Rhodora
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Conducted and published for the Club, by MERRITT LYNDON FERNALD, Editor-in-Chief
CHARLES ALFRED WEATHERBY LUDLOW GRISCOM Associate Editors STUART KIMBALL HARRIS
VOLUME 43
1941
The New England Botanical Club, Ine. 8 and 10 West King St., Lancaster, Pa. Room 1001. 53 State St., Boston, Mass.
JAN
18 1944
Hodova
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Conducted and published for the Club, by MERRITT LYNDON FERNALD, Editor-in-Chief
CHARLES ALFRED WEATHERBY
LUDLOW GRISCOM \ Associate Editors
STUART KIMBALL HARRIS f Vol. 43. January, 1941. No. 505. CONTENTS: Revision of the Genus Pteridium. ZA. M. Tryon, Jr. ........... 1 Walker Prize in Economic Botany (Notice) ................... 31 The Red Azalea of the Cumberlands. E. Lucy Braun .......... 31 Geranium nemorale, var. Bicknellii. M. L. Fernald ............ 35 Lachnanthes in Nova Scotia. C. A. Weatherby ................ 36
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Rhodora Plate 650
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Segments and their characters in PrERIDIUM.
Rhodora
JOURNAL OF
THE NEW ENGLAND BOTANICAL CLUB
Vol. 43. January, 1941. No. 505.
CONTRIBUTIONS FROM THE GRAY HERBARIUM OF HARVARD UNIVERSITY—NO. CXXXIV
A REVISION OF THE GENUS PTERIDIUM R. M. TRYON, JR. (Plates 650—653) INTRODUCTION
THE only species of the monotypic genus Pteridium, P. aqui- linum, is one of the most familiar plants in the world. Its distri- bution is world-wide in temperate and tropical regions and it is common to the point of becoming a weed in most of the regions in which it grows. Many authors complain of its weedy char- acter in the British Isles, northwestern United States and New Zealand. In these regions and probably in others it invades fields and most especially pastures and it is probably the most obnoxious native weed with which the farmers have to deal. Apparently, it immediately assumes the role of a weed as soon as man tries to turn to his own uses any land that it finds suit- able for its growth. However, the weedy character seems to vary with the variety, or region, for var. latiusculum and var. pseudo- caudatum in eastern North America only rarely, if ever, become a serious menace to farmers,
Among the many species of ferns that spread by means of rhizomes or runners, the Bracken is particularly well adapted to the role of a weed. In favorable soil conditions there is a widely ereeping, deep, main rhizome from which lateral, ascending branches arise. "These branches usually rebranch and produce
2 Rhodora [JANUARY
the fronds fairly near the surface. A dormant bud is produced at the base of each frond so that each year the plant has a po- tential second set of fronds. If, through burning or plowing, the lateral branches are destroyed, the main deep-seated rhizome can continue the life of the plant.
A few excerpts from a recent article on the Bracken in Britain! will illustrate its economic importance as a weed and some of the methods taken to control it. “Bracken ranks as the most plentiful and widespread of British ferns. It is generally dis- tributed in most rough grazings and in many old permanent pastures in almost every part of Britain. It is plentiful in most open woodlands and on the sites of formerly wooded areas. . . . Many of the older writers have dealt with the loss of grazing due to Bracken. . . . ‘I have seen the Roots of it in some Grounds, eight Foot deep. The best cure is often mowing of it while in grass. If you plow it up, plentiful dunging of it and Ashes are very good: but the certainest cure for it is Urine.’ J. Mortimer, The Whole Art of Husbandry, 2nd Ed., 1708." At the present time, the authors recommend cutting the fronds when they are fully mature and following this with heavy grazing; and also the use of phosphate and lime fertilizers to build up the ground and make it unsuitable for the growth of the Bracken. Sodium chlorate and sulphuric acid are recommended as killing sprays. “In closing this paper, however, the writers desire very definitely to record their view that the bracken menace has grown so rapidly, and has now attained such proportions, that it is of real national importance: at the International Grassland Congress of 1937 one speaker referred to it as ‘the ghost stalking silently at our side, which nobody dares to discuss.’ . . . A million pounds sterling devoted to the reduction of the area infested by bracken would be well spent."
Many species of fungi are parasitic upon Pteridium aquilinum” and attempts have been made to use them in its control. Fusarium sp. and Corticium anceps (Bres. and Syd.) Gregor have been tried? but due to the faet that the host seems to be
1Long and Fenton, The Story of the Bracken Fern. Journ. Royal Ag. Soc. Eng. 99: 15-36 (1938).
2S8ee Seymour, Host Ind. Fungi N. Am. (1929) and Faull, Gen. Urediniopsis. Contrib. Arn. Arb. 11 (1938).
з See Gregor in Verdoorn, Man. Pterid. 146 (1938).
19411 Tryon, Jr.—Revision of the Genus Pteridium 3
more durable under varying environmental conditions than the fungus, they have not met with success.
To compensate for its economic liabilities, the Bracken has made itself available for many uses, though at the present time none are of real importance. The fronds have been used for stock-food, when mixed with grass; as silage; for packing fruits; in the place of hops in brewing; medicinally; in glass making and as litter for both man and stock. The fronds and rhizomes have been used in thatching and the green fronds burned as a source of potash. Nakait says that in Java the fronds are laid on racks over the Cinchona beds to protect the young plants from the sun and that: “This shelter is not liable to rotten and decays harmoniously with the growth of the Cinchona. When Cinchona reaches to the height of the roof, it want the shelter no more, and the fronds are so good and perfect as they then become bones only, dropping all of the leaves.” The rhizomes have been used in brewing a beer and also as swine-food. It is reported that the swine prefer the rhizome plus soil. The hairs from the base of the petiole of var. caudatum are used for stuffing pillows and the fronds are used as packing in the tops of char- coal bags. A solution of the boiled fronds has been used effec- tively as a rose spray. Bracken is often accused of being poisonous to stock but apparently this is true only when it is eaten exclusively and in large amounts. The rhizomes, which contain a considerable amount of starch, have been used as food, at one time or another, in Switzerland, France, Canary Islands, western North America, Society Islands, New Zealand, Tasmania, New South Wales and China. It is said that formerly it was an important food-item of the Maoris of New Zealand. One doctor is reported as using the starch in place of arrowroot. However, the ordinary rhizome would undoubtedly be poor fare and probably was usually eaten only to stave off actual starva- tion. Forster? says that: “Radices ab incolis inopibus et famelicis exsuguntur, insipidae, parum nutrientes lignoso-fibrosae.” The eroziers have been used as "asparagus" in Japan and western North America. A number of botanists have reported eating the broiled eroziers and say the flavor is quite agreeable. I can
t Nakai, Critical Notes on Japanese Ferns. Bot. Mag. Tokyo 39: 110 (1925). 5 Forster, Pl. Escul. 74 (1786).
4 Rhodora [JANUARY
attest this myself but must add that the dense pubescence is rather troublesome.
In the Canary Islands, Orobanche trichocalyx (Webb & Berth.) G. Beck and O. Schultzii Mut. are parasitic upon Pteridium aquilinum.
REPRODUCTION
Vegetative propagation by means of the creeping rhizome is the predominate method of reproduction of the Bracken, al- though in favorable localities ordinary sexual reproduction is quite as effective. Although the Bracken habitually grows in rather dry places these are apparently not suitable for its normal reproduction, the prothallial stage developing only in relatively damp situations. Long and Fenton® say that in the eastern side of Britain, the prothallial stage is rare or unknown, while in the western side, where there is considerably more rainfall, young plants are not uncommon. In eastern North America young plants are only rarely seen. The young stages are not common as herbarium specimens. It is an indication of their relative rarity that at least two species and one variety have been described on the basis of young plants, although their authors no doubt little suspected they were redescribing the familiar Bracken. If, in the past, the Bracken has reproduced largely by vegetative means, this might well explain its present vigor and the lack of specific segregation within the genus. Judging biological age by the number of generations, it would still be a relatively young species.
NECTARIES
Nectaries are present at the base of the lower pinnae. These were apparently first noticed by Charles and Francis Darwin‘ but they have been mentioned by few authors? since then. I had an opportunity to observe them on var. latiusculum in the Har- sard University Botanic Garden in the spring of 1940. On the sixth of May, the basal pinnae were just starting to unroll and * Op. cit.
т Francis Darwin, Journ. Linn. Soc. 15: 407, Tab. 6, Fig. 6 (1876). 5'The only other original observation I have seen is that of Victorin, Les Filicinées du Québec. Contrib. Lab. Bot. Univ. Montréal No. 2, 71 (1923), who says: “Au printemps
les nectaires basilaires des segments sont presque toujours visités par une grosse fourmi noire."
19411 Tryon, Jr.—Revision of the Genus Pteridium 5
the nectaries were clearly visible at the base of the pinnae as dark brown, smooth, somewhat swollen areas on the dorsal side of the then densely pubescent rachis. As the fronds continued to unroll proportionately smaller nectaries could be seen at the base of the second and third pair of pinnae. These nectaries were functional in that they exuded a liquid in the form of droplets from small pores. The nectaries at the base of the second pair of pinnae were not as active as those below and those at the base of the third pair of pinnae seemed to be entirely inactive. Large red and black ants were attracted by the exuda- tion and observed in considerable numbers feeding upon it. Al- though an analysis was not made, the fact that ants feed upon the liquid undoubtedly indicates that it contains sugar. A free- hand section of the nectary area showed parenchymatous tissue but there was no evidence of glands. For these reasons I think it is likely that the exudation is neither a guttation of nearly- solute-free water nor a modified secretion but rather an excretion of the stipe-sap. The nectaries are active in the sun and also in the shade. They were quite active on a cloudy humid day. They were observed to remain active until the fronds were almost completely unrolled and may be active for a longer period of time. By midsummer they have become quite insignificant and entirely inactive. I believe this phenomenon is physiological, perhaps related to root pressure.
PRIMITIVE AND ADVANCED CHARACTERS
Phylogenetically considered, Pteridium has a curious combina- tion of relatively primitive external characters and relatively ad- vanced internal characters. In the Pteridoideae it is considered to be a primitive genus? on the basis of the following characters: presence of a more or less vestigial inner indusium; initial basi- petal succession of sporangia; hairs, but no scales, present on the rhizome; equal dichotomy of the axis in its early develop- ment; and open venation. On the other hand, the highly de- veloped vascular structure of the stem and petiole and the presence of true vessels!’ indicate an advanced condition. The highly developed internal structure, however, is undoubtedly, at least in part, related to the relatively large size of the plant.
9 See Bower, The Ferns. 3: 41—45 (1928). 10 Bliss, The Tracheal Elements in the Ferns. Am. Journ. Bot. 26: 620-624 (1939).
6 Rhodora [ JANUARY
HisTORICAL ACCOUNT
Most of the taxonomie work on the Bracken has been done by authors working in a limited area or on a limited number of forms. "There have been only a few comprehensive treatments. This has resulted in a general tendency to treat the various groups as species and to recognize, in one rank or another, trivial varia- tions, a condition which naturally has led to a multiplicity of names. Not only has it been unnecessary to describe any new varieties in the present treatment, but in 1839 Agardh likewise found all of the groups he recognized already named. Most of the critical work has been done by local authors.
As in the ease of so many groups, Linnaeus, Sp. Pl. 2 (1753), laid the foundations for the present treatment. He recognized, in the genus Pteris, two species, P. aquilina and P. caudata, which represent the ssp. typicum and caudatum of this treatment. Willdenow, Sp. Pl. 5 (1810) recognized, besides Pteris aquilina and P. caudata, some additional species described since 1753: P. capensis, P. esculenta and P. lanuginosa. Under P. aquilina he recognized, in var. 8, a combination of var. latiusculum and var. pseudocaudatum of this treatment, but most later authors did not take up this segregate.
Agardh, Rec. Pterid. (1839) was the first author really to study material from most of the regions of the world and his treatment is in elose agreement with the present one although he regarded most of the groups as species. He followed Linnaeus in treating the Brackens under the genus Pteris, but set them apart from the other species as the section Ornithopteris. He recognized P. esculenta, P. arachnoidea, P. caudata and P. decomposita and applied the names in essentially the same sense as in the present treatment. In P. semihastata and P. recurvata he recognized, respectively, var. yarrabense and var. Wightianum. Under P. aquilina he mentioned, but did not describe, var. pseudocaudatum. In the main his treatment differs from the present one only in the reeognition of P. lanuginosa and in uniting var. latiusculum with P. aquilina. Later authors would have done well to follow Agardh's treatment more closely.
Hooker, Sp. Fil. 2 (1858) presented the next, and really the last, comprehensive treatment of the genus. He followed Agardh in treating the species under section Ornithopteris of Pteris, but
19411 Tryon, Jr.—Revision of the Genus Pteridium 7
except for P. coriifolia and P. psittacina, which he says he did not understand, having seen no specimens, he reduced all names under P. aquilina and recognized only varieties. The present treatment agrees with his in this respect, but his varietal lines were rather poorly drawn. АП varieties in the present ssp. typicum he treats under var. glabra, fronds glabrous or nearly so beneath, and var. lanuginosa, fronds quite pubescent beneath. In the present ssp. caudatum he recognized var. caudata, apply- ing it in the present sense, and var. esculenta, including the present var. esculentum, var. arachnoideum and var. yarrabense. This treatment was long followed and led especially to a confusion of var. esculentum of Polynesia, Australia and New Zealand and var. arachnoideum of South America.
Diels, Nat. Pfl. 1* (1899) and Christensen, Ind. Fil. (1905) recognized the single species Pteridium aquilinum. Nakai, op. cit. (1925) in a rather rambling and non-critical discussion, made several new combinations under Pteridium, reviewed much of the literature and discussed the status of various species and varieties.
Under various names, Seopoli, Fl. Carn. (1760), Gleditsch, Syst. Pl. (1764), Newman, Phytol. 2 (1845) and John Smith, Hist. Fil. (1875) separated the present genus Pteridium from Pteris, but it was not until Kuhn in v. d. Decken, Reis. Ost.-Af. 3° (1879) took up and defined Pteridium that the Braekens were widely treated as a distinct genus.
‘TERMINOLOGY
Certain terms have been used in the descriptions in a strictly limited sense and certain unusual terms and characters have been used. These are explained below:
The vernation of the frond equal, all of the pinnae becoming unrolled at essentially the same time; subgleichenioid, the tips of the upper pinnae still inrolled while the lower pinnae are com- pletely unrolled; or gleichenioid, the basal pinnae being com- pletely unrolled while the third or fourth pair of pinnae are completely inrolled.
A costa is the midnerve of a pinna.
A costule is the midnerve of any segment of lesser order than a pinna, with the exception of the midnerve of the ultimate seg- ments, which is called the m?dnerve,
8 Rhodora [JANUARY
In some varieties there are lunate or sublunate, entire lobes along part of the rachis, costae and costules, between the divisions of the blade. These are called free lobes. Prate 650, Fic. 10.
The ultimate segments are adnate, broadest, but not dilated, at the base (PL. 650, ric. 7) ; decurrent, dilated at the base toward the base of the midnerve (PL. 650, FIG. 16); surcurrent, dilated at the base toward the tip of the midnerve (Pr. 650, FIG. 3) ; or narrowed at the base (PL. 650, ric. 6).
The upper surface of the ultimate segments includes the mid- nerve but excludes the margin.
The margin, although morphologically on the upper surface of the segment, is described separately. It is the chlorophyll-bear- ing tissue closest to the base of the outer indusium.
Peculiar, irregular, wing-like epidermal outgrowths occur on the veins and midnerve on the lower surface of the ultimate seg- ment in some varieties. These are called membranous wings PLATE 650, Fic. 14.
Some varieties have a farinaceous appearance on the lower surface of the ultimate segments. This is a minute epidermal outgrowth, occurring between the veins, which, under high magni- fication, resembles a sparsely branched coral fungus. It may be an excretion but it is not affected by alcohol or dilute acid. PLATE 650, ric. 13.
The term indusium is applied to the outer, functional indusium, the modified margin of the segment. The outer indusium is pres- ent in both fertile and sterile fronds and throughout partially fertile fronds. If it covers sporangia it is spoken of as the fertile indusium, if not, as the sterile indusium. The inner in- dusium is non-functional and usually nearly obsolete. When the inner indusium is meant it is called such.
The costules and lower side of the midnerves in ssp. caudatum are described as pubescent with white, dark or bicolorous hairs. The white hairs are long, thin, terete, whitish, multicellular hairs with oblique cross-walls. These are essentially of the same type that occur on the tissue on the lower surface of the segments. The dark hairs are rather large, stout, dark or reddish brown, usually terete, multicellular hairs with straight cross-walls, which are similar to those that occur on the rhizome. As these mature they often become moniliform, with whitish walls, only the septae
19411 Tryon, Jr.—Revision of the Genus Pteridium 9
remaining brown. These are called bicolorous hairs. When one of these dark or bicolorous hairs breaks off, it often leaves a swollen base that is dark reddish brown and has the appearance of a sessile gland.
SYSTEMATIC TREATMENT
Although Pteridium is world-wide in distribution and the vari- ous populations show very strong phytogeographic affinities, and although it is undoubtedly a rather old genus, three lines of evi- dence lead me to believe that specifie segregation has not taken place. First, in all areas where two varieties of the same sub- species overlap in range intermediate specimens are found (with the exception of var. africanum in ssp. typicum). Such inter- mediates are not common but they are of such frequeney that, assuming indiscriminate rather than critical collecting, they must form a percentage of the Pteridium population of the area that cannot be overlooked. Second, in areas occupied exclusively by one variety, individuals, perhaps representing local populations, occur that have one or more characters of some other variety. That is, characters of one variety crop up occasionally in other varieties. In some cases a specimen will resemble another variety so closely that it would certainly be identified with it except by using the geographic "character." In other words, the critical characters of the varieties are not thoroughly stable. "Third, the critical characters show little constancy within the group as a whole. That is, ciliation of the indusium in var. Fee?, or non- ciliation in var. africanum, are critical characters but both con- ditions are commonly found in var. caudatum; pubescence on the undersurface of the blade in var. pubescens and the lack of it in var. latiusculum are critical characters, but var. Wightianum, while most commonly pubescent, is not infrequently nearly glabrous; pubescence on the margin of the segments in var. Feet and the lack of it in var. typicum are important characters, but var. Wightianum frequently exhibits both conditions; and the free lobes in var. arachnoideum and their absence in var. cauda- tum are diagnostie characters but both conditions are found in var. yarrabense. It seems to me that, when the critical charac- ters of a group show such inconstancy, they are materially weak- ened for use in specific segregation.
10 Rhodora [JANUARY
The treatment of each variety follows a fixed order somewhat different from the conventional one. First, the accepted name and its synonymy and then any necessary discussion of the nomenclature. Second, the description of the variety followed by a discussion of taxonomic problems and a comparison with closely related varieties. Third, a short phytogeographie discus- sion, if desirable, a statement of habitat preference and a general statement of range. Last, a citation of specimens. In this ar- rangement, the critical discussion of nomenclature and taxonomic problems follows the synonymy and description, respectively ; and the usually long citation of specimens comes at the end.
This study is based on the 2,500 sheets of Pteridiwm in the Gray Herbarium, the Herbarium of the New York Botanical Garden, the United States National Herbarium and the Her- barium of the Field Museum of Natural History.
DESCRIPTIONS
The critical characters in the descriptions have been italicised. The descriptions are based on mature specimens and do not take into account juvenile forms. The cutting of the blade is the same in the juvenile state of all varieties. The margins of the segments are flat, not revolute, and there is no sterile indusium. None of the various epidermal outgrowths are present except pubescence, and that is only weakly developed and not distinctive.
Maps
The maps of the varieties have been compiled primarily from the cited specimens. These localities have been augmented by localities taken from other specimens examined and in a few cases by localities taken from the literature. These are used only when a specific locality was mentioned and where there was no doubt as to the identity of the variety. Localities based on specimens are indicated by a dot and those based on a printed record by a cross. The map of the species includes all of the localities on the varietal maps and in addition several localities taken from the literature where the identity of the variety was in doubt. On this map no distinction is made between localities based on specimens and those based on the literature. No at- tempt has been made to make the maps absolutely complete.
19411 Tryon, Jr.—Revision of the Genus Pteridium 11
The series of specimens studied has in most cases been sufficient to plot the general range of each variety. The maps used have been selected from Goode’s Series of Base Maps, Henry M. Lep- pard, Editor, Copyright by the University of Chicago, published by the University of Chicago Press.
