies the plants in the field) and the
constancy of the floral ch , NO segreg: on the
basis of vegetative ch ters is ted with our present
knowledge.
2. Erigeron tucumanensis Cabrera, Notas Mus.
La Plata Bot. 19: 196. 1959
several from the base, 2-3 mm. in diameter, hollow, grooved, slightly
pu unl
leaves short-lived, not different from the cauline; cauline leaves sessile,
» elongate-lanceolate, 10-15 em. long, 1-2 em. wide, margins
ace pubescent to slightly scabrous. Capitula
borne singly at the end of long pedicels, 2-5 at the end of each shoot.
Tnvolucre hemispheric 0.9-1.2 em. wide, 0.6-0.8 em. high; involucral
apparently in one row, triangular-elongate, punctate or short-
THE SOUTH AMERICAN SPECIES OF ERIGERON 15
Fic. 12. Distribution of Erigeron mazimus, manensis, E. fasciculatus and
. ecuadoriensis. Goode base map, ear Ly fie Ucmeeie at Chase
16 OTTO T. SOLBRIG
pubescent, with dark-brown or slightly reddish midrib portion; ligulate
flowers numerous, in one series, yellow, tube-portion 2-3 mm. long,
ligule 6-8 mm. long; tubular flowers numerous, yellow, 3-4 mm. long;
achenes terete, red-ochraceous in color, 1-2 mm. long, glabrous.
TyPE: Argentina, Venturi 4745 “Tucuman, Dept. Chicligasta, Las
Pavas, 2500 i Spa LP!, Isotypes GH!, LIL!, Us!)
MATERIAL IED: Argentina. Catamarca, dept. Andalgala (sic;
Tucuman, faves eeuecy Rio Cochuna, Jorgensen 1514 (GH, MO,
us).
This rare species from Tucuman in Argentina (fig. 12)
is closely related to E. maximus. It differs by its smaller
eads, borne singly or in groups of not more than five, the
yellow ligules, the slender stems, the narrow leaves and the
small size. It is also completely geographically isolated from
that species.
3. Erigeron tweediei Hook. et Arn., Comp. Bot.
Mag. 2:50. 1836
(Fig. 14)
Erigeron seneciiformis Blake, Proc. Biol. Soc. Wash. 36: 51. 1923,
sei cs =e “White 1206, css Rosario, near Reyes,” 11-IV-
-alhenal green to greenish-brown in color, leafy, unbranched ©
branched only at the top. Basal leaves forming a conspicuous pad
10-30 em. long, 5-30 mm. wide, lanceolate, entire, acute, margins entire
or dentate, surface smooth, glabrous or somewhat tt pubescent, especially
along the midvein and margins; petiole elongate, 1-5 em. long, expand-
ing gradually into the lamina, 0.5-3 mm. in diameter, the dry bases
the shoot, usually rather tight; pedicels 2-50 mm. long, pubescent.
Involucres large, 10-15 mm. wide, 5-10 mm. high. Ano chae bracts
, acute, 3-5 mm. long, 1-3 mm. wide, borders entire or slight-
g-
LOCALITY: Uruguay, Tweedie 1058 “Maldonado in boggy
ground” (not seen).
: SELECTED SPECIMENS: Argentina. Co: rrientes, Hoge Batel, ee?
3101 (ws) ; Entre Rios, Concordia, Burkart 1 5 (DARW); Misiones,
San José, Bertini 2732 (F); Jujuy, Toldos bei ae Fiebrig 2378
: ae eh Bolivia. ara, lomas de Buenavista, Steinbach
6712 (Mo, BM, GH), 2684 (DaRw, ms 5293 (GH), 3166 (LiL). Brazil.
THE SOUTH AMERICAN SPECIES OF ERIGERON
Fic. 13. Erigeron tucumanensis, general aspect X 4% (Venturi 4745, UL).
18 OTTO T. SOLBRIG
Goias, Gardner 4332 (K); Minas Gerais, Caldas, Henschen 203 (US) ;
Serra da Matuca, Williams 5314 (GH); Paranda, Curytiba, Hatschbach
609 (LIL) ; Rio de Janeiro, Rio, Glaziou 11052 (K); Rio Grande do Sul,
Lagoa Vermelha, Bornmiiller 704 (GH) ; Santa Catarina, Lages, Rambo
49587 (LIL). Paraguay. Carapegua, esteros Calistro, Rojas 3297
(DARW, LIL) ; Caaguaza, Colonia Sommerfeld, Sparre & Vervorst 2260
(IL). Uruguay. Montevideo, Arechavaleta 4182 (K) ; San José, Barra
de Santa Lucia, Herter 1540 (GH, LIL).