CITATIONS OF SPECIMENS
The citations of specimens have been limited, when necessary, to two or three from each minor political division, i. e. state, province or department. The most widely distributed collections are cited in preference to the more locally distributed ones. Full citations are given in the discussions only for specimens that are not listed in the formal citations. Herbarium-sheet numbers, such as, Herb..Field Mus. no. 47805, are used in cases where the data on the label is insufficient to identify the specimen accu- rately. The usual abbreviations for the herbaria are used: F, for the Herbarium of the Field Museum of Natural History, Chicago, Ш.; G, for the Gray Herbarium, Harvard University, Cambridge, Mass.; NY, for the Herbarium of the New York Botanical Gar- den, Bronx Park, N. Y.; and US, for the United States National Herbarium, Smithsonian Institution, Washington, D. C. In the few cases where other herbaria are cited, abbreviations are not used.
ACKNOWLEDGEMENTS
I wish to thank Dr. P. C. Standley, Dr. H. A. Gleason and Dr. W. R. Maxon for the generous loan of material and for privileges extended at the herbaria under their care; and es- pecially to express appreciation to Mr. C. A. Weatherby, without whose constant help and encouragement this study could scarcely have been completed.
SYSTEMATIC TREATMENT
PTERIDIUM Gled. ex. Scop. Fl. Carn. Ed. 1, 169 (1760), nomen abortivum(?); Kuhn in v. d. Decken, Reisen in Ost- Afrika 3?: 11 (1879).
Pteris L. Sp. Pl. 2: 1073 (1753), in part. Fili Ludwig, Instit. Hist. Ed. 2, 149 (1757), nomen dubium; sensu Woynar, Hedwigia эб: 383 (1915). Cincinalis Gled. Syst. Pl. 290 (1764), emend Trevis. Atti Soc. Ital. sc. nat. 17: 239 (1875). Asplenium Bernh. Sehrader's Journ. 1799!: 309 (1799), in part. Allosorus Bernh.
12 Rhodora | JANUARY
Schrader’s neues Journ. 1?: 36 (1806), sensu Pr. Tent. Pterid. 151 (1836), in part. Paesia St.-Hil. Voy. Brésil 1: 381 (1833), sensu Moore, Gard. Chron. 878 (1858), in part. Eupteris Newm. Phytologist 2: 278 (1845). Ornithopteris (Ag.) J. Sm. Hist. Fil. 297 (1875). Aquilina Pr. ex Diels in Engl. & Prantl, Nat. Pfl. 1%: 296 (1899), in synonymy. Filix-foemina Farwell, Am. Mid. Nat. 12: 290 (1931).
Although it is generally considered that Scopoli took up Pteridium Gled. for the Bracken, as a segregate genus from Pteris L., and it would therefore be the earliest valid generic name under the International Rules, some have argued that he proposed it as a substitute name for Pteris L. In order to meet this argument, Pteridium has been conserved (International Rules of Botanical Nomenclature, 131 (1935) ).
Rhizome subterranean, extensively creeping, repeatedly branched, invested with hairs but not scales, the stele a perforated solenostele, true vessels present; fronds alternate, large; stipe relatively long, the vascular bundles numerous; blade coarse, pinnately divided, usually tripinnate, the lower pinnae with nectaries at the base; the segments very numerous, ovate to linear, the margin revolute; veins free; sori marginal, mostly continuous; sporangia borne between the outer indusium, the modified margin of the segment, and the inner indusium;" re- ceptacle a vascular strand connecting the vein-ends, the inner indusium arising at its inner side; spores brown, very finely spinulose, tetrahedral-globose.—Represented by a single world- wide species, found in all temperate and tropical regions.
PrERIDIUM AQUILINUM (L.) Kuhn in v. d. Decken, Reisen in Ost-Afrika 3?: 11 (1879). МАР 1. BRACKEN. Characters of the genus.
KEY TO THE SUBSPECIES AND VARIETIES”
a. Ultimate segments adnate, or equally decurrent and surcur- rent, or surcurrent, or narrowed at the base, not having a farinaceous appearanee beneath; pubescence of the lower surface lanuginose or absent. (ssp. typicum) b.
b. Fertile and sterile indusium ciliate, or pubescent on the
outer surface, or both; ultimate segments densely or sparsely pubescent beneath between the margin and the midnerve; pinnules nearly at right angles to the costae. с.
u The inner indusium is quite variable in its structure. In some specimens it is a continuous membrane, as figured by Luerss. in Rabenh. Krypt. Fl. Ed. 2, 3: 101, fig. 81 and Bower (after Luerss.), The Ferns, 3: 43, in others it is an irregularly broken mem- brane and in some it consists only of a few hairs.
1? The user can expect that a considerable number of intermediate and non-typical speci- mens, especially shade-forms, will not run down satisfactorily.
19411 Tryon, Jr..—Revision of the Genus Pteridium 13
c. Pinnae and pinnules long-acuminate; ultimate segments faleate or subfalcate; fertile indusium 0.3 mm. or less wide, the sterile 0.2 mm. or less wide (India to Java, New Guinea and Formosa)...2. var. Wightianum, р. 22 c. Pinnae and pinnules short-acuminate to obtuse; ulti- mate segments straight; fertile indusium 0.3 mm. or more wide, the sterile 0.2 mm. or more wide. d. d. Upper surface of the ultimate segments glabrous or subglabrous, or if pubescent, the margin gla- brous. e. e. Rachis more or less pubescent; blade lanuginose be- neath (Europe, Africa and adjacent islands). 1. var. typicum, p. 15 e. Rachis glabrous; blade subappressed-lanuginose be- neath (Hawaiian Islands) .... 5. var. decompositum, p. 40 d. Upper surface of the ultimate segments pubescent, at least near the margin, and the margin pubescent. f. f. Sterile indusium 0.4 mm. or more wide; the fertile . portion of the indusium no broader than the sterile on the same segment (Mexico, Guate- ШААП Honduras) co p. 4. var. Feei, p. 37 f. Sterile indusium 0.25 mm. or less wide; the fertile portion of the indusium broader than the sterile on the same segment (western North America and northern Mexico)...3. var. pubescens, p. 26 b. Fertile and sterile indusium glabrous; ultimate segments glabrous beneath between the margin and the mid- nerve, or slightly pubescent on part of the segment; pinnules at an oblique angle to the costae. g. g. Blade usually quadripinnate, ovate; most of the ultimate segments narrowed at the base, the midnerve beneath glabrous or sub- glabrous CATION)» 4 t ee 8. var. africanum, p. 51 g. Blade usually bipinnate-pinnatifid or tripinnate, rarely tripinnate-pinnatifid, broadly triangu- lar; the ultimate segments adnate or more often broadest at the base, the midnerve'be- neath moderately pubescent, sometimes glabrous. h. h. Margin of the ultimate segments moderately pubescent; longest entire segment or en- tire part of a segment about four times as long as broad; the terminal segments mostly 5-8 mm. wide (North America, northern Europe and eastern Asia). 6. var. latiusculum, p. 41 h. Margin of the ultimate segments glabrous or subglabrous; longest entire segment or part of a segment about nine times as long as broad; the terminal segments mostly 2-4.5 mm. wide (southern and eastern United States) ..7. var. pseudocaudatum, p. 48 a. Some of the ultimate segments decurrent only, or more strongly decurrent than surcurrent, usually having a farinaceous appearance beneath; pubescence of the lower surface straight, appressed or arachnoid, rarely sub- lanuginose or absent (ssp. caudatum). 7.
| JANUARY
Rhodora
14
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1941] Tryon, Jr.,—Hevision of the Genus Pteridium 15
i. Ultimate segments having a farinaceous appearance beneath, except in some glabrous forms; pubescence straight, ap- pressed or arachnoid beneath. 7. j. No free lobes present on the rachis, costae and costules; the midnerve on the lower surface of the ultimate segments usually glabrous; fertile portion of the in- dusium broader than the sterile on the same segment; cells of the sterile indusium in rather definite rows, large, averaging 31 per mm. on the margin, the in- dusium not more than 5 cells wide (southernmost United States, West Indies, Mexico, Central America and northern South America) ........... 9. var. caudatum, p. 54 j. Free lobes present on the rachis, costae and costules; the midnerve on the lower surface of the ultimate segments usually dark-pubescent; fertile portion of the indusium usually not broader than the sterile on the same segment; cells of the sterile indusium irregularly arranged, small, averaging 48 per mm. on the margin, the indusium at least 6, usually 8 cells wide. k. k. Undersurface of the segments with membranous wings on the veins and midnerve, arachnoid-pubescent ; margin of the segments often pubescent (West Indies, Mexico, Central America and South Amer- ICON NE ате Е 10. var. arachnoideum, p. 57 k. Undersurface of the segments without membranous wings on the veins and midnerve, appressed-pubes- cent with straight hairs; margin of the segments glabrous (Australia, New Zealand and Polynesia). 1l. var. esculentum, p. 61 1. Ultimate segments not having a farinaceous appearance beneath, sublanuginose-pubescent beneath (India to Sumatra, Philippine Islands and northeastern Aus- Unna Puts reg M 12. var. yarrabense, p. 63
PTERIDIUM AQUILINUM ssp. typicum
Growing tip of the rhizome usually with a tuft of dark hairs, sometimes naked; rachis glabrous to densely pubescent; pinnules at right angles or at an oblique angle to the costae; ultimate segments broadly ovate to oblong, sometimes linear, adnate or surcurrent, or equally surcurrent and decurrent, or narrowed at the base, the upper surface glabrous to quite pubescent, the lower surface lanuginose-pubescent or glabrous, not having a farina- ceous appearance, cells of the sterile indusium of medium size, usually irregularly arranged. Map 1 (dots).
1. PrERIDIUM AQUILINUM var. typicum,'? PLATE 650, ric. 2,
13 Only the more important, the more obscure and scattered, and the more widely used subspecifie names and combinations are cited. About one hundred new subspecifie names and combinations in Lowe, Our Native Ferns, Druery, British Ferns and their Varieties, Junge, Die Pteridophyten Schleswig-Holsteins, Kaulfuss, Die Pteridophyten and Gandoger, Flora Europae terrarumque adjacentium are omitted from the synonomy. Some names from other well known but less detailed works are also omitted. All of these names ap- parently represent trivial variations. Gandoger’s treatment (op. cit. 27: 180 (1891)), the opening portion of which is here quoted, amounts almost to a naming of individuals:
16 Rhodora [JANUARY
PLATE 651, ric. 4, МАР 4. Illustrations: Fl. Dan. 13: t. 2303 (1840); Waldner, Deutschl. Farne t. 17 (1883); Schk. Krypt. Gew. t. 95 (1809); Lowe, Native Ferns 2: 407, t. 61 (1867) ; Marloth, Fl. S. Africa 4: pl. 37 (1915), habitat.
Pteris aquilina L. Sp. Pl. 1075 (1753). Cincinalis aquilina (L.) Gled. Verm. Abh. 1: 24 (1765), reference taken from Woy- nar, Hedwigia 56: 381 (1915). Pteris nudicaulis Güld. Reisen Russ. 1: 421 (1787), according to C. Chr. Ind. Fil. 603 (1906). Pteris borealis Salisb. Prod. 402 (1796). Asplenium aquilinum (L.) Bernh. Schrader's Journ. 1799!: 310 (1799). Pteris capen- sis Thunb. Prod. 2: 172 (1800) ; fragment of type at NY. Pteris lanuginosa Bory ex Willd. Sp. Pl. 5: 403 (1810); fragment of type at NY. Pteris aquilina L. var. ciliata Opiz, Kratos 2!: 19 (1820) 1+ Pteris aquilina L. var. sinuata Opiz, Kratos 2': 19 (1820). Pteris foemina Gray, Nat. Arr. Brit. Pl. 2: 16 (1821). Allosorus aquilinus (L.) Pr. Tent. Pterid. 153 (1836). Allosorus lanuginosus (Bory ex Willd.) Pr. Tent. Pterid. 154 (1836). Allosorus hottentottus Pr. Tent. Pterid. 154 (1836). Allosorus acutifolius Pr. Tent. Pterid. 154 (1836), nomen nudum. Al- losorus villosus Pr. Tent. Pterid. 154 (1836), nomen nudum. Allosorus tauricus Pr. Tent. Pterid. 154 (1836), nomen nudum. Pteris brevipes Tausch, Flora 19°: 427 (1836). Pteris lanuginosa Bory ех Willd. var. capensis (Thunb.) Ag. Rec. Pterid. 51 (1839). Pteris plebia R. Br. ex Ag. Rec. Pterid. 52 (1839), in synonymy. Pteris coriifolia Kze. Linnaea 182: 120 (1844). Pteris aquilina L. var. vera Moore, Handb. Brit. Ferns, Ed. 1, 134 (1848), refer- ence taken from Moore, Pop. Hist. Brit. Ferns, 167 (1851). Pteris aquilina L. var. integerrima Moore, Handb. Brit. Ferns, Ed. 1, 134 (1848), reference taken from Moore, Pop. Hist. Brit.
“20, Pteris L. 1. PTERIS AQUILINA L. Hab. Europa tota.
Lobi sup. integri 4 mm. longi 2
Lobi sup. integri 5 mm. longi 5
Lobi sup. integri 6 mm. longi 8
Lobi sup. integri 7 mm. longi 17 1 Lobi sup. integri 8 mm. longi 28
Lobi sup. integri 9 mm. longi 34
Lobi sup. integri 10 mm. longi 40
Lobi sup. integri 11 mm. longi 45
Lobi sup. integri 12 mm. longi 47
Lobi sup. integri 14 mm. longi— Dordogne, Payzac (Gdgr.) Pteris attenta Gdgr.
Pinnae remoti . 3 2 \ Pinnae contiguae 4
Lobi obtusi- Pedem., Vallées Vaudoises (Rostan) P. polystichoides Сайт. 3 Lobi acuti- Basses-Pyren., m. Lestibette (Gdgr.) P. oreodoxa Сарт.”
“I am indebted to Prof. Dr. Adolf Pascher for the references to Kratos, a periodical apparently not in the United States.
19411 Tryon, Jr..—Revision of the Genus Pteridium I
Ferns, 167 (1851). Pteris aquilina L. var. pubescens Spreng. ex Liebm. Vid. Selsk. Skr. s. 5, 1: 225 (1849), as to basinym, Pteris lanuginosa Bory ex Willd., not as to plant. Pteris aquilina L. var. multifida Moore, Handb. Brit. Ferns, Ed. 3, 226 (1857). Pteris aquilina L. var. crispa Moore, Handb. Brit. Ferns, Ed. 3,226 (1857). Pteris aquilina L. var. pubescens Afz. fil. ex Hook. Sp. Fil. 2: 198 (1858), in synonymy. Pteris aquilina L. var. lanuginosa (Bory ex Willd.) Hook. Sp. Fil. 2: 196 (1858). Al- losorus capensis (Thunb.) [incorrectly attributed to Presl by] Pappe & Raws. Syn. Fil. 32 (1858). Allosorus coriifolius (Kze.) Pappe & Raws. Syn. Fil. 31 (1858). Paesia corufolia (Kze.) Moore, Gard. Chron. 1858: 878 (1858). Aquilina vulgaris Pr. ex Milde, Fil. Eur. 45 (1867), in synonymy. Pteris Heredia Clem. ex Colm. Enum. crypt. Esp. y Port. 1: 16 (1867), refer- ence taken from C. Chr. Ind. Fil. 599 (1905) and Colmeiro, Enum. Plantas Hisp.-Lusit. 5: 437 (1889). Paesia aquilina (L.) Keys. Pol. Cyath. Hb. Bung. 22 (1873). Ornithopteris aquilina (L.) J. Sm. Hist. Fil. 298 (1875). Cincinalis lanuginosa (Bory ex Willd.) Trevis. Atti Soc. Ital. se. nat. 17: 239 (1875). Pteris gracilis Paterson in Hennedy, Clydesd. Fl. Mem. Ed. 255 (1878). Pteridium aquilinum (L.) Kuhn var. lanuginosum (Bory ex Willd.) Kuhn in v. d. Decken, Reisen in Ost-Afrika 3?: 11 (1879). Pteris aquilina L. var. abbreviata Gillot, Bull. Soc. Bot. France 29: June, xxii (1882). Pteris abbreviata (Gillot) Gerard, Bull. Soc. Bot. France 29: June, xxii (1882). Pteris aquilina L. var. transsilvanica Schur, Enum. Pl. Transs. 841 (1885), nomen nudum. Pteridium aquilinum (L.) Kuhn var. brevipes (Tausch) Luerss. in Rabenh. Krypt. Fl. Ed. 2, 3: 107 (1889). Pteridium aquilinum (L.) Kuhn var. wmbrosum Luerss. in Rabenh. Krypt. Fl. Ed. 2, 3: 107 (1889). Pteris aquilina L. var. normalis О. Ktze. Rev. Gen. 2: 820 (1891). Pteris aquilina L. f. lanuginosa (Bory ex Willd.) O. Ktze. Rev. Gen. 2: 820 (1891). Pteridium aquilinum (L.) Kuhn var. osmundaceum Christ, Beiträge Krypt. Schweiz 12: 54 (1900). Pteridium aquilinum (L.) Kuhn var. capense (Thunb.) Christ, Beiträge Krypt. Schweiz 1°: 55 (1900). Pteridium capense (Thunb.) Krasser in Zahlbruckner, Ann. Nat. Hofm. Wien, 15!: 4 (1900). Pteridium lanuginosum (Bory ex Willd.) Clute, Fern Bull. 8: 38 (1900), as to name-bringing synonym, not as to plant. Pteridium ceheginense Barnola, Bol. Soc. Aragonesa Cienc. Nat. 11: 35 (1912). Pteridium aquilinum (L) Kuhn var. crispulatum Barnola, Bol. Soc. Aragonesa Cienc. Nat. 11: 35 (1912). Pteridium aquilinum (L.) Kuhn ssp. capense (Thunb.) Bonap. Notes Ptérid. 2: 66 (1915). Ат aquilina (L.) Woynar, Hedwigia 56: 383 (1915). Filix-foemina aquilina (L.) Farwell, Am. Mid. Nat. 12: 290 (1931).
In 1848 Moore published Pteris aquilina L. var. vera but he
18 Rhodora [JANUARY
did not base it on Pteris aquilina L. He described it as having the secondary pinnules pinnatifid, while the Fuchs plate, the type of Pteris aquilina, has the secondary pinnules entire (in fact the pinnules are only pinnatifid). In contrast to his var. vera he had var. integerrima, secondary pinnules entire. He apparently thought of var. vera as the more common and hence more "typical" phase and his description of it excludes the type of Pteris aquilina.
Kuntze published a Pteris aquilina L. var. normalis in 1891 but did not actually base it on Pteris aquilina L. and circum- scribed it much more broadly than the typical variety in the present treatment. “U. St.: Oil City" includes var. latiusculum and “Hongkong” includes either var. latiusculum or var. Wightianum.
Because of the above objections to taking up either var. vera Moore or var. normalis O. Ktze., I am designating the typical variety as var. typicum, definitely based on the type of Pteris aquilina L.
Growing tip of the rhizome with a tuft of dark hairs; frond 0.4—4 m., usually about 1—1.5 m. high, the taller fronds scandent, vernation gleichenioid; stipe longer or shorter than the blade; blade 2-20 dm., usually about 6—10 dm. long, usually ovate- triangular, less often ovate or broadly triangular, not ternate, bipinnate-pinnatifid or tripinnate, less often tripinnate-pinnatifid, rarely quadripinnate-pinnatifid ; rachis usually moderately pubes- cent, less often densely pubescent or subglabrous; pinnae and pinnules short-acuminate to obtuse; pinnules usually nearly at right angles to the costa, sometimes at an oblique angle; costules slightly to densely pubescent beneath and less so above; penulti- mate segments usually pinnatifid, less often pinnate or pinnate- pinnatifid; longest entire segment or part of a segment from three to six, usually about four, times as long as broad; ultimate segments usually straight, rarely subfalcate, adnate or broadest at the base, the upper surface usually glabrous or slightly pubescent, rarely moderately pubescent, the margin glabrous or subglabrous, rarely moderately pubescent, the lower surface usually densely pubescent, less often slightly pubescent, rarely glabrous or pubescent only on the midnerve; fertile and sterile indusium ciliate and sometimes also pubescent on the outer sur- face, the fertile usually 0.8 mm. or more wide, the sterile usually 0.2 mm. or more wide, the fertile portion broader than the sterile on the same segment, or no broader.
19411 Tryon, Jr..—Revision of the Genus Pteridium 19
Type: Figure of Filix femina Fuchs, Hist. 596, misprinted 569 (1542). Linnaeus did not have a specimen of Pteris aquilina in his herbarium in 1758 and the only illustration cited by him is taken as the type.
TYPE LOCALITY: Europe.
I have not been able to separate the African Bracken from the European. Extreme specimens from Africa differ from the typical European plant (Tidestrom 13295, France) in having a densely pubescent, flexuous rachis and the fertile indusium no broader than the sterile on the same segment. However, these characters are not always correlated and they show no definite geographic segregation. Specimens with the rachis densely pubescent and flexuous have been seen from Mauritius, “Ex. Herb. Wm. Boott" (G) and Madagascar, Herb. Field Mus. no. 595745. Specimens from British East Africa, E. Heller; Natal, Abraham 28; Cape of Good Hope, U. S. Nat. Herb. no. 22419; St. Thomas, Moller 33 and Sierre Leone, Johnston 7 have the rachis densely pubescent but straight. Distributed over es- sentially the same area are specimens that have a straight glabrous rachis: Egyptian Sudan, Lynes 186; British East Africa, Mearns 1229; Kilimanjaro, Abbott; Madagascar, Webb (?) 113 (G); Bourbon, Halsey (U.S. Nat. Herb. no. 1285441) ; and Cape of Good Hope, Sim 1579. Most of the European and about half of the African specimens have the fertile indusium broader than the sterile on the same segment. Specimens from the Azores, Madeira and Canary Islands and from northernmost Africa are distinctly of the European type.
Shade forms often approach var. latiusculum in some charac- ters: they may be almost completely glabrous (Ziesché, Herb. Field Mus. no. 764960; Prov. Westfalen (G); Kaulfup, Bavaria (US)) or only slightly pubescent with the sterile indusium con- siderably less ciliate than normally (Ziesché, Reisengebirge, Silesia; Richter, Hungary).
Intermediates between var. typicum and var. latiusculum are only shghtly pubescent beneath except on the midnerve and have the fertile and sterile indusium only slightly ciliate. Such are Danzig, July 24, 1876, Baenitz (F); Pl. Scand. ex insula Got- landia, June 16, 1894, Johansson (US); Plantae Goreanae, Taquet 2317 (US); and Hungary, Aug., 1893, Marton (F).
20 Rhodora [JANUARY
Some specimens, here referred to var. latiusculum, approach var. typicum in one or more characters. The sterile indusium may be slightly ciliate and the blade pubescent beneath between the margin and the midnerve and the blade ovate: Pl. Sibiricae Exsiec. 1288, Tomsk; Honig, Bavaria; near Berlin, 1844. In A. O. Olson, Vestrogothia, Sweden, Aug. 17, 1915 (F) the in- dusium is slightly pubescent.
Several minor variations have been named, some of which may be worthy of recognition as forms, but I do not feel justified in recognizing them in view of the limited amount of material I have seen and the lack of field experience. Some of the varia- tions that have been described certainly do not deserve recogni- tion. Pteris aquilina var. abbreviata Gillot, for instance, is a much stunted plant found growing under extremely unfavor- able conditions, on rocks and bathed in a sulphurous vapor at 20? C. Pteridium ceheginense Barnola is a young plant with ab- normal leaf-cutting and Pteris gracilis Patterson is also а young plant.
Var. typicum differs from var. latiusculum in its gleichenioid rather than equal vernation, in having the undersurface of the segments densely pubescent rather than pubescent only on the midnerve, in the glabrous rather than pubescent margin of the segments and the ciliate rather than glabrous fertile and sterile indusium. Also the frond is considerably taller on the average and the blade is ovate to ovate-triangular and evenly pinnate rather than broadly triangular with the basal pinnae considerably longer and broader than the second pair. The pinnules are usually at right angles to the costa rather than at an oblique angle and the penultimate segments are usually evenly pinnatifid rather than pinnate or pinnate-pinnatifid.
The critical differences between var. typicum and vars. Wightianum, pubescens and Feei, with which it is also closely related are discused under those varieties. It is perhaps most closely related to var. pubescens of western North America— probably representing another example of the well-known re- lationship of the floras of western North America and Europe.
Var. typicum grows in open woods, in clearings, at the edge of woodlands, in thickets, in pastures, in abandoned fields, in grassy places, on open hillsides, in forests and in recently burned-
19411 Tryon, Jr.—Revision of the Genus Pteridium 21
over areas. It is most common in dry places and in acid soils. It grows from sea-level to about 1800 m. in the Alps and to 3000 m. in the mountainous regions of Africa.
It ranges throughout Europe and all but the drier regions of Africa, and adjacent islands.
EURASIA.— SWEDEN: Helsingborg, July, 1862, Suetberg (Herb. Field. Mus. no. 335698). InELANp: Holywood, Oct. 20, 1884, (G). Great BRITAIN: Snowdon (Wales), July 19, 1905, Pease 8031 (О); Isle of Man, Sept., 1895, Underwood (NY). NETHER- LANDS: Haaksbergen, July 20, 1918, Boetje van Ruyven (Herb. Field Mus. no. 840219). Germany: Heidelberg, Sept. 19 (Herb. Field Mus. no. 756408) ; Münsterland, Prov. Westfalen, Aug. 19, 1905 (G); Zabtengebirge, Mittelberg, Silesia, Aug. 29, 1906, Ziesché (Herb. Field Mus. no. 764960) ; Reisengebirge, Silesia, July 20, 1903, Ziesché (G) ; Spandau, Brandenburg, Sept., 1880, Ruhmer (F. Schultz herb. norm. no. 975) (G, NY); Nürnberg, Bavaria, Aug. 26, 1902, Каміјир (US). CZECHOSLOVAKIA: Aug., 1911, Petrak (Fl. Bohemiae et Moraviae exsice. no. 603) (G). Huxcanv: Klausenberg, Aug. 16, 1901, Richter (G); Piliscaba, Sept., 1916, Filarszky & Kiimmerle (Fl. Hung. exsice. no. 431) (F, б, US); Orawieza, Wierzbicki (Herb. Field Mus. no. 47805). SWITZERLAND: Wilazek (G). France: Bures, Sept. 26, 1909, Jeanpert (Herb. Field Mus. no. 815531); Perignae, Charente, Sept. 4-11, 1893, Guillon (Mangier, Fl. selecta exsiec. no. 3395) (F); July 11, 1934, Tidestrom 13295 (F, G, US); May 9, 1935, Tidestrom 13509 (NY); Vicinity of Paris, June 28, 1911, Jean- pert (Herb. Field Mus. no. 815529). Sparx: San Sebastian, Barbour (NY). Porrucau: Porto, 1891, Buchtien (U. S. Nat. Herb. no. 1095027). Corsica: July 20, 1933, Aellen 1306 (F, US) ; June 17, 1933, Aellen 1316 (Е, US). IraALv: Venetia, Nov. 10, 1907, Fiori, Béguinot & Pampanini (Fl. It. exsicc. no. 705) (С); Capri, Aug. 29, 1909, Ware (G, US); Near Rome, Pisotta 34 (F). GmnkECE: Olympia, April 27, 1906, Pease 9081 (G). Turkey: Taurus Mts., Aug. 12, 1907, Wankow (NY); “Plantae Tauricae,” Wankow (US). Union or Soviet Soctauist RE- PUBLICS: Dist. of Chernomose (Black Sea), Kousenetzoff (G); Caucasus, Kuban Prov., May 9, 1907, Busch & Klopotow (Сб); Western Grusia, Caucasus, Meffert 20 and 910 (G).
AFRICA.—ALGERIA: Oran, May 9, 1935, Faure (U.S. Nat. Herb. no. 1674024, NY). Morocco: 1889, T. Williams 118 (US). Azores: 1890, Chute (G); July 18, 1894, Brown 334 (С). Mapema: 1853-56, Wright (Сб); 1888-42 (U. S. South Pacific Ex- ploring Expedition) (G). Canary IsraNps: Grand Canary, Feb., 1897, Cooke 107 (F, G, NY, US). ӨтЕВЕА Leone: Oct., 1875 (Herb. Field Mus. no. 69396) ; Freetown, April 8, 1882, H. H.
22 Rhodora [JANUARY
Johnston 7 (G). LiBERIA: Grand Bassa, Dinklage 1628 (G); along Dukwai River near Firestone Plantations, 1928, G. P. Cooper 21 (F, NY, US). Cameroon: Jan., 1919, Gocker 133 (О). Sr. THoMas (Is.): 1885, Moller 33 (US). FRENCH EQUATORIAL ArRICA: Fernand Vaz, French Congo, 1917, Aschemeier 19 (US). Union or боотн Arrica. САРЕ or Соор Hore: Vicinity of Cape Town (U. 8. South Pacifie Exploring Expedition) (U. S. Nat. Herb. no. 22419); King Williamstown, 1892, Sim 1579 (G). NaTAL: Mapumulo, 1867—69, Abraham 28 (С, US); Buchanan 534d (US). 'TnaNsvaarL: Bokfontein, Aug. 3, 1934, Mogg 4755 (US); Rustenburg, (District of) Rustenburg, Sept., 1910, Col- lins (Herb. Field Mus. no. 653635).—Mapacascar: 1897 (Herb. Field Mus. nos. 595715, 595725, 595745, U. S. Nat. Herb. nos. 1431040, 1431121); Webb(?) 113 (G). Восввох: (Herb. Field Mus. no. 830986); Halsey (U. S. Nat. Herb. no. 1285441). MaunririUs: 1867, Pike (U. S. Nat. Herb. no. 593158); (U. S. Nat. Herb. no. 516236). TANGANYIKA (GERMAN East AFRICA): Kilimanjaro, 1889-90, Abbott (U. S. Nat. Herb. nos. 22433, 22472); Ulugurus, Bunduki, Aug. 2, 1935, Bruce 96 (US). ZANZIBAR: Oct., 1873, Hildebrandt (NY). Ucawpa: Mt. De- basien, Jan., 1936, Eggeling 2703 (G); Vicinity of Kigomma, Dec. 30, 1909, Mearns 2617 (US); Kikayo, Diimmer 551 (US). Kenya (Bnrrisu. East Arrica): Mau Range, July 27, 1923, A. G. Curtis 848 (G); Lake Naivasha, July 17 to Aug. 15, 1909, Mearns 929 (US); Wambugu, Sept. 13, 1909, Mearns 1229 (G, US); Mbololo, Nov. 7-11, 1911-12, E. Heller (U. S. Nat. Herb. no. 634399). ЕсүртІАХ Supan: Jebel Marra, Dafur, Dec., 1921, Lynes 186 (US). ABvssiNIA: 1842, Schimper 856 (US).
2. PTERIDIUM AQUILINUM var. Wightianum (Ag.), n. comb. PLATE 650, ric. 1, PLATE 651, Fic. 3, мар 2. Illustration: Bedd. Ferns $. India, t. 42 (1863).
Pteris revoluta Bl. Enum. Pl. Jav. 214 (1828). Pteris excelsa Bl. Enum. Pl. Jav. 213 (1828). Pteris lanigera Bl. Enum. PI. Jav. 214 (1828). Pteris densa Wall. List no. 99 (1829), nomen nudum, isotype at US. Pteris firma Wall. List no. 100 (1829), nomen nudum. Pteris terminalis Wall. List no. 101 (1829), nomen nudum. Pteris recurvata Wall. List no. 113 (1829), nomen nudum; ex Ag. Rec. Pterid. 50 (1839), isotype at US. Pteris Wightiana Wall. List no. 2178 (1829), nomen nudum. Allosorus recurvatus (Wall.) Pr. Tent. Pterid. 154 (1836), nomen nudum. Pteris recurvata Wall. ex Ag. var. Wightiana Ag. Rec. Pterid. 50 (1839), as Wigtiana, epithet taken from Pteris Wightiana Wall. Pteris villosa Fée, Gen. Fil. Mém. Fam. Foug. 5: 128 (1850-1852). Cincinalis villosa (Fée) Trevis. Atti Soc. Ital. se. nat. 17: 239 (1875). Pteridium aquilinum (L.) Kuhn
19411 Tryon, Jr.,—HRevision of the Genus Pteridium 23
var. osmundoides Christ ex Léveillé, Bull. Acad. Geogr. Bot. 20: no. 243, 9 (1910), nomen nudum. Pteridium capense (Thunb.) Krasser var. densa Nakai, Bot. Mag. Tokyo 39: 109 (1925), epithet taken from Pteris densa Wall. Pteridium revolutum (Bl.) Nakai, Bot. Mag. Tokyo 39: 109 (1925).
Growing tip of the rhizome with a tuft of dark hairs; frond 0.60—4 m. high, the taller fronds scandent, vernation gleichenioid; stipe usually shorter than the blade; blade 0.3-3 m., usually about 1.1—5 m. long, ovate-triangular, or less often pentagonal, not ternate, usually tripinnate-pinnatifid, less often bipinnate- pinnatifid or tripinnate; rachis usually densely or very densely pubescent, rarely becoming subglabrous; pinnae and pinnules usually long-acuminate, infrequently short-acuminate; pinnules usually nearly at right angles to the costa, less often at a some- what oblique angle; costules slightly to densely pubescent be- neath and less so above; penultimate segments usually pin- natifid, less often pinnate or pinnate-pinnatifid; longest entire segment or part of a segment from three to seven, usually about four, times as long as broad; ultimate segments usually falcate or subfaleate, adnate or broadest at the base, the upper surface usually glabrous or slightly pubescent, the margin glabrous to moderately pubescent, the lower surface usually densely pubes- cent, less often slightly pubescent, rarely glabrous or pubescent only along the midnerve; fertile and sterile indusium ciliate and sometimes also pubescent on the outer surface, rarely gla- brous, the fertile usually 0.3 mm. or less wide, the sterile usually 0.2 mm. or less wide, the fertile portion no broader than the sterile on the same segment.
Tyre: Wallich 2178, at Kew (not seen).
Type Locarrry: Dindygul, India.
Specimens from the Himalayas (R. R. & I. D. Stewart 4761, 4048; Stewart 6492) are almost completely glabrous and Stewart 6492 has the ultimate segments narrowed at the base. How- ever, they have the long-acuminate pinnae and pinnules of typieal var. Wightianum and probably represent only a local 'ariation.. Other material from the same region is entirely typical. Some specimens approach var. typicum and var. latiusculum in having a nearly glabrous rachis (Henry, China; Fang 3281, China) or acuminate to obtuse pinnules (Bartsch 153, Philippine Is.; Sallet, Indo-China).
Var. Wightianum can be separated from var. typicum, with which it is apparently closely related, by its much more densely pubescent rachis, its long-acuminate rather than subacute or
Rhodora [JANUARY ‘ ame
Re / Ag
Ranges of PrTERIDIUM AQUILINUM, var. WiGHTIANUM (map 2); var. PUBESCENS (map 3).
19411 Tryon, Jr.—Revision of the Genus Pteridium 25
obtuse pinnae and pinnules, its faleate rather than straight ultimate segments and the narrower fertile and sterile indusium. The critical differences between var. Wightiamum апа var. latiusculum, with which it intergrades, are discussed under that variety.
Var. Wightianum grows on dry hillsides, in jungle clearings, on waste slopes, on eraters and in grassland, usually in sterile, often dry, soil. It occurs from 700 m. to 3300 m. in India and up to 2500 m. in China and 2800 m. in Papua.
It ranges from northwesternmost India east to Formosa, south to Ceylon, Java and British New Guinea.
Inpa: N. W. India, 1871, J. L. Stewart 3626 (NY); Nazara, N. W. Himalayas, May 16, 1896 (US Nat. Herb. no. 1274966) ; Simla Region, Simla, Sept. 18, 1883, Blanford (МҮ); Sonamarg, Kashmir, July 28, 1921, R. R. Stewart 6492 (NY, US); Tragbal, Kashmir, July 31, 1919, R. R. & I. D. Stewart 4761 (NY, US); Murree Hills, Changla Gali, Punjab, Sept. 12, 1918, R. R. & I. D. Stewart 4048 (NY, US); Halann, Kulu, Punjab, May 21, 1931, Koelz 1980 (US); Kumaon, Strachey & Winterbottom 8 (G); Dalhousie, Punjab, June 9, 1917, R. R. & I. D. Stewart 2182 (NY, US); Nepal, 1820, Wallich 113 (US), isotype of Pteris recurvata Wall.; Khasi Hills, Shillong, Assam, Sept., 1888, Mann (NY, US); Sikkim, Hooker (G); Nilghiri, Thomson (б); 40 miles north of Darjeeling, Dec., 1884, native collector (NY). CEvLoN: Beckett 182 (G); (U.S. Nat. Herb. no. 816885); (U. 5. Nat. Herb. no. 22423); С. Wall (NY). білм: Doi Sutep, Dec. 15, 1928, H. M. Smith 410 (G, US); Doi Chang Mt. near Hue San, Chiengmai Prov., Jan. 10, 1922, Rock 1720 (NY, US); Jan Khien, Doi Sootep, Chiengmai, Aug. 14, 1931, Cunniff 7 (NY). FnENcH Iwpo-CHia: Annam, Massif de Bah-Na, 30 miles southwest of Tourane, Aug., 1920, Sallet (G, NY, US) ; Cochin- ehina, May, 1870, Pierre 5793 (US) ; Angkor, Cambodia, Jan. 14, 1926, Н. M. Smith 295 (US). FEDERATED MALAY STATES: Perak, 1887, Wray 1486 (US); Penang, 1822, Wallich 99 (US), isotype of Pteris densa Wall.; Pulan Tulai, Pahang, May 27, 1927, Henderson 18525 (US). CHiNA. Kwanctune: Canton and vicinity, Nov. 11, 1917, Levine 1834 (С). Hartnan: Hainan, Dee. 18, 1933, Wang 35562 (NY). Kwricuow: Kyingtenshan, Tsunyi, Jan. 3, 1930, T'siang 5241 (G, NY); Liang Feng Yah, Tsunyi Hsien, July 31, 1931, Steward, Chiao & Cheo 53 (F, NY, US). Yunnan: Between Mohei and Moakai, March 20-Аргі 7, 1922, Rock 2906 (G). SzecHuan: Mt. Omei, Omei Hsien, 1928, Fang 3281 (G); Patung District, Feb., 1887, Henry (G).—Fon- mosa: Arisan, Dec. 4, 1933, Kanehira 2995 (NY, US); Shakko,
26 Rhodora [JANUARY
Dec., 1913, Faurie (E. Rosenstock exsice. no. 51) (G); Mt. Taihei, Taiheisan, Sept. 28, 1926, Bartlett 6084 (US).
PuiniPePINE ISLANDS: Dumaguete, Island of Negros, Prov. Negros Oriental, June, 1908, Elmer 10349 (G, NY, US); Port of Dos Amigos, Twai Twai Island, Feb. 19, 1908, Bartsch 153 (G, NY, US); Twin Peaks, Benguet Road, Luzon, March 2, 1908, Bartsch 210 (G, NY, US); Bosoboso, Prov. Rizal, Luzon, June, 1896, Ramos (Herb. Phil. Bureau Sci. no. 1054) (US); Bukidnon, Mindanao, July-Aug., 1913, Escritor (Herb. Phil. Bureau Sci. no. 21414) (US). Borneo: Korthals (NY). SUMATRA: Aels Kanopan, Loendoet Concession, Koealoe, March 19, 1927, Bartlett 6993 (US); Toba, Owwchand 204 (G). Java: Tjiboeroem, Preanger Prov., April 12, 1909, Palmer & Bryant 171 (US); vicinity of Goenoeng Boender, Batavia Prov., May 16, 1909, Palmer & Bryant 529 and 541 (US). Papua (BRITISH New Guinea): Murray Pass, Wharton Range, Central Division, June-Sept., 1933, Brass 4634 (G, NY).
3. PTERIDIUM AQUILINUM var. PUBESCENS Underw. Our Nat. Ferns, Ed. 6, 91 (1900). PrarE 650, ric. 3, PLATE 652, FIG. 5, MAP З. Illustration: Ashton, Pl. Rocky Mt. Nat. Pk. 22 (1933), habitat.
Pteris aquilina L. var. lanuginosa Bong. Mém. Acad. St. Petersb. s. 6, 2: 176 (1832), isotype at G. Pteridium lanuginos- um (Bory ex Willd.) Clute, Fern Bull. 8: 38 (1900), as to plant, not Pteris lanuginosa Bory ex Willd.; nomen provisorium. Pteris lanuginosa sensu Clute, Fern Bull. 8: 37 (1900), not Bory ex Willd. Pteris aquilina pubescens (Underw.) Clute, Fern Bull. 15: 124 (1907). Pteridium latiusculum (Desv.) Hieron. ex Fries var. pubescens (Underw.) [combination incorrectly at- tributed to Underw. by] Seymour, Host Ind. 25 (1929). Filiz- foemina aquilina (L.) Farwell var. lanuginosa (Bong.) Farwell, Am. Mid. Nat. 12: 290 (1931). Pteris aquilina L. var. pubescens (Underw.) [combination apparently incorrectly attributed to О. Ktze. by] Hanna, Am. F. Journ. 22: 6 (1932). Pteridium aquilinum (L.) Kuhn var. lanuginosum (Bong.) Fernald, Rho- dora 37: 247 (1935), not (Bory ex Willd.) Kuhn (1879).