This species occupies the largest geographical area of
all the South American species of Erigeron, growing in
swamps and wet places, in the states of Goids, Minas Gerais,
Rio de Janeiro, Parana and Rio Grande do Sul in Brazil, in
Uruguay, in eastern Argentina, in Paraguay and in the low-
lands of eastern Bolivia (fig. 20). In spite of its great range
it does not seem to be too common (being much less fre-
quent than the partly sympatric E. maximus), although
one should remember that the area is in general poorly
explored and collected. The species is quite distinct and
certain doubts as to its status in Erigeron have been raised
(see Solbrig, 1960). It is related to Erigeron maximus, from
which it can be separated easily by the short yellow ligulate
flowers and the tight inflorescences. The heads are also
smaller than in that species.
4, Erigeron meyeri Cabrera, Notas Mus. La Plata
Bot. 19: 198. 1959
(Figs. 1, 15-18)
Perennial herb from a short stout rhizome, 40-60 em. tall. Shoots
several from the base, slightly grooved, ioe to greenish brown.
Basal rosette of leaves Jax; leaves sword-shaped, 15-25 cm. long, 8-15
mm. wide, acute, eae minutely dentate-pubescent, surface
glabrous, petiole short, expanding gradually into the lamina; cauline
leaves shorter and narrower than the basal, sessile. Capitula several
shoot in an open panicle, pedicels 5-15 em, long, slightly pubescent.
Involucre 8-12 mm. wide, 5-7 mm. high; involucral bracts in 1-2
series, 0.5-1 mm. wide, 2-4 m. long, acuminate, dorsally
Sameer stiff; ligulate flowers in one series, tube 2-3 mm. ee
‘igule 3-4 mm. long, 1 mm. wide; tubular flowers numerous, 3-4 mm.
a ye Prov. Chaco, Meyer 2234 (Holotype LP!, Isotype
ca STUDIED: Argentina, Prov. Chaco, Fontana, Meyer 2125
19
AMERICAN SPECIES OF ERIGERON
THE SOUTH
Fic.
! V7
~
14. Erigeron tweediei, general aspect X 14 (Dusen 14493, GH). Drawing by
Ruth Hsu.
20 OTTO T. SOLBRIG
This species, known from only two collections from one
locality (fig. 19), is very distinct, and can be easily recog-
nized by the basal leaves. Its affinities are unclear, but it
is likely that it might be related to Erigeron tucumanensis
or Erigeron maximus, the first one in particular having
certain similarities in leaf shape and aspect of the capitula,
although it lacks a basal rosette. Its distribution is restricted
to a small area in the western part of the province of Chaco
in Argentina, where it apparently is abundant (Meyer, per-
sonal communication).
5. Erigeron camposportoi Cabrera, Arq. Jard.
Bot. Rio de Janeiro 15:75. 1957
Perennial erect herb, 70 cm. high. Stem grooved, solid, with white,
soft pith, brownish to brownish-green in color. Leaves 5-10 cm. long,
1.5-2.5 cm. wide, lanceolate, dentate, glabrous on both surfaces, short-
petiolate; upper leaves smaller, entire, sessile. Heads borne at the tip
the ches in open panicles with many heads. Involucre hemi-
spheric, 7-9 mm. wide, 4-5 mm. high; involucral bracts numerous,
linear, in one to three series, 2-4 mm. long, 0.5-1 mm. wide, glabrous
or very short pubescent, with short-ciliate margins; ligulate flowers
numerous, white, in two series, tube 2-4 mm. long, ligule 3-4 mm. long,
0.5-1 mm. wide; tubular flowers yellow, 3-4 mm. long; styles triangular
tipped; achenes dorsally compressed, turbinate, 1-2 mm. long, slightly
seni siae
estado do Rio de Janeiro, Cabrera 12294 “Serra dos
ate selva, 1800 m.s.m.” (LP!)
MATERIAL ED: Brazil, estado do Rio de Janeiro, Serra do Ita-
tiaia, 900 m. s.m., Malme 312 (rR).
A rare species known only from two collections (fig. 42), it
is very distinct from any other South American species. It
grows in an area relatively well collected, and it is strange
that it has not been found at any other time.
6. Erigeron fasciculatus Colla, Mem. Acad.
Torino 38:27. 1834
(Fig. 20)
. Erigeron. berterianus DC., Prodr. 5: 286. 1836, based on Bertero sn.
lollium [sie] prope Valparaiso Chilensium” ( isotype BM!)
suspected 4
spiculosus Hook. & Arn. var. a (var. “liguli longiori”
are Comp. Bot. Mag. 2: 49. 1836, based on Cuming 407 “Valparaiso”
_Haplopappus hispidulus DC., Prodr. 5: 348. 1836, based on “in Chile
prope Coquimbo legit el. pa 87” (Isotype GH!)
Erigeron subandinus Phil., Linnaea 28: 723. 1858, based on Gay 761
“in subandinis Sta. Barbara, prov. Concepcién, Jan. 1839” (sco!)
THE SOUTH AMERICAN SPECIES OF ERIGERON
21
Fics. 15-18. Erigeron meyeri. 15. Capitulum X 2. 16. General aspect X %4. 17. Ligu-
late flower
bular flower X 4 (All from Meyer 2125, LP).