Growing tip of the rhizome usually with a tuft of dark hairs; frond 0.3-5 m., usually about 0.8-2 m. high, the taller fronds scandent, vernation subgleichenioid; stipe usually shorter than the blade; blade 0.2-4 m., usually about 0.6—1 m. long, usually ovate-triangular, less often pentagonal or ovate, not ternate, usually tripinnate or tripinnate-pinnatifid; rachis slightly to moderately pubescent; pinnae and pinnules subacute to obtuse; pinnules usually nearly or quite at right angles to the costa, sometimes somewhat at an oblique angle; eostules usually slightly
Rhodora Plate 651
Wig. 1, middle pinna of PTERIDIUM AQUILINUM, var. Peer X 1; FIG. 2, upper pinna of var. DECOMPOSITUM, X 10; FIG. 3. tip of middle pinna of var. WiGHTIANUM, X 14; FIG. 4, upper half of middle pinna of var. TYPICUM, X 1
Rhodora Plate 652
Fic. 1, basal pinna. of PTERIDIUM. AQUILINUM, Var. LATIUSCULUM, X lá; FIG. 2, basal pinna of var. PSEUDOCAUDATUM, X 15; FIG. З, next to basal pinnule of a basal pinna of var. AFRICANUM, X 4%; FIG. 4, frond of variant of var. LATIUSCULUM, X 1/6; FIG. 5,
next to basal pinna of var. PUBESCENS, ХОМ.
19411 Tryon, Jr.—Revision of the Genus Pteridium 27
to moderately pubescent beneath and less so above; penultimate segments often pinnatifid, less often pinnate or pinnate-pin- natifid; longest entire segment or entire part of a segment from three to five, usually about four, times as long as broad; ultimate segments usually straight, adnate or broadest at the base, the upper surface slightly to quite pubescent, at least near the margin, the midnerve usually glabrous, the margin moderately pubescent, the lower surface usually densely pubescent, less often slightly pubescent, rarely glabrous except along the midnerve; fertile indusium usually slightly ciliate and slightly pubescent on the outer surface, rarely becoming glabrous with age, sterile indusium ciliate and sometimes also pubescent on the outer surface, the fertile usually 0.25-0.3 mm. wide, rarely up to 0.5 mm. wide, the sterile 0.25 mm. or less wide, very rarely 0.4—0.5 mm. wide, the fertile portion broader than the sterile on the same segment.
ТҮРЕ: No type designated, specimens labeled by Underwood in the Herbarium of the New York Botanical Garden typify the variety.
Type Locauiry: “Utah, California and northward.”
Shade forms! approach var. latiusculum in having the sterile
indusium glabrous or nearly so and the lower surface of the blade nearly glabrous, or in some cases more glabrous than in typical var. latiusculum—for example, Merrill & Wilcor 923. Wyoming; Nelson & Nelson 6733, Wyoming.
Intermediates between var. pubescens and var. latiusculum occur along the eastern border of the range of var. pubescens. Hanna! remarked that the Wyoming Bracken was “rather in- termediate between the eastern P. aquilina [var. latiusculum | and the western Р. aquilina, var. pubescens (Underw.) Kuntze.” In a more recent article, on the ferns of Colorado where var. latiusculum also occurs, Wherry! says that “The relations be-
15 І regard to ecological variation I think it is worthwhile to put on record the results of the following experiment. As part of the Carnegie Institution transplant experiments, the late Prof. Hall sent two plants of var. pubescens from Mather, California, to Mr. C. A. Weatherby at East Hartford, Connecticut. Mr. Weatherby received the plants in Septem- ber, 1922, grew them successfully, and had them under observation until July, 1929. One plant was grown next to an unprotected east wall and the other by a fence, exposed to the sun for the greater part of the day. The reactions of the two plants were similar. The outline of the blade did not change and the segments did not become noticeably broader or thinner. The fronds grew to be six feet tall and dependent upon a support to remain erect. The only marked change was that the density of the pubescence gradually decreased so that in 1929 the plants were very noticeably less pubescent than when first received. The general differences between the Connecticut and California localities are a more humid climate, a lower elevation and about a four-degree higher latitude at the former.
16 Hanna, Distribution of the Ferns of Wyoming. Am. F. Journ. 22: 6 (1932).
17 Wherry, Colorado Ferns. Am. F. Journ. 28: 136 (1938).
28 Rhodora [JANUARY
tween these two Brackens need further study, for their inter- gradation in this region is so marked as to cast doubt on their specific distinctness.” Some of the intermediates have the general characters of var. latiusculum except that they are pubes- cent beneath or have the pinnules set nearly at right angles to the costa. Others have an ovate blade and the sterile in- dusium slightly ciliate as in var. pubescens but are glabrous beneath except along the midnerve; or have the general char- acters of var. pubescens but have the pinnules at an oblique angle to the costa. Intermediate specimens are: Miller's Canyon, Huachuca Mts., Arizona, July 10, 1909, Goodding 170 (G, NY); San Francisco Mts., Arizona, Aug. 17, 1889, Knowlton 14 (б, US); Buckskin Mts., Arizona, June 30, 1909, Tidestrom 2336 (US); Central, New Mexico, Aug., 1895, Mulford 398 (NY); Rindoso Creek, Lincoln Co., New Mexico, July 1, 1895, Mearns (US); Buffalo Pass, Colorado, Aug. 11, 1898, Shear «€ Bessey (US) ; Southwest of Franktown, Douglas Co., Colorado, June 17, 1937, Wherry (US); Silver Reef, Utah, May 5, 1894, M. E. Jones 5176 (US) ; Gibbon Canyon National Park, Wyom- ing, Carleton 204 (F); Centennial Valley, Wyoming, Aug. 18, 1896, Nelson 2662 (NY).
Some specimens of var. latiusculum approach var. pubescens in characters of indusium and vestiture. A specimen from a burnt-over hillside, Moore 2221, West Virginia, is pubescent beneath between the margin and the midnerve and the fertile and sterile indusium is slightly ciliate and pubescent. The fol- lowing specimens from New York, Ohio, Massachusetts and Indiana are pubescent beneath between the margin and the midnerve: Washington Co., New York, July 30, 1890, Burnham (Herb. Cornell U.)18; Otsego Co., New York, Frost (Herb. Cor- nell U.); Cleveland, Ohio, Aug. 15, 1875 (Herb. Cornell U.); Line Station, Indiana, Aug. 2, 1876, Grassly (F); Granville, Hampden Co., Massachusetts, Sept. 19, 1913, Seymour 60 (G).
An interesting phase, approaching var. pubescens in outline of blade and pubescence is discussed under var. latiusculum.
Var. pubescens differs from var. latiusculum in having the tip of the rhizome with a tuft of dark hairs rather than nearly
18 Т am indebted to Dr. R. T. Clausen for bringing to my attention this material in the Cornell University Herbarium.
19411 Tryon, Jr..—Revision of the Genus Pteridium 29
naked, and an ovate-triangular, fairly evenly pinnate blade rather than a broadly triangular, ternate blade. Also the pin- nules are nearly at right angles to the costa rather than at an oblique angle, the lower surface is densely pubescent between the margin and the midnerve rather than pubescent only on the midnerve and the fertile and sterile indusium is ciliate and sometimes also pubescent rather than glabrous. The frond is a little taller on the average and certainly reaches an extreme that var. latiusculum never approaches.
Var. pubescens differs from var. typicum in the pubescent rather than usually glabrous margin of the segments and the markedly less ciliate and pubescent fertile and sterile indusium. The penultimate segments are usually not as evenly pinnatifid or pinnate.
Var. pubescens is also closely related to vars. Feei and de- compositum and the differences are discussed under those varieties.
In Michigan, Ontario and Quebec var. pubescens probably occurs as a pre-glacial relic on or related to local nunatak areas.!?
Var. pubescens grows in pastures, open forests, burnt-over areas, on open slopes, in thickets, and in woods, in damp or dry places; mostly in the Transition and Canadian Zones, from sea level up to 3000 m.
It ranges from southern Alaska to Mexico, east to Wyoming, Colorado and western Texas; isolated eastward in South Dakota, northern Michigan, Bruce Co., Ontario and Megantie Co., Quebec.
ALASKA: New Metlapatla, July 5, 1895 (Herb. Field Mus. nos. 366780, 366786) ; Sitka, Bongard (G), isotype of Pteris aquilina var. lanuginosa Bong.; Sitka, Aug. 2, 1916, J. P. Anderson 263 (US); Heyder, June 25, 1924, Whited 1284 (US); McDonald Lake, July 15, 1921, H. M. Smith (US); Petersburg, July 3, 1918, J. P. Anderson 652 (NY).
DoMINION oF CANADA.—QvUEBEC: Caribou Hill, Black Lake, Megantic Co., Aug. 26, 1915, Fernald & Jackson 11960 and 11961 (С). Ontario: Tobermory, Bruce Peninsula, July 28, 1933, Krotkov 6319 (US); Dunk’s Bay, Tobermory, Bruce Co., Aug. 20, 1933, Т. M. C. Taylor 6100 (G). ALBERTA: Vicinity of Banff, Aug. 29, 1899, McCalla 2434 (NY). British Cotum-
19 See Fernald, Critical Plants of the Upper Great Lake Region of Ontario and Michigan. Rhodora 37: 247 (1935).
30 Rhodora [JANUARY
BIA: Tulameen River, 1900, Kemp (NY); Revelstoke, Selkirks, July 3, 1905, Shaw 786 (G, NY, US); Near Victoria, Vancouver Island, July 28, 1908, Macoun 84096 (F).
UNITED STATES OF AMERICA.—MICHIGAN: Lake Manganese, 1 mile southeast of Copper Harbor, Keweenaw Co., July 23, 1936, Hermann 8231 (US); West Bluff, 315 miles west of Copper Harbor, Keweenaw Co., June 18, 1936, Hermann 7520 (NY); West Bluff, Keweenaw Co., July 4, 1934, Fernald & Pease 3033 (G, NY); Mackinac Island, July 28-29, 1898, Millspaugh 85 (F, G). Sours Daxora: (undoubtedly from the Black Hills) Gifford (G). Texas: Mt. Livermore, Davis Mts., Jeff Davis Co., Oct. 6, 1926, E. J. Palmer 32010 (С). Monrana: Near Apgar, Glacier National Park, July 30, 1937, Т. С. & Е. С. Yuncker 7006 (F); Belton, Aug. 22, 1903, Umbach 732 (F, NY, US). Ipamo: Trinity, Elmore Co., Aug. 13, 1910, Macbride 599 (F, G, NY, US) ; Lake Pend Oreille, Kootenai Co., Aug. 23, 1892, Sandberg, MacDougal & Heller 943 (US). Wyomina: Gibbon Canyon, Yellowstone National Park, Aug. 28, 1899, E. & A. Nelson 6733 (G, NY, US); Leigh's Lake, July 24, 1901, Merrill & Wilcox 923 (С, NY, US); Alpine, Lincoln Co., July 13, 1923, Payson & Armstrong 3486 (G); Bradley Creek, Grand Teton National Park, July 17, 1932, L. Williams 887 (G, NY). CoL- ORADO: Near Pagosa Peak, Aug. 25, 1899, С. F. Baker 128 (С, NY, US); Park Range above Steamboat Spa, Aug. 11, 1898, Shear & Bessey (NY). Uran: La Motte Peak, July 24, 1926, E. B. & L. B. Payson 5078 (G); Granite Canyon, Deep Creek Mts., Juab Co., June 20, 1933, Maguire & Becraft 2466 (G). Nevapa: East Humboldt Mts., Aug., 1868, Watson 1359 (G); Little Valley, Washoe Co., Aug. 14, 1902, C. F. Baker 1460 (С. NY, US). New Mexico: Jemez Canyon, Aug. 10, 1932, A. «€ R. A. Nelson 212 (С); Sacramento Mts., Lincoln Co., July 30, 1923, Eggleston 18881 (NY). Arizona: Chiricahua Mts., Bare- foot Park, Oct. 19, 1906, Blumer 1450 (F, G, NY, US); Washall Gulch, Santa Catalina Mts., Oct. 2, 1917, Shreve 5397 (F, G). CALIFORNIA: Avalon, Santa Catalina Island, Mar., 1897, Trask (US); Near Forest Ranch, Sierra Foothills, Butte Co., Sept. 23, 1916, A. A. Heller 12660 (F, G, NY, US); Plains of Mendocino, Aug. 10, 1882, Pringle (F, G, NY, US). Овесох: Portland, June 25, 1920, Fisher 14 (С) ; Salem, 1871, E. Hall 678 (F, G). Wasu- INGTON: Wenatchee, Chelan Co., July 6, 7, 1916, Eggleston 12930 (F) ; Seattle, King Co., Sept. 19, 1931, J. W. Thompson 8106 (G).
Mexico.—Baga CALIFORNIA: Sierra San Pedro Martir, 1923, Gallegas (US). Cninvanva: Majalea, Aug. 20, 1935, LeSuer Мех-498 (F); Southwestern Chihuahua, Aug.—Nov., 1885, Е. Palmer 447 (б, US); Madera, May 27-June 3, 1908, E. Palmer 288 (F, G, NY, US); Majarachic, April 24, 1938, Knobloch
19411 Braun,—The Red Azalea of the Cumberlands 3l
5114 (F). DunmawNco: Metates, north of Cueva, Aug. 29—30, 1934, Pennell 18410 (US).
(То be Continucd)
WALKER PRIZE IN Economic Borany.—Original unpublished essays on any subject in the field of plants useful to man are eligible for the Walker Prize competition for 1941. Further in- formation may be had from the Secretary, New England Mu- seum of Natural History, 234 Berkeley Street, Boston, Massa- chusetts. "The closing date is May 1, 1941.
ТНЕ RED AZALEA OF THE CUMBERLANDS E. Lucy BRAUN
Кок a number of years the writer has been referring to an Azalea of the Cumberland Mountains and Cumberland Plateau under the name “ red azalea,"! for want of something more spe- cific. W. Н. Camp also mentions “red azalea.”2 In Small’s flora, the statement is made that “a red azalea native in the Cumberland Mountains . . . may be distinet.'? To refer this plant to Rhododendron calendulaceum is to ignore its outstanding differences from that species. The red azalea of the Cumberlands is sufficiently different from А. calendulaceum as to be readily distinguished at any season. It also differs in habitat and in time of bloom from that species. R. calendulaceum in south- eastern Kentucky blooms at the beginning of May. "The natives there call it ^ honeysuckle " or “ yellow honeysuckle.” When the writer first saw bushes of the red azalea, in August, 1931, on the summit of Black Mountain in Harlan County, Ky., and inquired of a mountaineer as to the color and time of bloom, the reply was that the flowers are red and bloom in summer, about the first of July. They are not there confused with the “ yellow honey- suckle.” The height of the bloom-period is the end of June,
! Braun, E. Lucy, Vegetation of Pine Mountain, Kentucky. Amer. Midland Nat. 16: 517-565. 1935. An ecological transect of Black Mountain, Kentucky. Ecol. Mon. 10: 193-241. 1940.
?'The red azalea of Black Mountain, Kentucky. Jour. N. Y. Bot. Garden 37: 164—165.
1936. ? Small, J. K. Manual of the southeastern flora, p. 994. N. Y. 1933.
82 Rhodora [JANUARY
though in early seasons, bushes at lower elevations may bloom the middle of June. "There is, then, an interval of five weeks to two months between the bloom-periods of these two azaleas.
The hesitancy in assigning specific rank to the red azalea has been due to its resemblance to the flame azaleas of the southern Blue Ridge and Great Smoky Mountains. The great range in time of bloom and in relative maturity of leaves at blossom-time which is displayed by flame azaleas of the southern Blue Ridge province, especially the Great Smoky Mountains, where there is a succession of bloom from May to July, suggest that all are referable to one species, or perhaps are hybrids. This last pos- sibility is mentioned by Camp.*
In a region where the bloom-periods do not overlap, hybrids are impossible, and specific stability is maintained. In south- eastern Kentucky (in Letcher County) both R. calendulaceum and red azalea occur. Specimens of the former were collected in full bloom on May 6, 1934, at an elevation of about 2700 ft. in Joe Day Branch, Black Mountain. The flowers open when the leaves are unfolding and when some are about half-grown, but before the leafy shoots of the season have elongated. On the same day, and at approximately the same elevation, specimens of red azalea were collected. "The buds which produce the flower- clusters are still in winter condition. The winter buds which pro- duce leafy shoots are just beginning to open. On June 21, 1933, (a very early season) specimens of the two azaleas were collected on Pine Mountain, Letcher County, Ky., at about 2000 ft. eleva- tion. At that time, the capsules of R. calendulaceum were about half-grown, some 2 cm. long; the flowers of red azalea were just beginning to fade. These differences at the same date and eleva- tion are mentioned in order to emphasize the improbability of hybridization in this region, hence the maintenance of specific dis- tinctness of the two azaleas. In a region where only one of these azaleas occurs, a pure strain should persist.
At the western edge of the Cumberland Plateau in southern Kentucky, the red azalea occurs, but not the better known flame azalea. This is a region of isolated occurrences of ancient species, some coastal plain, some southern Appalachian.” The red azalea
i Camp, W. Н. On Appalachian trails. Jour. N. Y. Bot. Garden, 37: 249-265. 1936. 5 Braun, E. Lucy. Some relationships of the flora of the Cumberland Plateau and Cum-
19411 Braun,—The Red Azalea of the Cumberlands 33
of this section may well represent an ancient species which here has remained distinet; hence a specimen from this area is desig- nated as the type of Rhododendron cumberlandense to be de- scribed here. In the southern Blue Ridge province it has hy- bridized freely resulting in the apparently variable flame azalea of that section. A genetical study might help to establish the specific identity of the red azalea and the hybrid nature of the flame azalea of the southern Blue Ridge. Rhododendron calen- dulaceum of southeastern Kentucky and the red azalea of Ken- tucky both differ from flame azaleas collected on Standing Indian Mountain in the Nantahala National Forest of North Carolina on June 26, 1940. The flame azalea of the Nantahala region has characters of both of the Kentucky azaleas under consideration, and resembles the red azalea in the habit of flowering when the leaves are about mature.
In 1937, W. P. Lemmon described as Azalea Bakeri a yellow- to red-flowered species from northern Georgia.® The red azalea of the Cumberlands and Lemmon’s species have some features in common, suggesting that perhaps A. Bakeri is by hybridization related to the red azalea (here designated as R. cumberlandense). It appears to be a much larger-flowered species.
RHODODENDRON cumberlandense, sp. nov. Shrub with twigs sparsely strigose, winter buds yellow-brown, scales ciliate, mucro- nate, the outer scales aristate. Leaves obovate, 3-5 cm. long, glabrous except on midrib above and midrib and lateral veins below, margin ciliate. Flowers opening 5—6 weeks after leaves, mostly red; corolla 3.5—4 (rarely 5) em. broad; upper lobe broad, with large orange blotch. Filaments carmine, nearly glabrous. Capsule 2 cm. long, strigose.'—Cumberland Plateau and Cumber- land Mountains, Kentucky, in mesophytie oak woods. Type (in
writer’s herbarium) and isotype (in herbarium of Arnold Ar- boretum), Yahoo ridge, McCreary Co., Ky., June 15, 1935 (Braun,
berland Mountains in Kentucky. Rhodora 39: 193-208. 1937. А remarkable colony of coastal plain plants on the Cumberland Plateau in Laurel County, Kentucky. Am. Mid- land Nat. 18: 363-366. 1937.
8 Lemmon, W. P. Notes on a study of the southeastern azaleas with descriptions of two new species. Bartonia, no. 19: 14—17. 1987.
7 RHODODENDRON cumberlandense, sp. nov., frutex, ramulis sparse strigosis, gemmis flavo- brunneis, paleis ciliatis mucronatis, exterioribus aristatis; foliis obovatis 8—5 cm. longis, costa supra costa venulisque lateralibus infra exceptis glabris, margine ciliatis; floribus 35-40 diebus post maturitate foliorum dehiscentibus; corollis 3.5-4 (rarius —5) cm. latis, lobo superiore lato, macula magna aurantiaca ornato; filamentis rubris fere glabris; capsulis 2 em. longis. Typus (in herb. scriptoris) ad Yahoo Ridge, Comitate McCreary, Kentucky, June 15, 1935, collectus, Braun 971.
34 Rhodora [JANUARY no. 971). Specimens in writer’s herbarium from Flat Rock, McCreary Co.; 10 miles east of London, Laurel Co.; Peabody, Clay Co.; Stony Fork of Leatherwood, Perry Co.; Buck Branch, Whitley Co.; Pine Mountain, Letcher Co.; Black Mountain, Letcher Co. Most abundant on the summit of Black Mountain in Letcher and Harlan counties; also on Log Mountain in Bell Co. A tabular comparison of the characters of R. calendulaceum (the early spring blooming plant of the Cumberland Mts., not the apparent hybrids of the southern Blue Ridge) and of R. cumber- landense will emphasize the characters of the latter species.
R. calendulaceum
А shrub of dry southerly slopes, growing in company with other heaths.
Loosely branched, straggly; leaves more or less evenly disposed along the twigs of the season.
Young twigs and petioles strigose, with dense fine pubescence between the coarse hairs.
Young winter buds (June) vellow, scales ciliate, the outer scales pubescent, mucronate.
Leaves bright green, 5-7 cm. long, 2.5-3 cm. wide, broadest in the middle or sometimes above, sparsely strigose above and below, densely so near the leaf margins and on veins beneath; midrib be- neath with dense fine pubescence between the coarse hairs.
Flowers opening with the leaves.
Corolla 5 cm. or more across, yel- low to orange; the upper corolla- lobe broader than the lateral.
k. cumberlandense
А shrub of more mesophytie (usu- ally uae? woods, often the only eath.
More compactly branched; leaves more crowded toward tips of twigs.
Young twigs sparsely strigose, otherwise glabrous or nearly so.
Young winter buds (June) yellow- brown, glabrous, scales ciliate, mu- cronate, the outer aristate, outer- p with awn as long as body of scale,
Leaves dark green, mostly 3-5 cm. long, occasionally 7 cm., 1.5-2 or sometimes 2.5 cm. wide, broadest above the middle; glabrous above except for short fine whitish pubes- cence along the midrib, glabrous below except on midrib and larger lateral veins which are finely pu- bescent and sparsely strigose; mar- gins ciliate, but without the stri- gose band near margin.
Flowers opening 5-6 weeks after the leaves, i.e. when leaves are about mature.
Corolla 3.5-4 сш. or sometimes 5 cm. across, prevailingly red, but ranging through all the nasturtium colors; the upper corolla-lobe broader than laterals, sometimes 2x, almost orbicular but con- tracted to a short acuminate tip, with an orange-yellow blotch oc- cupying most of the area.
1941] Fernald,—Geranium nemorale, var. Bicknellii 35
R. calendulaceum
Corolla-tube loosely and coarsely glandular-pubescent, scattered glandular hairs continuing in a band almost to the apex of corol- la-lobes in the midrib region.
Style and filaments orange shad- ing to yellow about half-way to tip.
Filaments conspicuously pubes- cent for 1-2 cm. beyond throat of corolla tube.
Calyx-lobes short-ovate, sparsely hirsute, ciliate.
Capsules narrowly elongate, 2.5 cm. long, strigose, with pedicels glandular-pubescent.
R. cumberlandense
Corolla-tube pubescent with short glandular and non-glandular hairs, the glands, almost sessile, con- tinuing in a band almost to apex of corolla lobes.
Style and filaments carmine.
Filaments glabrous or sparsely pu- bescent for about 1 em. beyond throat of corolla tube.
Calyx-lobes similar. Capsules broader and shorter, 2
em. long, strigose, with pedicels strigose.
The smaller and more brilliantly colored flowers, the summer blooming even at low elevations (1200-1300 ft. at the western edge of the Cumberland Plateau), the small, nearly glabrous leaves, and brown, glabrous winter-buds with awned scales will generally distinguish this azalea from R. calendulaceum.
THE UNIVERSITY OF CINCINNATI,
Cineinnati, Ohio.
GERANIUM NEMORALE Suksd., var. Bicknellii (Britton), comb. nov. C Bicknellii Britton in Bull. Torr. Bot. Cl. xxiv. 92 (1897).
When I published a study of Geranium carolinianum and Allies in northeastern North America, in Ruopora, xxxvii. 295- 301 (1935), I was guilty of a lapse, due to long familiarity with the name C. Bicknell. I there showed that there are no specific distinctions but good varietal characters separating the more 'astern plant (C. Bicknellii) from the more western G. Bicknellii, var. longipes (Wats.) Fern. in Кнорока, 1. e. 297 (1935). Under var. longipes I included as a synonym G. nemorale Suksd. in Deuts. Bot. Monats. xvi. 222 (1892), being then quite blind to the fact that Suksdorf’s binomial antedated that of Britton by five years! My attention was most kindly directed to this
error a year ago by Mr. S. J. Smith, who suggested that I make
36 Rhodora [JANUARY
the eorreetion. Since the point was his discovery, I have urged that he clarify it; but, as he has modestly refrained from doing so in print, I am making the needed correetion.—M. L. FERNALD.
LACHNANTHES IN Nova Scoria.—In the course of some recent motoring in Nova Seotia, my wife and I traversed an area in Queen's County, northwest of Liverpool, whieh had not been visited either by the Gray Herbarium Expedition of 1920 or by Messrs. Fernald and Long in the following year.’ Shortly before reaching it, we had chanced upon the station for Lophiola at Faney Lake? and our appetite for collecting had been whetted thereby. Accordingly, we made stops, rather brief but as long as our schedule allowed, in a patch of red-oak woods, on a pebbly beach of Ponhook? Lake, and at another point on the same lake where the strand was composed of solidly compacted black peat, cut off so sharply at the water’s edge as to form there a miniature cliff a foot or two high.
The woods yielded nothing of interest. The pebbly beach had Aster Tradescanti L. (A. vimineus var. saxatilis Fern.) somewhat farther east in Nova Scotia than recorded by Prof. Fernald. The peaty strand rewarded us with Lachnanthes tinctoria (Walt.) Ell., apparently new to Nova Scotia and to Canada. The nearest known stations are in southeastern Massachusetts.
This adds one more to the considerable list of coastal-plain species with similarly interrupted ranges known from Nova Scotia and suggests that Prof. Fernald's propheey as to the increased proportion of such species in the flora of the province which fur- ther collection would reveal, might easily be fulfilled.—C. A.
WEATHERBY, Gray Herbarium.
1 Ruopona, xxiii. 90ff. (1921); xxiv. 157ff. (1922).
2 RHopons, xxiv. 167.
3 So spelled on the official highway map of Nova Scotia; “Penhook” in Rand-MeNally’s Atlas.
4 RHopona, xxiii. 168.
Volume 42, no. 604, including pages 503-650 and the title-page of the volume, was issued 17 December, 1940.
Hovova
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Conducted and published for the Club, by MERRITT LYNDON FERNALD, Editor-in-Chief
CHARLES ALFRED мшезш LUDLOW GRISCOM Associate Editors STUART KIMBALL HARRIS f Vol. 43. February, 1941. No. 506. CONTENTS: Revision of the Genus Pteridium (concluded). R. M. Tryon, Jr. 37 An old Forest in Stonington, Connecticut. Hugh M. Raup. .... 67 Remarks on the Name Phlox nivalis. Edgar T. Wherry. ........ l Reappearance of rare New England Marine Algae. Wm kandolph Taylora se E ИЛ EE ru M CM 72 Nymphaea tetragona in Somerset County, Maine. Санет ы. шл E E т CS 74 Deam's Flora of Indiana (Review). C. A. Weatherby. ......... 75 New Form of Bromus inermis. John W. Moore ................ 76
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Gray Herbarium of Harvard University, Cambridge, Mass.
Rhodora
Plate 653
MU
NU He
E
INS
TS
Fia. 1, upper half of middle pinna of PrERIDIUM
X lo; FIG. 2, basal pinna of sm: all | pl int of var. YARRABENSE, X 10; FIG.
of var. ARACHNOIDEUM, X lá; F
‚ 4, tip of frond of var. CAUDATUM,
AQUILINUM, Var. ESCULENTUM,
3, upper pinna x М.
IRbodora
JOURNAL OF
THE NEW ENGLAND BOTANICAL CLUB
No. 506.
Vol. 43. February, 1941.
A REVISION OF THE GENUS PTERIDIUM R. M. Tryon, JR.
(Continued from page 31)
4. PrERIDIUM AQUILINUM var. Ferr (Schaffn. ex Fée) Maxon ex Yuncker, Field Mus. Pub. Bot. 17: no. 4, 308 (1938). PLATE 650, FIG. 4, PLATE 651, FIG. 1, МАР 7.
Pteris aquilina L. var. pubescens Kze. Linnaea 13: 142 (1839), as to plant, not as to basinym, Pteris lanuginosa Spreng. Pteris aquilina L. var. pubescens Spreng. ex Liebm. Vid. Selsk. Skr. s. 5, 1: 225 (1849), as to plant, not as to basinym, Pteris lanuginosa Bory ex Willd. Pteris Feei Schaffn. ex Fée, Mém. Fam. Foug. 8: 73 (1857). Pteridium Feei (Schaffn. ex Fée) [combination incorreetly attributed to Maxon by] Faull, Contrib. Arn. Arb. 11: 87 (1938).
Pteris aquilina L. var. pubescens Spreng. ex Liebm. based on Pteris lanuginosa Bory ex Willd. and Pteris aquilina L. var. pubescens Kze. based on Pteris lanuginosa Spreng. are both earlier varietal names than var. Кее, and were originally applied to this variety, but their basinyms refer them, respectively, to var. typicum and var. latiusculum. Also, of course, they could not to be used under Pteridium because of var. pubescens Underw.
Growing tip of the rhizome with a tuft of dark hairs; frond 0.2-1 m., usually about 0.5—0.7 m. high, vernation subgleichen- 1011; stipe usually shorter than the blade; blade 1—5 dm., usually about 3 dm. long, usually broadly ovate or pentagonal, less often ovate or broadly triangular, not ternate, usually bipinnate- pinnatifid to tripinnate, less often tripinnate-pinnatifid; rachis usually slightly pubescent, sometimes strongly pubescent or
38 Rhodora [FEBRUARY
glabrate; pinnae and pinnules short-acuminate to obtuse; pin- nules usually nearly at right angles to the costa, sometimes at an oblique angle; costules slightly to moderately pubescent be- neath and less so above; penultimate segments pinnatifid, often pinnate, or pinnate-pinnatifid; longest entire segment or entire part of a segment from three to eight, usually about four, times as long as broad; ultimate segments usually straight, rarely sub- faleate, adnate or broadest at the base, the upper surface slightly to moderately pubescent, at least near the margin, the midnerve glabrous or slightly pubescent, the margin usually moderately pubescent, rarely glabrate, the lower surface usually densely pubescent, rarely slightly pubescent or pubescent only on the midnerve; fertile and sterile indusium ciliate and sometimes also pubescent on the outer surface, the fertile usually 0.3 mm. or more wide, the sterile usually 0.4 mm. or more wide, the fertile portion no broader than the sterile on the same segment.
Түрк: Schaffner 138, 141. Probably at Rio de Janeiro (not seen).
Tu Locanrrv: Huatusco, Mexico.
Some specimens of var. Feei are not entirely typical, having some character of var. pubescens : Heyde & Lux from Guatemala and Mohr from Vera Cruz, Mexico have the sterile indusium narrow, 0.2-0.3 mm. wide; Rose 2212, Tepie, Mexico has the sterile indusium only slightly ciliate; and Palmer 67, San Luis Potosi, Mexico has an ovate blade. Ortega 7400, Sinaloa, Mexico, 1934 (F) is intermediate between the two varieties.
Var. Feei may be separated from var. pubescens by its con- spieuously ciliate rather than only slightly, if at all, ciliate fertile indusium, the usually much more ciliate sterile indusium, which is also almost twice as broad, and the fertile portion of the indusium no broader than the sterile on the same segment rather than broader. On the average, the fronds are consider- ably smaller.
It differs from var. typicum in having the margin of the seg- ments pubescent rather than glabrous, the sterile indusium usually considerably broader, and in its usually smaller size. Its differences from var. decompositum, with which it is also closely related, are discussed under that variety.
Var. Feei grows in the mountains of Mexico, Guatemala and Honduras, up to 2800 m.
Mexico.—Sawn Luts Potosi: San Miguelito Mts., 1876, Schaff- ner 925 (С); Alvarez, Sept. 5—10, 1902, E. Palmer 67 (F, G, NY,
Tryon, Jr..—Revision of the Genus Pteridium 39
1941]
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‘(2 deur) ay "194 IKalvdavoodadsd вл (р dew) WAJdAL /
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came р
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40 Rhodora [FEBRUARY
US). Terr. Теріс: near Santa Teresa, Aug. 12, 1897, Rose 2212 (G, NY, US). Guanasuato: 1905, Duges 6 (US). Hiparco: Durango, Aug. 13, 1937, Fisher (NY, US) ; El Chico, July, 1927, Lyonnet 98 (G, NY); between Somoriel and Las Lajas, Aug. 5, 1905, Rose, Painter & Rose 9204 (NY, US). Vera Cruz: Huatusco, April, 1857, Mohr (U. S. Nat. Herb. no. 724103) ; Cordoba, 1889-91, Fink 18 (G, NY, US) ; near Jalapa, May 12, 1900, Pringle 8342 (US). Mexico: Oct. 1875, Schaffner 59 and 116 (NY); below Ajusco, Sept. 19, 1903, Rose & Painter 7214 (G, US). Puesta: Teziutlan, Sept. 7, 1910, Orcutt 4029 (US). Oaxaca: Cerro de San Felipe, Sept. 26, 1897, Conzatti & Gonzales 487 (С); Cuicatlan, June 16 and 22, 1898, Conzatti & Gonzales 747 (G, US).—GuvarEMALA: Between Solola and Chidueadenango, Aug. 13, 1936, Hatch & Wilson 322 (US); Chichavae, Chimaltenango, Nov.-Dec., 1930, Skutch 12 (US); Laguna de Avarza, Jalapa, Sept., 1892, Heyde & Lux (J. D. Smith no. 4080) (G, NY, US). Honpuras: Near Siguatepeque, Dept. Comayagua, July 3, 1936, Yuncker, Dawson & Youse 5600 (F, б, NY, US).
5. PrERIDIUM AQUILINUM var. decompositum (Gaud.), n. comb. Prats 650, Fic. 5, PLATE 651, ric. 2. Illustration: St. John & Hosaka, Weeds Pineapple Fields. Haw. Is., Univ. Haw. Res. Pub. 6, 24 (1932).
Pteris decomposita Gaud. in Freye. Voy. Bot. 393 (1829). Pteridium capense (Thunb.) Krasser var. decompositum (Gaud.) Nakai, Bot. Mag. Tokyo 39: 110 (1925).
Growing tip of the rhizome with a tuft of dark hairs; frond 0.5-2 m., usually about 0.7—1 m. high, vernation subgleichenioid; stipe usually shorter than the blade; blade 2-10 dm., usually about 4 dm. long, usually ovate or broadly ovate, not ternate, usually tripinnate or tripinnate-pinnatifid; rachis glabrous or slightly pubescent; pinnae and pinnules short-acuminate to ob- tuse; pinnules usually nearly at right angles to the costa, some- times at an oblique angle; costules slightly pubescent beneath and less so above; penultimate segments pinnatifid, pinnate or pinnate-pinnatifid; longest entire segment or entire part of a segment from three to five, usually about four, times as long as broad; ultimate segments usually straight, adnate or broadest at the base, the upper surface glabrous, rarely very slightly pubes- cent along the midnerve, the margin glabrous, slightly pubes- cent, or rarely quite pubescent, the lower surface usually densely subappressed-lanuginose pubescent, sometimes only slightly so; fertile and sterile indusium ciliate and sometimes also pubescent on the outer surface, rarely becoming glabrous with age, the fertile usually about 0.3 mm. wide, the sterile
19411 Tryon, Jr.,—Revision of the Genus Pteridium 41
usually about 0.2 mm. wide, the fertile portion broader than the sterile on the same segment, or no broader.
Type: Gaudichaud, in Herb. Muséum d'Histoire. Naturelle, Paris (not seen).
Typr LocaLiTY: Hawaiian Islands.
Var. decompositum is closely related to vars. Кее and pubes- cens. However, it has a nearly glabrous rachis rather than a pubescent one as in those two varieties and the upper surface of the segments is glabrous or rarely slightly pubescent only along the midnerve rather than pubescent and usually with a glabrous midnerve. The sterile indusium is only about half as broad as that of var. Feei and on the average it is not as large a plant as var. pubescens. The margin of the segments is some- times glabrous or nearly so rather than pubescent. The sub- appressed pubescence on the lower surface of the segments is characteristic though not always well defined.
Var. decompositum is a part of the small element of the Hawaiian flora that is related to the American flora.
It occurs exclusively in the Hawaiian Islands, where it grows along field borders, in virgin land, on craters, on bare eroded slopes, on open grassy slopes and in thickets from 300 m. up to 2700 m.
Hawaiian IstaNDs.—Kavar: June 22, 1895, A. A. Heller 2416 (F, G, NY, US). OaHv: Waianae Mis. Honouliuli, May 2, 1937, Fosberg 13810 (G); Koalau Mts., March 29, 1933, Fosberg 9320 (С); Wahiawa, June 3, 1909, Forbes (NY). Могокат: Kahuaawi, May 30, 1928, Degener 3558 (NY, US). Lawar: Kaokahi, Nov. 28, 1935, Fosberg 12429 (G). Maur: Aug 22, 1933, Fosberg 9943 (C). Hawan: Kilauea Bird Park, Aug. 5, 1925, Neal (NY).
6. PTERIDIUM AQUILINUM var, LATIUSCULUM (Desv.) Underw. ex Heller, Cat. N. Am. Pl. Ed. 3, 17 (1909). Plate 650, ric. 8, PLATE 652, ric. 1, МАР 8. Illustrations: Tilton, Fern Lover's Comp. 48, 49 (1922); Tryon et al., Ferns of Wis. 18, 19 (1940), habitat; Svensk Bot. 2: t. 90 (1803).
Pteris caudata L. sensu Schk. Krypt. Gew. 88 (1809), in part, pl. 96b, a. Pteris ciliata Willd. ex Schk. Krypt. Gew. 89 (1809), in synonymy. Pteris lanuginosa Spreng. Nova Acta 10: 231 (1821). Pteris Sprengelii Steud. Nom. Bot. 2: 358 (1824). Pteris latiuscula Desv. Mém. Soc. Linn. 62: 303 (1827). Pteris aquilina L. var. pubescens Kze. Linnaea 13: 142 (1839), as to
[FEBRUARY
Rhodora
42
‘(QT dew) я$мяяуннул * (6 dem) XALNWIAOSS4 “IBA :(g dew) WOATAOSAILVI IBA ипмгипбу ия jo saduvy
19411 Tryon, Jr.j—Revision of the Genus Pteridium 43
basinym, Pteris lanuginosa Spreng., not as to plant. Pteridium latiusculum (Desv.) Hieron. ex Fries, Wiss. Ergebn. Schwed. Rhodesia-Kongo Exp. 1': 7 (1914). Cincinalis latiuscula (Desv.) Vict. Contrib. Lab. Bot. Univ. Montréal no. 2; 41 @1923); nomen provisorium. Pteridium aquilinum (L. Kuhn var. japonicum Nakai, Bot. Mag. Tokyo 39: 106 (1925), ex char. Pteris latiuscula lanuginosa Small, Ferns N. Ү. 241 (1935), in synonymy. Pteridium latiusculum (Desv.) Hieron. ex Fries var. verum Wherry, Am. Е. Journ. 27: 58 (1937). Pteridium aquil- inum (L.) Kuhn f. glabrum Tardicu-Blot and C. Chr. in Lecomte, Fl. Gen. Indo-Chine 72: 138 (1939). Pteridium japonicum (Nakai) Tardicu-Blot and C. Chr. in Lecomte, Fl. Gen. Indo- Chine 77: 138 (1939), in synonymy.
The earhest varietal name, Pteris aquilina L. var. pubescens Kze., cannot be transferred to Pteridium aquilinum because of Pteridium aquilinum var. pubescens Underw.
Growing tip of the rhizome usually naked, or with a few whitish hairs, rarely with a tuft of dark hairs; frond 0.3-1.5 m., usually about 0.5—1 m. high, vernation equal; stipe longer or shorter than the blade; blade 2-8 dm., usually about 5 dm. long, usually broadly triangular, rarely broadly ovate or ovate, ofteu ternate, usually tripinnate or tripinnate-pinnatifid, sometimes bipinnate-pinnatifid; rachis usually glabrous or subglabrous, sometimes slightly pubescent; pinnae and pinnules subacute to obtuse; pinnules usually at an oblique angle to the costa, rarely nearly at right angles; costules slightly pubescent beneath and less so above, or glabrous; penultimate segments usually pinnate or pinnate-pinnatifid; longest entire segment or entire part of a segment from three to seven, usually about four, times as long as broad; ultimate segments usually straight, adnate or broad- est at the base, the upper surface glabrous or subglabrous, the margin pubescent, or rarely subglabrous, the lower surface usually pubescent only along the midnerve, rarely shghtly pubescent between the margin and the midnerve; fertile and sterile indusium usually glabrous, rarely the fertile slightly pubes- cent on the outer surface or ciliate, and the sterile slightly ciliate, the fertile 0.25-0.4 mm. wide, the sterile 0.1-0.2 mm. wide, the fertile portion broader than the sterile on the same segment.