OTTO T. SOLBRIG
‘rigeron meyeri, E. luzurians, E. pazensis and E. karwin-
base map, copyright by the University of Chicago.
Fic. 19. Distribution of E;
THE SOUTH AMERICAN SPECIES OF ERIGERON 23
Aster breviflorus Phil., Linnaea 33: rites 1864, based on Volkmann
s.n. “Prov. Coquimbo, aestate 1860/61” (sco
Aster litoralis Phil., Anal. Univ. Sg 43: 486. 1873; Erigeron
litoralis (Phil.) Skottsb., Acta Horti Gothob. 18: 158. 1950; based on
Philippi s.n. “in scopulis maritimis Guayacan, Nov. 1864” (sco!)
Subshrub, 20-70 cm. high. Root woody; stems, especially near the
from the base, or one or two, with few, sparse hairs or glabrous.
Leaves spirally arranged, borne singly on shoots, or in fascicles in the
axils of branchlets or older leaves, linear-lanceolate to spathulate,
margins entire, or slightly and irregularly serrate or dentate, grayish-
green in color, covered with short multicellular, uniseriate hairs on
both surfaces, or glabrous, hairs usually dense but varying from col-
lection to collection, leaves 5-30 mm. long, 1-5 mm. broad; petiole short,
gradually expanding into the lamina, 1-2 mm. broad, 1-5 mm. long,
pubescent or glabrous. Heads arranged in panicles at the end of the
branchlets or occasionally solitary; peduncles up to 10 cm. long,
glabrous to pubescent, covered with a few bracts or short leaves, 10
mm. long, up to 3 mm. broad, sessile, pubescent; involucre 5-12 mm.
wide, 4-8 mm. high, involucral bracts triangular-lanceolate, 2-10 ) me.
long, 05-20 m mm. wide, imbricate, arranged in two loose series, co’
with medium to long, unicellular or more commonly multicellular ony
lense, giving a white-woolly appearance, or occasionally
glabrous; ligulate corolla white or yellow, 10-15 mm. long, 0.5-1.0 mm.
—— glabrous, tips of ligules sometimes slightly notched; tubular
rolla 3-5 mm. long, whitish or yellowish; style of tubular flower with
ian tips covered with collecting hairs, pappus white-yellowish
in herbarium material, 2.5-4.5 mm. long; achenes terete, pubescent, 1-2
mm. long.
TYPE: Bertero “in Chili locis sylvaticis collium Valparaiso” (not
seen).
MATERIAL STUDIED: Chile, Prov. Coquimbo, Coquimbo, Gaudichaud
87 (GH), Harvey sn. (paRw, GH), Ball sn. (GH, K), Coppinger s.n.
(kK), Phillippi sn. (sGo), Geisse s.n. (SGO), Hastings 596 (US), Hicken
8689 (DARW), Joseph 5422 (US), Jaffuel 1207 (GH), Montero 1890
(GH), Werdermann 109 (BM, DARW, F, GH, LIL, MO); La Serena, yan
5483 (us), Dept. Ovalle, coders Jiles P. 1948 (LP), Tulahwen, Geisse
(sco); Los Vilos s.c. (Sco); p ov. Aconcagua, Zapallar, Betin (F);
playa Papudo, Looser 2470 i); Pree Valparaiso, Valparaiso, que-
brada del Licumo, Looser 3761 (LP), Quintero, Albert s.n. opened
Valparaiso, Cuming 407 fae 406 (BM), Wilkes sn. Joseph 1.
(us), Phillippi sm. (sco); collium loco dicto Bertero nt
(BM, GH); Cerro Campana, Morrison & Wagenknecht 17153 (DARW,
GH, K), Looser 586 (GH), Garaventa 1875 (GH).
Erigeron fasciculatus grows along the Chilean coast from
Valparaiso to Coquimbo, on hills near the coast, or on the
beach (fig. .
Since we are dealing with a shrubby species, the speci-
mens available are still less satisfactory than the average
24
OTTO T. SOLBRIG
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Fic. 20. Erigeron fasciculatus, general aspect X 44 (Werdermann 109, GH).
THE SOUTH AMERICAN SPECIES OF ERIGERON 25
herbarium specimen. It is evident that at least in some cases
they are misrepresentative of the species. The basal leaves
(which are wider) are often missing, giving an entirely
different aspect to the specimen. Also data as to habit, size
of plant, color of flowers, etc. are almost entirely missing.
The collections from Cerro Campana near Valparaiso are
quite interesting. Although the species has in general
slightly to heavily pubescent leaves and stems, the specimens
from Cerro Campana are almost completely glabrous. In
addition the leaves are spathulate, at least the basal ones,
and not in fascicles. Nevertheless, in view of the scarcity
of material, it is thought best not to ascribe taxonomic status
to this material for the present.
Skottsberg (1950) has expressed certain doubts that F.
berterianus ( which . Signe sige with E. litoralis)
and E. twithstanding the
fact that De Candolle i in