Type: Sheet labeled Pteris latiuscula Desv., Herb. Desvaux in Herb. Muséum d'Histoire Naturelle, Paris (not seen). Photo- graph of type in U. S. National Herbarium and Gray Herbarium (seen).
TYPE LocaLity: Newfoundland and St. Pierre.
44 Rhodora [ FEBRUARY
In northern Europe, Kamtchatka and occasionally through- out its range in North America, plants of var. latiusculum occur that have the sterile indusium slightly ciliate and the lower sur- face of the blade somewhat pubescent between the margin and the midnerve. Such plants in North America are discussed under var. pubescens. Also, occasionally, the blade is ovate rather than broadly triangular. These are apparently normal variations in any large population of var. latiusculum.
In northern Wisconsin and adjacent Michigan, and perhaps more widely distributed, plants with an ovate blade, pubescent beneath between the margin and midrib, and with the sterile indusium ciliate are not uncommon (PLATE 652, ric. 4). Repre- sentative specimens are: Boulder Junction, Vilas Co., Wisconsin, July 3, 1938, Tryon 3914 (G); Hersey, Osceola Co., Michigan, June 25, 1938, Fassett 19244 (С); Northwest of L’Anse, Baraga Co., Michigan, Fassett 19251 (G). They constitute a rather noticeable proportion of the var. latiusculum population. An attempt to identify such plants led me into this study of Pteridium but I am still unable to give a satisfactory interpreta- tion of them. In the summer of 1940 I made an effort to study them in the field more closely than I had in 1938? but heavy late frosts had killed or mutilated most of the Bracken. They may be regarded as a scattered population intermediate between var. pubescens and var. latiusculum, closely related to the former in the characters given above but, I believe, derived from the latter by rhizomes or spores. Or, there is considerable evidence that they are merely the result of adverse growing conditions such as burning, pasturing, and extremes of exposure and soil sterility. They are found in especially dry, sunny places, often in pastures, fields, railroad rights-of-way and recently burnt- over land.
There is a certain amount of intergradation between var. latiusculum and var. Wightianum. Some specimens with the leaf-cutting of var. latiusculum are slightly pubescent beneath between the margin and the midnerve and have the sterile in- dusium slightly ciliate, while others have tapering pinnules set at right angles to the costa, as in var. Wightianum, and are pubescent beneath only on the midnerve. Such intermediates
20 See Tryon, Notes on the Ferns of Wisconsin. Am. F. Journ. 29: 1 (1939).
1941] Tryon, Jr.,—Revision of the Genus Pteridium 45
are: Kwangtung, China, Jan. 4, 1928, Tsang 16704 (F); Kwangtung, China, Lau 2353 (G); Canton, China, 1874, Рой (Herb. Field Mus. no. 593622); near Kau Fung, Loh Ch’ang Dist., Kwangtung, China, Nov. 2-30, 1932, Tsang 20872 (NY, US); Foochow, Fukien Prov., China, Metcalf 7406 (US); Wang Shan, Anhwei Prov., China, Aug. 28, 1923, Ip (US); Mt. Reng- ger, Java, Sept. 25, 1907, Buysman (US).
Var. latiusculum may be separated from var. Wightianum by its equal rather than gleicheniold vernation, its broadly tri- angular and ternate rather than ovate-triangular and evenly pinnate blade, its nearly glabrous rather than densely pubescent rachis and its subacute or obtuse rather than long-acuminate pinnae and pinnules. Also the ultimate segments are straight rather than faleate, the lower surface 1s glabrous except along the midnerve rather than densely pubescent, the fertile and sterile indusium are glabrous or nearly so rather than quite ciliate and the pinnules are at an oblique angle to the costa rather than at right angles.
Vars. typicum, pubescens and pseudocaudatum are also closely related to var. latiusculum and the critical differences are dis- eussed under their treatments.
Var. latiusculum, in eastern North America and eastern Asia, is another example of the well-known relationship of the floras of those two areas. The localities in the Black Hills of South Dakota and the mountains of Wyoming, Colorado and Nuevo Leon undoubtedly represent relies of а once continuous range, the intervening population perhaps having been wiped out by aridity in the Great Plains Region. I do not have enough data at hand to interpret the occurrence of var. latiusculum in north- ern Europe. It may have survived glaciation in local nunatak areas in Seandinavia and spread since the disappearance of the ice, or it may have spread, since glaciation, westward from un- glaciated areas in western Siberia. Var. latiusculum is probably more widely distributed in central Asia than Map 8 indicates. Several of the localities in central Asia on Map 1 probably represent var. latiusculum.
Var. latiusculum grows in pastures, open woods, thickets, on open slopes, in woods, on grassy slopes in abandoned fields and in burnt-over areas, in damp or more often dry, usually sterile
46 Rhodora [FEBRUARY
soll; from sea level up to 1500 m. in eastern North America, 2300 m. and 2700 m. in the mountains of Wyoming and Colorado and up to 2000 m. in China.
It ranges from Newfoundland to Minnesota, south to Okla- homa and Tennessee; isolated in Mississippi, Wyoming, South Dakota, Colorado and Nuevo Leon; Sweden south to Germany, east to western Russia; Siberia; Kamtchatka to Amur, south to Formosa, Hainan and Szechuan.
М№овтн AMERICA.—SAINT Pierre: Cape Noir, July 10, 1901, Arsène 6 (G); July 10, 1909, Arsène 5 (NY). Пе MIQUELON: July 27, 1882, Delamare 364 (NY). NEwrouNDpLANp: Holy- rood, South Arm River, Aug. 23, 1894, Robinson & Schrenk 4 (F, G, NY, US); Grand Falls, July 10, 1911, Fernald & Wiegand 4281 (G). Dominion or Canapa. QuEnBEC: Boishébert, Mut- ton Bay, Saguenay Co., Aug. 15, 1915, Н. St. John 90010 (О); Seven Islands, Saguenay Co., Aug. 12, 1907, C. B. Robinson 873 (NY); Bie, Rimouski Co., Aug. 15, 1927, Rousseau 26884 (С); Northwest of 'Three Rivers, Champlain Co., Aug. 1, 1923, Cham- berlain & Knowlton (G); Grindstone, Grindstone Island, Mag- dalen Islands, Aug. 23, 1912, Fernald, Long & St. John 6645 (G). Prince Epwarp Isand: Dundee, Kings Co., Aug. 26, 1912, Fernald, Long & St. John 6646 (G). New Brunswick: Shediac Cape, July 23, 1914, F. T. Hubbard (G). Nova Scotia: Brazil Lake, Yarmouth Co., July 16, 1921, Bartram & Long 23003 (G) ; Digby, July 2-7, 1901, Howe & Lang 258 (G, NY). ONTARIO: Moore Lake, Bruce Peninsula, Aug. 26, 1934, Krotkov 9606 (G, US) ; east end of Timagami Lake, Timagami Provincial Forest, Aug. 4-11, 1935, E. C. & T. С. Yuncker 5499 (F) ; Laurier, Parry Sound District, Aug. 13, 1905, Moyer (NY). UNITED STATES OF AMERICA. Marne: Mt. Katahdin, July 14, 1900, Fernald (G); Boundary Lake, Aroostook Co., Aug. 12, 1902, Eggleston & Fernald (С). New Hawrsurgk: Randolph, Sept. 1, 1903, A. H. Moore 1454 (G); Jaffrey, July 25, 1897, B. L. Robinson 287 (С). VrgnMOoNT: Manchester, July 27, 1898, Day 260 (G); Brandon, May 23, 1908, E. F. Williams (G). Massacuuserts: Sharon, July, 1905, S. F. Poole 307 (G); Tisbury, Martha's Vineyard, June 16, 1917, F. C. Seymour 1001 (G, US) ; Granville, Sept. 19, 1913, Seymour 60 (NY). Кноре IsrawNp: Barrington, Bristol Co., Sept. 15, 1906, M. H. Grant (G). CowwEcrICUT: North Guilford, Sept. 30, 1906, G. H. Bartlett (G) ; Southington, Sept. 20, 1898, Bissell 880 (G). New York: Ithaca, Tompkins Co., Sept. 3, 1914, Metcalf 1405 (G); Staten Island, July 16, 1906, Dowell 4506 (G, US). New Jersey: Budd’s Lake, Sussex Co., Aug. 12-14, 1890, Small (F). PENNsvLvANIA: Wissahickon
19411 Tryon, Jr.—Revision of the Genus Pteridium 47
Ravine, Philadelphia Co., July 19, 1924, Lang 626 (С); Reading, Berks Co., Sept. 11, 1929, E. J. Palmer 36311 (G). MARYLAND: between Oakland and Thayerville, Garrett Co., July 5, 1913, Tidestrom 6457 (G); Cumberland, 1894, Shriver (NY). Dıs- TRICT OF CoLuMBiA: Terra Cotta, June 17, 1888, Holm (С). West VinGiN1A: White Sulphur Springs, Greenbrier Co., May 14-17, 1914, Hunnewell (G); Whitmer, Randolph Co., Sept. 13, 1904, A. H. Moore 2221 (Сб). Vernia: Bull Run Mts. Fau- quier Co., June 9, 1935, Allard 598 (G, NY); Marion, Smyth Co., June, 1892, Britton, Britton & Vail (NY). NoRrH Caro- LINA: near Waynesville, Sept. 5, 1910, Standley 5529 (US). MicHiGAN: 4 miles northwest of Calumet, Houghton Co., July 24, 1936, Hermann 8264 (NY); Ludington, Mason Co., Sept. 17, 1910, Chaney 256 (F, G, US). Onto: Berea, July, 1897, Ash- croft (Herb. Field Mus. nos. 140093, 140094) ; Hiram, Portage Co., Aug. 15, 1897, Webb 265 (С). IwxpiaNa: Millers, Lake Co., July 7, 1908, Lansing 2759 (F, G); Lake Oliver, July 16, 1933, Shoop (Herb. Field Mus. nos. 907912, 907922). KENTUCKY: Burnt Bridge Ridge, Madison Co., July 7, 1937, Smith, Hodgdon & Brown 3625 (G); Pine Mt., Bell Co., Sept., 1893, Kearney (US). TENNEssEE: South of Craggie Hope, Cheatham Co., Aug. 20, 1922, Svenson 342 (G); Henderson, June, 1892, Bain (NY). MississiPPr: Biloxi, June 3, 1898, PT acy, эЛЕ eNOS) Wisconsin: Delavan, July 13, 1919, Hollister 146 (G, US); Solon Springs, Douglas Co., Sept. 7, 1930, Somerville 41 (G). Пллхоѕ: Starved Rock, La Salle Co., sept. 7, 8, 1914, Lansing 3786 (Е) ; Joliet, Sept. 20, 1904, Skeel 549 (Е); Pine Hills, Union Co., May 6, 1902, Gleason 2899 (G). МіххЕѕотА: St. Cloud, July, 1896, Campbell (F); Itasea Park, Clearwater Co., July 16, 1933, Mayle 654 (G, NY). Iowa: Fayette Co., July, 1894, B. Fink 444 (G, US); Lebanon, July 5, 1897, Sample 502 (G, US). Missovnr: Monteer, Oct. 24, 1907, Bush 1146 (G, NY, US); Ironton, June 23, 1897, Savage & Stull 328 (F). ARKAN- sas: Jasper, Newton Co., June 18, 1932, D. M. Moore 32503 (С). SoutH Dakora: Custer, Black Hills, Aug. 19, 1892, Rydberg 1192 (NY, US); Pinecrest Camp, Deadwood, 1927, Haywood 1115 (F). Wxvowiwc: Jackson's Hole, Lincoln Co., Aug. 11, 1920, L. B. & E. B. Payson 2275 (G); Laramie Peak, Albany Co., July 10, 1900, Nelson 7518 (С, NY). Согоклро: Rabbit Ear Range, Routt Co., July 18, 1903, Goodding 1595 (G, NY, US) ; Brush Creek, Sept. 9, 1910, Tidestrom 4166 (б).—МЕхтсо: Sierra Madre Mts., Monterey, Nuevo Leon, July 6, 1933, C. H. & M. T. Mueller 366, in part (G).
Europr.—Norway: Bygdo, June 28, 1907, O. Anderson (US). SWEDEN. (STOCKHOLM): near Stockholm, Sept. 18, 1887, A. F. Carlson (US). ÓsrERGÓTLAND: Aug. 8, 1915, А. O. Olson (Herb.
48 Rhodora [FEBRUARY
Field Mus. no. 821335) ; Ljushult, July 21, 1911, A. O. Olson (NY); Asunden, Aug. 17, 1915, A. O. Olson (NY). KALMAR (SMALAND): Kalmar, 1882, Linddorff (G).—FiNLAND: Nyland, Aug. 20, 1908, H. Lindberg 409 (NY). Germany: near Berlin, July, 1844, Gausauge (G); Hanau, Sept., 1910, Peipos (Herb. Field Mus. no. 756316); Nürnberg, Bavaria, Aug. 15, 1910, Honig (G). CzkEcnosrovakiA: Moravia, Sept. 13, 1925, J. Bily 103 (С). Оміом or Soviet SocrarnisT REPUBLICS: St. Petersburg, 1860 (Herb. Field Mus. nos. 29377, 162025).
ASIA.—SIBERIA. Tomsk: near Titovka, Aug. 29, 1928, Proto- popova (G). KamTcHatKa: Savoiko, Aug. 29, 1928, Eyerdam (G, NY, US); Petropavlovsk, Aug. 6, 1928, Eyerdam (F, G, NY, US). Primorsk: Vladivostok, May—Oct., 1919, Top- ping 2343 (US). Amur: Blagowjeschtschensk, 1906, Karo (G, US).—SacHaniN: 1872, Augustinowicz (G). Japan: Maries (U. S. Nat. Herb. no. 022422) ; June, 1896, Halbrook 40 (NY); Atago, Oct. 13, 1894, Stanford (Herb. Field Mus. no. 825006) (NY); Sakamoto, Aug. 8, 1929, Dorsett & Morse 897 (US); Kano San, Kadsusa, Sept., 1888 (U. S. Nat. Herb. no. 22432) ; Mt. Kano San, Kadsusa, Sept. 7, 1908 (U. S. Nat. Herb. no. 1095365); Yase near Kyoto, June, 1921, Husimi (U. S. Nat. Herb. no. 1704754) ; Nanokawa, Tosa, July 3, 1892 (U. S. Nat. Herb. no. 22439). Formosa: Hancock 56 (US). Cuna. MawvcHnunia: 1931, Chen 494 (NY). Снїнїл: Tungling Mts., May 18, 1921, Cowdry 1214 (US). KrawNc Su: Poa Wha Mt., Chu-Yung, Oct. 10, 1915 (U. S. Nat. Herb. no. 1094030). CHEKIANG: Mo Kan Shan, June 28, 1926, Cheo & Wilson 12663 (G). Anwer: Chiu Hua Shan, June 28, 1925, Ching 8478 (G, NY). Huren: 1885-88, Henry 3146 (G); Wuchang, June, 1932, Chung 9058 (F). KraNcsr: Lu Shan, Sept. 19, 1922, Steward 2724 (US). Kwanerune: Hong Kong, 1853-56, Wright (G, US). Hamax: Ka Chik Shan, April 25, 1933, Lau 1637 (С). Yunnan: Ping-pien Hsien, June 5, 1934, Tsai 60128 (С). SzecHuan: Mt. Omei, Omei Hsien, 1928, Fang 3034 (G), 3231 and 3317 (G, US); Nanchuan Hsien, 1928, Fang 5841 (Q).— FnENcH INpo-Cutna: Nov., 1921 (U.S. Nat. Herb. no. 1505970).
7. PTERIDIUM AQUILINUM Var. PSEUDOCAUDATUM (Clute) Heller, Cat. N. Am. PI. Ed. 2, 12 (1900). PLATE 650, FIG. 7, PLATE 652, FIG. 2, MAP 5. Illustration: Blomquist, Ferns of N. Car. 42 (1934).
Pteris caudata L. sensu Schk. Krypt. Gew. 88 (1809), in part, Pl. 96b, b. Pteris novae-angliae Bory ex Hook. Sp. Fil. 2: 197 (1858), in synonymy. Pteris aquilina L. var. pseudocaudata Clute, Fern Bull. 8: 39 (1900). Pteridium aquilinum pseudo- caudatum (Clute) Clute, Fern Bull. 8: 39 (1900), nomen pro-
19411 Tryon, Jr..—Revision of the Genus Pteridium 49
visorium. Pteris pseudocaudata (Clute) Anon. in Index, Proc. Biol. Soc. Wash. 14: 200 (1901). Pteris latiuscula pseudo- caudata (Clute) Clute, Fern Bull. 11: 62 (1903), nomen pro- visorium. Pteridium — latiusculum pseudocaudatum (Clute) Maxon, Am. F. Journ. 9: 44 (1919). Filix-foemina aquilina (L.) Farwell var. pseudocaudata (Clute) Farwell, Am. Mid. Nat. 12: 290 (1931). Pteris latiuscula Desv. var. pseudocaudata (Clute) E. P. St. John, Am. F. Journ. 25: 40 (1935).
Growing tip of the rhizome usually with a tuft of dark hairs; frond 0.3-1.5 m., usually about 0.5-1 m. high, vernation equal; stipe longer or shorter than the blade; blade 2—7 dm., usually about 5 dm. long, usually broadly triangular, rarely broadly ovate or ovate, sometimes ternate, bipinnate-pinnatifid or tri- pinnate, rarely tripinnate-pinnatifid; rachis glabrous; pinnae and pinnules acute to obtuse; pinnules usually at an oblique angle to the costa, rarely at right angles; costules glabrous or less often slightly pubescent; penultimate segments pinnatifid or pinnate, rarely pinnate-pinnatifid; longest entire segment or entire part of a segment from six to fifteen, usually about nine, times as long as broad; ultimate segments usually straight, ad- nate or broadest at the base, the upper surface glabrous, the margin usually glabrous, rarely slightly pubescent, the lower surface glabrous, or sometimes pubescent along the midnerve; fertile and sterile indusium glabrous, the fertile 0.3-0.4 mm. wide, the sterile 0.1-0.2 mm. wide, the fertile portion broader than the sterile on the same segment.
Type: Clute 339, isotype in Herb. New York Botanical Garden (seen).
ТҮРЕ Locanrrv: Babylon, Long Island, New York.
It is interesting to note that in 1899, one year before Clute described var. pseudocaudatum, Maxon identified Ball 511 as "Pteris aquilina L. var. nov." “not typical—approaching P. caudata Linn."
Var. pseudocaudatum intergrades to a considerable extent with var. latiusculum. Intermediate specimens have the leaf- cutting of var. latiusculum but are nearly glabrous or have the leaf-cutting approaching var. pseudocaudatum and are either glabrous or have a pubescent margin and midnerve. Such specimens are: Hammonton, New Jersey, May 30, 1919, Killip 2260 (US); Bladensburg, Maryland, July 31, 1919, Mazon 6461 (G); Table Rock, North Carolina, June, 1879 (Herb. Field Mus. no. 315115, С. S. Nat. Herb. no. 22450); near White Sulphur Springs, Greenbrier Co., West Virginia, Aug. 29, 1903, Mackenzie
50 Rhodora [FEBRUARY
381 (NY); Henderson, Tennessee, June, 1892, Bain 162 (G); Wasioto, Bell Co., Kentucky, Sept., 1893, Kearney (NY); Bowl- ing Green, Kentucky, July, 1891, Price (NY). Lansing 518, West Pullman, Illinois, Sept. 18, 1898 (F) approaches var. pseudocaudatum. This strongly suggests that typical var. pseudo- caudatum occurs at the southern tip of Lake Michigan, and it has been reported from the Dunes Region,” but I have not seen any specimens.
Var. pseudocaudatum may be separated from var. latiusculum by the glabrous or subglabrous rather than pubescent margin of the segments, the usually glabrous rather than pubescent mid- nerve and the long and narrow rather than relatively short and broad segments. Also the growing tip of the rhizome usually has a tuft of dark hairs.
Although not closely related, var. africanum approaches var. pseudocaudatum in some characters and the differences are dis- eussed under the former variety.
Var. pseudocaudatum grows in open woods, pastures, thickets, in burnt-over areas and abandoned fields, usually in dry sterile soil but sometimes in fairly damp or rich places.
It is primarily of Coastal Plain distribution: Cape Cod, Massachusetts, and Long Island, New York, to Florida and Texas; also inland in North Carolina, Tennessee, West Virginia, Ohio, Indiana, Missouri, Arkansas and Oklahoma.
UNITED STATES OF AMERICA.—MassACHUSETTS: Harwich, Barn- stable Co., Sept. 2, 1918, Fernald & Long 15914 (Herb. New Eng. Bot. Club). New York: Babylon, Long Island, Sept. 8, 1898, Clute 339 (NY), isotype of Pteris aquilina var. pseudo- caudata Clute! New Jersey: Hammonton, Aug. 19, 1879, Kitchel (G) ; Atsion, Burlington Co., Aug. 10, 1926, Benner, Long & Bassett (G). Devaware: Seaford, Aug., 1874, Canby (Herb. Field Mus. no. 149427) ; Laurel, Sussex Co., Aug. 19, 1880, Com- mons (G). MaRYvLAND: 3 miles southeast of Ridgely, Caroline Co., Sept. 24, 1938, Wherry (G). West VriRGINI1A: Rickett's Place, Cabell Co., Sept. 13, 1936, F. A. Gilbert 519 (F, NY). VinGINIA: Ocean View, Norfolk Co., Oct. 4, 1912, Tidestrom 6184 (G); Buckroe, May 18, 1912, B. L. Robinson 341 (G); Great Dismal Swamp, June 18, 1936, Fulling (NY). Norru CangoLiNA: Tryon, Polk Co., May, 1918, Millspaugh 4083 (F); 4 miles east of Hamlet, Richmond Co., July 2, 1927, Wiegand &
71 Peattie, Fl. Ind. Dunes, 29 (1930) : “асс. to Clute”.
19411 Tryon, Jr..—Revision of the Genus Pteridium 51
Manning 21 (G); Goldsboro, Wayne Co., June 21, 1935, Cor- rell 1382 (G). Зостн Caronina: near Navy Yard, Charleston, May 4, 1912, B. L. Robinson 198 (С); Myrtle Beach, Horrey Co., June 12, 1936, Correll 5218 (©); Laurel Hill, July 6, 1936, Tar- box 735 (NY). GEORGIA: Sumter Co., July 24, 1901, Harper 1110 (F, G, NY, US); Near Darien, McIntosh Co., June 20, 1936, Correll 5456 (G). Fiorina: Warrenton, May 23, 1903, Tracy 8633 (F, G, US); Eustis, Lake Co., May 1-15, 1894, Nash 638 (БЕ, С, NY, US). Kentucky: Mammoth Cave Road, Edmonson Co., July 2, 1916, King 121 (F). TENNESSEE: Look- out Mt., Eggleston (NY). Onto: Salem Township, Meigs Со, Oct. 10, 1931, C. H. Jones (Herb. Ohio U.). INDIANA: 1 mile east of "aswell, Crawford Co., Aug. 17, 1913, Deam 13976 (Deam Herb.) ; 15 mile south of Emison, Knox Co., Sept. 2, 1939, Tryon 4268 (G). ALanAMa: Mobile Co., June, 1905, Dukes (G); near Fairfax, Chambers Co., Aug. 17, 1936, Correll 6562 (G) ; Auburn, Lee Co., Oct. 14, 1897, Earle & Baker (NY). MuississiPPI: West of Kosciusko, Attala Co., May 17, 1933, C. A. & U. F. Weatherby 6300 (G, NY, US) ; French Camp, April 28, 1899, 7. M. Clute 54 (F, NY). Міѕѕосві: Monteer, May 14, 1901, Bush 474 (G); Chadwick, Christian Co., Oct. 5, 6, 10, 1915, Eggleston 12187 (NY); Webb City, Jasper Co., Aug. 22, 1920, E. J. Palmer 18788 (NY). Arkansas: Nashville, Howard Co., Oct. 19, 1932, Demaree 9952 (С); West Otis, Sevier Co., July 26, 1937, Brinkley 256 (T); Wilmar, Drew Co., Oct. 12, 1936, Demaree 14008 (NY). LovisiANA: Alexandria, May 31, 1899, C. R. Ball 511 (F, G, NY, US); Chapin, Natchitoches Parish, Oct. 5, 1915, E. J. Palmer 8845 (NY). OkLaAHoMa: Page, LeFlore Co., Sept. 9, 1913, Stevens 2715 (G, US); Idabel, MeCurtain Co., May 29, 1916, Houghton 3909 (G, NY). Texas: 10 miles south of Yellow Pine, Sabine Co., Oct. 3, 1934, Cory 10750 (G) ; Huntsville, June 3-12, 1908, Dixon 122 (Е); Houston, Harris Co., May 18, 1917, E. J. Palmer 11942 (NY).
8. PTERIDIUM AQUILINUM var. AFRICANUM?? Bonap. Notes Ptérid. 1: 62 (1915). Prata 650, ric. 6, PLATE 652, FIG. 3, MAP 6.
Pteridium *centrali-africanum Hieron. ex Fries, Wiss. Ergebn. Sehwed. Rhodesia-Kongo Exp. 1': 7 (1914). Pteridium aqui- linum (L.) Kuhn var. caudatum (L.) Sadebeck f. africanum (Bonap.) Bonap. Notes Ptérid. 14: 321 (1923).
Growing tip of the rhizome with a tuft of dark hairs; frond 0.6-1 m. high, vernation gleichenioid; stipe shorter than the blade; blade 4-8 dm. long, ovate to broadly ovate, not ternate, tripinnate-pinnatifid, or more often quadripinnate; rachis gla- brous or subglabrous; pinnae and pinnules acute to obtuse;
22 Described under ssp. caudatum.
52 Rhodora [FEBRUARY
pinnules at ап oblique angle to the costa; costules glabrous to slightly pubescent; penultimate segments pinnate; longest entire segment or entire part of a segment from five to eight times as long as broad; ultimate segments usually straight, at least some, often many, narrowed at the base, the upper surface glabrous, the margin glabrous, the lower surface glabrous or very slightly pubescent on the midnerve; fertile and sterile indusium glabrous, the fertile usually 0.3 mm. wide, the sterile 0.1-0.2 mm. wide, the fertile portion broader than the sterile on the same segment, or no broader.
Түрк: Busse 944, in Herb. Muséum d'Histoire Naturelle, Paris (not seen).
Type Locauiry: “Afrique Orientale allemande. Magaba- Thal.”
Var. africanum may be separated from var. typicum by its more finely cut, often quadripinnate, blade, its glabrous rather than pubescent rachis and its glabrous rather than ciliate fertile and sterile indusium. Also the pinnules are at an oblique angle to the costa rather than at right angles and the ultimate segments are usually narrowed at the base and glabrous or subglabrous rather than adnate or broadest at the base and densely pubescent beneath. It differs from var. pseudo- caudatum in its gleichenioid rather than equal vernation, its ovate or broadly ovate rather than broadly triangular blade and in that the ultimate segments are usually narrowed rather than adnate or broadest at the base.
Var. africanum grows in dry moderately light woods, in virgin forest and in open grassland in tropical Africa, up to 1400 m.
Arrica.—BELGIAN Conco: Stanley Pool, Sept., 1883, H. Н. Johnston (U. S. Nat. Herb. no. 22427); Elizabethville, Dec. 30, 1919, Shantz 513 (US) ; Albertville, 1931, Lugen 89 (G). Ровто- GUESE West ArRICA: Near Cuanza River, Sept. 24, 1923, А. G. Curtiss 358 (G). Union or Ѕоотн Arrica: Ndola, northern Rhodesia, Stevensen 400 (US). TANGANYIKA (GERMAN EAST Arrica): N’Gano-N’Gano, Urundi, March 17, 1919, Schantz 757 (US).
PTERIDIUM AQUILINUM ssp. CAUDATUM (L.) Bonap. Notes Ptérid. 1: 62 (1915)
Growing tip of the rhizome with a tuft of dark hairs; rachis
glabrous to slightly pubescent, rarely densely pubescent; costae
glabrous to moderately pubescent; pinnules usually nearly at right angles to the costa, rarely at a somewhat oblique angle;
19411 Tryon, Jr.,—Revision of the Genus Pteridium 53
Ranges of PTERIDIUM AQUILINUM var. CAUDATUM (map 11); var. ARACHNOIDEUM (map 12).
54 Rhodora [FEBRUARY
longest entire segment or entire part of a segment variable, from four to seventeen times as long as broad; ultimate segments usually linear to long-linear, sometimes ovate or oblong-ovate, mostly decurrent or more strongly decurrent than surcurrent, sometimes most, but not all, of the segments adnate, the upper surface glabrous to moderately pubescent, the lower surface usually appressed-pubescent with straight hairs or arachnoid- pubescent, rarely sublanuginose or glabrous, usually having a farinaceous appearance (PLATE 650, ric. 13), fertile indusium 0.3-0.5 mm. wide, the sterile 0.1—0.35 mm. wide. Map 1 (stars).
9. PTERIDIUM AQUILINUM var. CAUDATUM (L.) Sadebeck,? Jahrb. Hamb. Wiss. Anst. 14: Beiheft 3, 5 (1897), as to indicated basinym Pteris caudata L., not as to plant. PLATE 650, Fics. 9, 12, PLATE 653, FIG. 4, MAP 11. Illustrations: Plumier, Pl. Amer. t. 22 (1693) ; Jacquin, Ic. Pl. Rar. 3: t. 645 (1786—1793); Brit- ton, Fl. Bermuda, 419 (1918).
Pteris caudata L. Sp. Pl. Ed. 1, 2: 1075 (1753). Pteris aqui- lina L. var. caudata (L.) Link, Hort. Berol. 2: 33 (1833). Al- losorus caudatus (L.) Pr. Tent. Pterid. 154 (1836). Pteris aquilina L. var. mexicana Fée, Mém. Fam. Foug. 8: 114 (1857). Pteris caudata L. var. mexicana Fée, Mém. Fam. Foug. 9: 8 (1857), nomen nudum. Ornithopteris caudata (L.) J. Sm. Hist. Fil. 298 (1875). Cincinalis caudata (L.) Trevis. Atti Soc. Ital. sc. nat. 17: 239 (1875). Pteridium caudatum (L.) Maxon, Proc. U. S. Nat. Mus. 23: 631 (1901). Pteridium aquilinum (L.) Kuhn var. caudatum (L.) Sadebeck f. glabratum Hieron. Hed- wigia 48: 246 (1909). Pteridium aquilinum (L.) Kuhn var. caudatum (L.) Sadebeck f. pubescens Hieron. Hedwigia 48: 246 (1909). Filix-foemina aquilina (L.) Farwell var. caudata (L.) Farwell, Am. Mid. Nat. 12: 290 (1931).
Frond 0.6-7 m., usually about 1.2-2.5 m. high, the taller fronds scandent, vernation not clearly observed, apparently gleichenioid; stipe usually about as long as the blade; blade 0.3— 4 m., usually about 0.6-1 m. long, triangular to broadly ovate or long-triangular in large plants, tripinnate, or more usually tripinnate-pinnatifid or quadripinnate; costules of the penulti- mate segments usually glabrous, sometimes slightly pubescent above and beneath with long white hairs, rarely pubescent above with short white hairs, or beneath with dark or bicolorous hairs; free lobes not present along the rachis, costae and costules; ulti- mate segments usually linear or long-linear, sometimes oblong- ovate or ovate, the margin glabrous or infrequently slightly pubescent, the lower surface usually densely appressed-pubescent with long straight hairs, rarely arachnoid-pubescent, sometimes glabrous, having a farimaceous appearance except in the glabrous
23 On many of my annotation labels the combination is accredited to Underwood (1900).
19411 Tryon, Jr.,—Revision of the Genus Pteridium 55
forms, the midnerve usually glabrous, rarely pubescent with dark or bicolorous hairs, only rarely membranous wings present along the veins and midnerve; fertile and sterile indusium usually glabrous, sometimes slightly or densely ciliate, the fertile portion broader than the sterile on the same segment; cells of the sterile indusium large, in fairly definite rows (PLATE 650, ric. 12).
TYPE: Specimen in the Linnaean Herb. (not seen). Linnaeus had two specimens labelled Pteris caudata in his herbarium in 1753.74 One is apparently var. pseudocaudatum and the other is clearly var. caudatum. The latter specimen is taken as the type. Since Linnaeus’ description in the Species Plantarum was taken from his Hortus Cliffortianus, a specimen, if there is one, in the Clifford Herbarium might be considered to be the type. However, since there is a perfectly good specimen available in Linnaeus’ own herbarium, it seems best to designate that as the type. Although one of Linnaeus’ specimens is apparently var. pseudocaudatum, the application of his name is perfectly clear from the figures and localities cited by him.
Type Locauity: West Indies. Linnaeus, Sp. Pl: “Jamaica, Dominica.”, Hort. Cliff.: “Santo Domingo, Jamaica ete."25
The typical phase of var. caudatum, with the segments long and narrow and remote, occurs mainly in the West Indies, Florida and in the coastal regions of northern South America, Central America and Mexico. At the higher altitudes, mostly in Cen- tral America and Mexico, there is a phase with the segments relatively short and broad and approximate. The extremes of this phase (Lago San José, Porto Rico, July 15, 1912 (U. S. Nat. Herb. no. 566772) ; San Jose, Tamaulipas, Mexico, 600-1100 m., 1902, Kemp (NY); San Rafael de Norte, Nicaragua, March 25, 26, 1917, Miller & Griscom 157 (US) ; Costa Rica, 1800 m., Aug. 2, 1933, Solis 277 (F); Columbia, Charetier 33 (NY, US); El Salvador, 1200—1500 m., Standley 21537; Nicaragua, 850 m., Mazon, Harvey & Valentine 7421) are well marked, but there is a great deal of intergradation from one phase to the other. In fact, almost a third of the specimens I have examined are intermediate, and the ranges overlap considerably in individual
% Mr. C. A. Weatherby has kindly made his notes on the Linnaean Herbarium avail- able to me. ?» Linnaeus, Hortus Cliffortianus, 473 (1737).
56 Rhodora [FEBRUARY
cases. I do not believe that this variation can reasonably be given varietal status.
Occasional specimens, especially of the “compact” phase men- tioned above, but also of the typical phase, have the segments only slightly decurrent but these can usually be placed in var. caudatum by the characteristic pubescence.
Anthony 400, Lower California, has the lower surface of the segments appressed-pubescent with short hairs as is often the case in var. esculentum.
Var. caudatum can be distinguished from var. esculentum by its lack of free lobes along the rachis, costae and costules; and the fertile indusium is broader than the sterile on the same seg- ment rather than usually no broader. Also the glabrous phases of var. caudatum do not have a farinaceous appearance.
Var. caudatum is most closely related to var. arachnoideum; the differences are discussed under that variety.
It grows in clearings, rough pastures, on dry hillsides, in cut- over forest land, in fresh-water marshes, in pinelands, scrublands and in shady rocky places, mostly at the lower altitudes but up to 2000 m. in Central America and Mexico, and 3000 m. in Venezuela; and from 1000 m. to 1300 m. in the Revillagigedo Islands.
It occurs from Bermuda to Florida, West Indies, Mexico, Central America and northernmost South America.
UNITED STATES or AMERICA.—FLoRIDA: Fort Myers, Lee Co., June 1, 1916, J. P. Standley 213 (F, G, NY, US); Cape Sable, July, A. H. Curtiss 3705* (F, G, NY, US).
Brermupa: Devonshire Marshes, Aug. 31—Sept. 20, 1905, Brown & Britton 159 (F, G, NY, US).
Wesr Inpres:—Baunama IsLanps: Near Nassau, Feb. 11, 1903, A. H. Curtiss 74 (F, G, NY, US); Orange Creek and vicinity, Cat Island, Feb. 27, 28, 1907, Britton & Millspaugh 5754 (F, NY). Cusa: Monte Verde, Jan.-July, 1859, Wright 872 (F, G, NY, US); Josephina, north of Jaguey, Yateras, Oriente, April 23, 1907, Maxon 4129 (G, NY, US). Jamaica: Mulgrove, north of Ipswich, St. Elizabeth, April 1, 1920, Maxon & Killip 1488 (F, G, NY, US). HisPANIOLA: Anse Galette, Gonave Island, Haiti, March 3-14, 1920, Leonard 3210 (F, G, US), 3208 (NY); Vicinity of Mission, Haiti, April 17-May 4, 1920, Leonard 3916 (US); San Lorenzo Bay, south coast of Samana Bay, Dominican Republie, April 5-11, 1921, Abbott 1275 (G, US); Province of
19411 Tryon, Jr.,—Revision of the Genus Pteridium 57
Barahona, Dominican Republic, July, 1911, Fuertes 1053 (F, G, NY, US). Porro Rico: Santurce, Jan. 22, 1903, A. A. Heller 6446 (F, G, NY, US). Monrtserrat: Turner (U. S. Nat. Herb. no. 428409). Awntiaua: (US).
Mexico.—Islands off the coast of Lower California and on the adjacent mainland, March-June, 1897, Anthony 400 (F, G, NY, US). Nuevo Leon: Sierra Madre, July 6, 1933, C. H. & M. T. Mueller 366, in part (G). Tamauuipas: La Vegonia, San Jose, July 5, 1930, Bartlett 10096 (US). SINALOA Sierra de Chabarria, 1927, Ortega 4079 (US). Nayarir (Terr. Терс): Jan. 5—Feb. 6, 1892, E. Palmer 1948 (С, US); Jalisco, Nov. 11, 1925, Ferris 5958 (G, US). Vera Cruz: Mt. Orizaba, Aug. 21, 1891, Seaton 110 (F, G, NY); near Jalapa, May 12, 1900, Pringle 8342 (G, NY, US). СоглмА: Socorro Island, Revillagigedo Is- lands, May 8, 1925, Mason 1662 (G, US). Guerrero: Montes de Oca, San Antonio-Buenos Aires, May 5, 1938, Hinton 14083 (G, US). Oaxaca: Tolosita, June, 1937, L. Williams 9614 (F). CuiaAPAS: Tacnalpan, July 28, 1890, Rovirosa 835 (G, NY). Yucatan: Tuxpena, Campeche, March 23, 1932, Lundell 1431 (F). Terr. Quintana Roo: Cozumel Island, Feb. 20, 1899, Millspaugh 1551 (F, G).
CENTRAL AMERICA.—BriTISH Honpuras: Big Creek, April 27, 1929, Schipp 190 (F, G, NY, US). GvarEMALa: Vaxactum, Dept. Peten, April 7, 1931, Bartlett 12521 (F, US); Vicinity of Puerto Barrios, Dept. de Izabal, June 2-6, 1922, Standley 25028 (US). Er SALvaApon: Volcan de San Vicente, Dept. San Vicente, March 7, 8, 1922, Standley 21537 (G, US). Honpuras: San Pedro Sula, Dept. Santa Barbara, May, 1888, Thieme (J. D. Smith no. 5650) (G, NY, US); Ceiba, Oct. 18, 1916, Dyer A134 (F, G, US). NicARAGUA: Casa Colorado and vicinity, south of Managua, June 27, 1923, Maxon, Harvey & Valentine 7421 (G, US). Costa Rica: 1881, J. J. Cooper (Herb. Field Mus. no. 347710, G, U. S. Nat. Herb. nos. 22440, 154190) ; San Jose, Prov. San Jose, 1887, J. J. Cooper (J. D. Smith no. 6018) (US). Panama: Pedro Miguel, Jan. 27, 1918, Killip 2825 (US); vicinity of El Boquete, Chiriqui, March 2-8, 1911, Maxon 4926 (US).
SourH AMERICA.—COLUMBIA: Santa Marta, near Onaca, Aug. 22, 1898-1901, H. H. Smith 1088 (F, G, NY, US); Vicinity of Medellin, 1911, Charetier 33 (NY, US). VENEZUELA: 215 miles east of Merida, State of Merida, Jan. 23, 1931, Reed 210 (US).
10. PTERIDIUM AQUILINUM var. ARACHNOIDEUM (Kaulf.) Her- ter, Rev. Sudam. Bot. 5: 21 (1937) ?9 PraTE 650, rics. 10, 13, 14, PLATE 653, ric. 3, MaP 12. Illustrations: Vellozo, Fl. Flum. 11: t. 80 (1827) ; Christ, Geog. Farne, Fig. 9 (1910), habitat.
?9 Combination incorrectly attributed to Baker.
58 Rhodora [FEBRUARY
Pteris psittacina Pr. Delie. Prag. 1: 185 (1822). Although I have not seen Presl’s type, his description clearly refers his name to Pteridium aquilinum and the locality, Rio de Janeiro, to var. arachnoideum. Material recently referred to Pteridium psittacinum represents juvenile leaf-forms of var. arachnoideum and var. caudatum. The leaf-cutting of young plants of these varieties is considerably different from the mature condition and the typical types of pubescence are not developed. Pteris campestris Schrad. Gott. gel. Anz. 18241: 871 (1824). Pteris arachnoidea Kaulf. Enum. Fil. 190 (1824). Allosorus psittacinus (Pr.) Pr. Tent. Pterid. 153 (1836), as A. psitaccinus. Allosorus arachnoideus (Kaulf.) Pr. Tent. Pterid. 153 (1836). Pteris aqui- lina L. var. arachnoidea (Kaulf.) D. C. Eaton, Proc. Amer. Acad. n.s. 8: 203 (1861). Pteris Gardneri Pr. ex Ettingsh. Denkschr. Ak. Wiss. Wien, 23: 42 (1864). Aquilina Gardneri Pr. ex Et- tingsh. Farnkr. 91 (1865), in synonymy. Pteris aquilina L. var. psittacina (Pr.) Baker in Martius, Fl. Brasil. 1?: 404 (1870). Cincinalis arachnoidea (Kaulf.) Trevis. Atti Soc. Ital. sc. nat. 17: 239 (1875). Pteridium aquilinum (L.) Kuhn var. esculen- tum (Forst.) Kuhn f. arachnoideum (Kaulf.) Hieron. Hedwigia 48: 246 (1909). Pteridium arachnoideum (Kaulf.) Maxon, Journ. Wash. Acad. Sci. 14: 89 (1924). Filix-foemina aquilina (L.) Farwell var. arachnoidea (Kaulf.) Farwell, Am. Mid. Nat. 12: 290 (1931). Pteridium psittacinum (Pr.) Maxon, Proc. Biol. Soc. Wash. 46: 141 (1933).
Frond 1-3 m. high, vernation not clearly observed, apparently gleichenioid; stipe usually shorter than the blade; blade 0.5-2 m. long, ovate-triangular to long-triangular in large plants, tripin- nate to quadripinnate; costules of the penultimate segments usually pubescent beneath and less so above with short white and also dark or bicolorous hairs, sometimes glabrous; free lobes present along the rachis, costae and costules; ultimate segments ovate to linear, the margin often pubescent, the lower surface arachnoid-pubescent, rarely appressed-pubescent with short straight hairs, or glabrous, nearly always having a farinaceous appearance, the midnerve usually pubescent with dark or bi- colorous hairs, membranous wings usually present along the veins and midnerve (PLATE 650, ric. 14); fertile and sterile indusium ciliate and sometimes also pubescent on the outer surface, or glabrous, the fertile portion no broader than the sterile on the same segment; cells of the sterile indusium small, irregularly arranged (cf. PLATE 650, FIG. 11).
Tyre: Chamisso, probably at Berlin (not seen).
TYPE LocALITy: Brazil.
19411 Tryon, Jr.,—Revision of the Genus Pteridium 59
Riedel, Brazil, “Ex. herb. hort. Petropolitani” (G) and Curran 128, Bahia, Brazil (G, US) differ from typical var. arachnoideum in having no free lobes along the rachis, costae and costules.
The following are intermediate between this and var. cauda- tum. Hitchcock 17031, Penal Settlement, British Guiana, Dec. 3-9, 1919 (G, US) has free lobes present and a farinaceous ap- pearance, even though glabrous, as in var. arachnoideum but has the fertile indusium broader than the sterile as in var. caudatum; Heller 4468, 14 miles northeast of Mayaguez, Porto Rico, Feb. 1, 1900 (С, F, NY, US) and Mazon 4075, San Piedra, Oriente, Cuba, April 14, 1907 (G, US) are similar to var. caudatum in characters of pubescence and of the indusium but they have a few free lobes as in var. arachnoideum; Rose & Painter 7595, Jalisco, Mexico (US) is arachnoid-pubescent but. has no free lobes; and Pennell 5162, La Cumbre, El Valle, Colum- bia (US) and Ariste-Joseph A207, Bogota, Columbia (US) are arachnoid-pubescent as in var. arachnoideum but have the fertile indusium broader than the sterile and have no free lobes as in var. caudatum.
Var. arachnoideum differs from var. caudatum in having free lobes along the rachis, costae and costules rather than not having them; the midnerve is usually pubescent with dark hairs rather than usually glabrous and the fertile portion of the indusium is no broader than the sterile on the same segment rather than broader. Also the lower surface is arachnoid-pubescent rather than appressed-pubescent with straight hairs, the cells of the indusium are smaller and irregularly arranged and membranous wings are usually present on the veins and midnerve beneath rather than usually not present. Even the glabrous phases have a farinaceous appearance beneath.
The differences between var. arachnoideum and var. esculen- tum, with which it has often been confused in the past, are dis- cussed under the latter variety.
Var. arachnoideum grows on open slopes, in open rocky places, in thickets, forests, grassland, in cleared land and on the edge of forests from the lower elevations up to 3000 m.; and from 300 m. to 700 m. in the Galapagos Islands.
It ranges from the West Indies, Cuba to Trinidad, to southern Mexico, Central America, Galapagos Islands and throughout
60 Rhodora [FEBRUARY
South America except the southern portion; also it is apparently absent from most of the Amazon Basin.
West Inpres:—Cusa: Monte Verde, Jan.-July, 1859, Wright 985 (G); Loma del Gato and vicinity, Sierra Maestra, Aug., 1923, Hioram & Clement 6497 (US); Santiago, Santa Ana, March 23, 1902, Hamilton 240 (NY). JaAmatca: Vicinity of St. Helens Gap, St. Andrew, March 4, 1920, Maxon & Killip 619 (F, G, NY, US). HisPANIOLA: vicinity of Furey, Haiti, May 26-June 15, 1920, Leonard 4339 (G, US); Prov. Monte Cristi, Santo Domingo, June 24, 1929, Ekman 12990 (NY, US). Ponro Rico: Aug. 28, 1885, Sintenis 2658 (G, US). Santa Lucia: Ventine Sulphur Springs (Soufrière), May, 1935, Box 449 (US). Trinipap: St. Ann, March 17, 1920, Britton, Hazen & Mendelson 676 (G, NY); 1877-8, Fendler 77 (G, NY, US).
Mexico.——Mexico: Nanchititla, Temascaltepec, Feb. 14, 1935, Hinton 7371 (G, NY). Vera Cruz: Zacuapan, Dec., 1912, Pur- pus 6191 (F, G, NY, US). Gurrrero: Montes de Oca, San Antonio-Buenos Aires, May 3, 1938, Hinton 14069 (G, US). Oaxaca: Cuicatlan, June 16—22, 1898, Conzatti & Gonzalez 748 (G).
CENTRAL AMERICA.—GUATEMALA: Volcan de Fuego, Salvin (G); Yzabal, Dept. Yzabal, J. D. Smith 1565 (G, US). Er SaLvApoRn: Cerro del Guayabal, Jan., 1924, Calderon 2008 (G, US). Номроваѕ: about 15 miles east of Ceiba, Dept. Atlantida, July 21, 1938, Yuncker, Koepper & Wagner 8555 (NY). Nicaracua: San Rafael de Norte, March 25, 26, 1917, Miller & Griscom 152 (US). Costa Rica: from Vara Blanca to La Con- cordia, July 23, 1923, Maxon & Harvey 8400 (US); 1901—1905, Werckle (U. S. Nat. Herb. no. 575231) ; San Ramon, April, 1913 (Herb. Field Mus. no. 404457). Panama: Vicinity of Monte Lirio, Prov. Chiriqui, June 27—July 13, 1935, Seibert 234, in part (G).
Ѕоотн AMERICA—CoLUMBIA: Palmira, Dept. El Valle, May 27, 1922, Pennell & Killip 6100 (G, NY, US); Santa Marta, Aug. 26, 1898—99, H. H. Smith 1091 (NY). VENEZUELA: Maracai, Vogl (G); Island of Margarita, Aug. 28, 1903, J. R. Johnston 177 (G); Tovar, 1854—5, Fendler 104 (G); 1917, Curran & Haman 1111 (G, NY). British Gutana: Malali, Demerara River, Oct. 30-Nov. 5, 1922, de la Cruz 2658 (F, G, NY, US); Mt. Iramaikpang, northwest part of Kanuku Mts., April 22, 1938, A. C. Smith 3657 (G). Frenc Gurna: Near Cayenne, Oct. 8, 1830, Leprieur 100 (F, G, US). Ecuapor: Western San Miguel Mts., Oct. 21, 1933, Schimpff 247 (F); Andes, 1857-9, Spruce 5601 (G, NY) ; Wreck Bay, Chatham Island, Galapagos Islands, July 6, 1905—6, A. Stewart 996 (F, G, NY, US). Рево:
19411 Tryon, Jr..—Revision of the Genus Pteridium 61
Estrella, Dept. Ayacucho, May 8, 14, 1929 Killip & Smith 23095 (NY, US); Cero de Cusilluyoc, Dept. Cusco, May 3-6, 1925, Pennell 13986 (G, NY, US); Tarapoto, Dept. San Martin, Dec., 1929, L. Williams 5971 (F). Braziu: Pará, Nov., 1913, Petelot (Herb. Field Mus. no. 593026) ; Near Petropolis, July 10—16, 1882, J. Ball (G); Mt. Itatiaya, vicinity of Monte Serrat, State of Rio de Janeiro, Dec. 31, 1928, L. B. Smith 1587 (F, G, NY, US) ; Sao Leopoldo, Rio Grande do Sul, Rick 24 (G). Borivia: Incachaca, Dept. Cochabamba, Prov. Chapare, Jan. 24, 1929, Steinbach 8927 (F, G); Lacotal, Dept. Cochabamba, Prov. Chapare, Feb. 25, 1929, Steinbach 9363 (F, G, NY); Tipuani, April, 1920, Buchtien 5271 (Е, US). Paraguay: Y-acá River, Cordillera Centralis, 1900, Hassler 6997 (G); Paraná River, 1909-10, Fiebrig 6138 (С). Uruauay: Catalan, Dept. Artigas, Nov., 1927, Herter 996 (NY); Pan de Azucar, Dept. Maldonado, Jan. 21, 1912, Osten 5688 (US). AmcENTINA: Fontana, Re- sistencia, Chaco, Feb., 1933, Schulz 727 (б); Dept. Punilla, Prov. Cordoba, March 16, 1939, Dawson 588 (G) ; Prov. de Catamarea, Nov. 11, 1910 (U.S. Nat. Herb. no. 1113401).
11. PTERIDIUM AQUILINUM var. ESCULENTUM (Forst. Kuhn, Chaetopt. 347 (1882). PEATE 650, Fics. 11, 15, PLATE 653, FIG. 1, MAP 9. Illustrations: Domin, Bibl. Bot. 85!: figs. 33, 34 (1914) ; Schk. Krypt. Gew. t. 97 (1809); Dobbie, New Zealand Ferns, Ed. 3, 183 (1930).
Pteris esculenta Forst. Pl. Escul. 74 (1786). Allosorus es- culentus (Forst.) Pr. Tent. Pterid. 154 (1836). Pteris auriculata Goldm. in Meyen, Nova Acta 19: supp. 1, 458 (1843). Pteris aquilina L. var. esculenta (Forst.) Hook. fil. Fl. N. Zel. 2: 25 (1854). Cincinalis esculenta (Forst.) Trevis. Atti Soc. Ital. sc. nat. 17: 239 (1875). Ornithopteris esculenta (Forst.) J. Sm. Hist. Fil. 298 (1875). Pteridium esculentum (Forst.) Diels in Engl. & Prantl, Nat. Pfl. 1*: 296 (1899). Pteris aquilina L. f. esculenta Christ 1n Warb. Monsunia 1: 68 (1900), without bibli- ography or reference. Pteris aquilina L. f. caudata Christ in Warb. Monsunia 1: 68 (1900), without bibliography or reference. Pteridium aquilinum (L.) Kuhn var. aequipinnulum Domin, Bibl. Bot. 851: 162 (1914). Pteridium aquilinum (L.) Kuhn var. pseudocaudatum Domin, Bibl. Bot. 85!: 161 (1914), not (Clute) Heller. Pteridium aquilinum (L.) Kuhn ssp. esculentum (Forst.) Bonap. Notes Ptérid. 4: 116 (1917).
Frond 0.6-3 m. high, vernation subgleichenioid; stipe about as long as the blade; blade 0.8-1.5 m. long, ovate to triangular, tripinnate to quadripinnate; costules of the penultimate segments glabrous above, glabrous to slightly pubescent beneath with white and often also dark hairs; free lobes usually present along
62 Rhodora [ FEBRUARY
the rachis, costae and costules; ultimate segments oblong or usually linear, the margin glabrous, the lower surface densely appressed-pubescent with long, or short, straight hairs, always having a farinaceous appearance, the midnerve usually glabrous, sometimes moderately pubescent with white and sometimes also dark hairs, no membranous wings present on the veins and mid- nerve; fertile and sterile indusium usually glabrous, rarely slightly ciliate, the fertile portion usually no broader than the sterile on the same segment; cells of the sterile indusium small, irregularly arranged (PLATE 650, ric. 11).
Tyre: Forster, location unknown. A fragment of the type “ex Forster Herb." “collected” by L. M. Underwood is at Herb. New York Botanical Garden (seen).
TYPE LOCALITY: Society Islands. Copeland?’ says that: “The sole Tahitian record is that of Forster . . . its absence from all later collections suggests that it does not beseem a wild plant to be edible."
Var. esculentum differs from var. arachnoideum in its pubes- cence,—appressed with straight hairs rather than arachnoid,— the absence rather than presence of membranous wings along the veins and midnerve and the midnerve usually glabrous or with white hairs rather than pubescent with dark or bicolorous hairs.
De la Cruz 2658, British Guiana and Gleason 423, Tumatu- mari, British Guiana, June 18-July 8, 1921 (G), var. arachnoide- um, approach var. esculentum in being appressed-pubescent with short hairs.
'The differences between var. esculentum and vars. caudatum and yarrabense are discussed under those varieties.
Var. esculentum grows in open places, pastures, thickets and clearings, from sea-level up to 1300 m.
It ranges from Australia to the Society Islands.
AUSTRALIA: Sassafras, Victoria, July 3, 1936, Lothian (G); Hall’s Gap, Grampian Mts., Victoria, Dec., 1912, Tilden 848 (F, б); Port Lonsdale, Victoria, Oct.-Nov., 1912, Tilden 762 (F, G); Bondi Bay, Sidney, New South Wales, Sept., 1912, Tilden 570 (F, О); near Mareton Bay, 1850—51, Strange (О); east coast (Nouvelle Hollande), 1845, Verreaux 267 (G, US). TASMANIA: Gunn (G). Norro_K IsLANp: 1884, Metcalfe (U. S. Nat. Herb. no. 22443). New ZkaALANDb: Whakarewarewa, Nov., 1909, Leland, Chase & Tilden 143 (F, G); South Island (Herb. Field Mus. no. 355839); Craig (Herb. Field Mus. no.
27 Copeland, Pteridophytes of the Society Islands. Bishop Mus. Bull. 93, 57 (1932).
19411 Tryon, Jr.,—Revision of the Genus Pteridium 63
596860, G) ; Taranaki, Heywood 56 (G); Mt. Ngongotaka, May- July, 1898, Prince (G); North Island (Ex. Herb. T. Kirk) (G). CHATHAM ISLAND: Oct., 1874, Kershner (US); Dec., 1874, Scott (US). New CaLEDoNiA: (Herb. Field Mus. no. 596487) ; 1861— 67, Deplanche 1563 (С); 1874-76, Germain (NY). New Hesripes: Aneiteum, Feb., 1859 (Herb. Field Mus. no. 596565). Кїлї IszANps: 1860, Seemann 809 (G); 1877—78, Horne 601 (G); (Herb. Field Mus. no. 593802). Navicaror Istanps (Samoa): 1873, McAlesber (NY). SociETv IstANps: “Ex. Forster Herb." (NY), fragment of type.
12. PrERIDIUM AQUILINUM var. YARRABENSE Domin, Bibl. Bot. 85': 161 (1914). PLATE 650, FIG. 16, PLATE 653, FIG. 2, MAP 10. Illustration: Domin, Op. cit. fig. 32.
Pteris lorigera Wall. List no. 103 (1829), nomen nudum; iso- type at US. Pteris semihastata Wall. List. no. 102 (1829), nomen nudum; ex Ag. Rec. Pterid. 48 (1839). Allosorus lorigerus (Wall.) Pr. Tent. Pterid. 154 (1836), nomen nudum.
Frond 0.5-3 m. high, vernation not observed; stipe about as long as the blade; blade 0.3-1.5 m. long, ovate to triangular, tripinnate to quadripinnate; costules of the penultimate seg- ments subglabrous above, moderately pubescent beneath with white and rarely also dark hairs; free lobes usually not present along the rachis, costae and costules; ultimate segments usually linear, sometimes oblong-ovate, the margin glabrous or rarely pubescent, the lower surface usually densely sublanuginose or rarely arachnoid-pubescent, not having a farinaceous appear- ance, the midnerve usually densely pubescent with white and infrequently also dark hairs, no membranous wings present on the veins and midnerve; the fertile and sterile indusium rather densely ciliate and usually also pubescent on the outer surface, the fertile portion broader than the sterile on the same segment, or no broader; cells of the sterile indusium small, irregularly ar- ranged.
'ТҮРЕ: 1910, Domin, probably at Praha (not seen).
TYPE LOCALITY: "Nordost-Queensland bei Yarraba."
Var. yarrabense differs from var. esculentum in having the lower surface of the blade sublanuginose rather than appressed- pubescent with straight hairs, the midnerve on the lower surface of the segments pubescent with white hairs rather than usually glabrous and the fertile and sterile indusium ciliate and pubes- cent rather than glabrous. Also it does not have a farinaceous appearance beneath and usually does not have free lobes along the rachis, costae and costules.
64 Rhodora [ FEBRUARY
Strange, Australia, and Seemann 809, Fiji Islands, var. esculen- tum, approach var. yarrabense in having the fertile and sterile indusium ciliate and the midnerve of the segments slightly pu- bescent beneath with white hairs.
Ching 5360, Kwangsi Prov., China (NY, US) is intermediate between var. yarrabense and var. Wightianum and therefore represents an intermediate between ssp. caudatum and ssp. typicum.
Var. yarrabense grows in clearings, thickets, open slopes and at the edge of woods, up to 2500 m. from northern India to Sumatra, east to the Philippine Islands and northeastern Australia.
India: Kumaon, Blinkworth, Wallich, 103 (US), isotype of Pteris lorigera Wall. Frenc Іхро-СніхА: Cochinchine, 1862— 66, Thorel (Herb. Field Mus. no. 540736); Bokor, Cambodia, Jan. 18, 1926, H. M. Smith 288 (G, US). Stam: Koh Chang, April 2, 1924, Н. M. Smith 197 (US). FrpERATED MALAY STATES: Penang, Dec., 1902 (U. S. Nat. Herb. no. 1097164) ; Penang (ex Herb. Oldfield) (NY); Larut, Perak, April, 1884, King’s col- lector 5926 (US); Tekik Sisih, Pahang, Aug. 19, 1929, Hender- son (US); Singapore (U. S. Nat. Herb. nos. 22437, 1097181). ANAMBA IsLANDS: Jemaja, Nov. 4, 1928, Henderson 20306 (US). Sumatra: Vicinity of Rantau, Parapot, Bila, March 28-May 10, 1932, Toroes 1832 (NY). РниплррІХЕ Istanps: Mindanao, Clemens 166 (F) ; Bucas Island, Oct. 4, 1906, Merrill 5264 (US) ; Cuming (U. 8. Nat. Herb. no. 853691); Cuming, “without a number” (G, NY). British Ховтн Borneo: Mt. Kinabalu, Kundasang, April 7, 1932, J. & М. 5. Clemens 29107 (NY, US) ; Kuching, Sarawak, Mjoberg (NY); Sandakan and vicinity, Sept.-Dec., 1920, Ramos 1697 (G, US). AvusrRALIA: Daintree River, North Queensland, Feb. 29, 1932, Brass 2199 (G).
DunBioUs AND REJECTED NAMES
Pteris aquilina L. var. mexicana Fée, Mém. Fam. Foug. 9: 8 (1857), nomen nudum.—Fée in Mém. Fam. Foug. 8: 114 (1857) described Pteris aquilina L. var. mexicana, which is clearly Pteridium aquilinum var. caudatum. In Mém. Fam. Foug. 9: 8 (1857) he lists Pteris caudata L. var. mexicana, with- out reference, but it undoubtedly represents a transfer of his earlier var. mexicana. However, he also lists Pteris aquilina L. var. mexicana which, while it apparently is not the same as his var. mexicana, Mém. Fam. Foug. 8: 114 (1857), cannot be defin-
19411 Tryon, Jr.—Revision of the Genus Pteridium 65
itely placed without an examination of the collections cited by him: “Orizaba, W. Schaffner (1834) No 136 et (1856) No 468.”
Pteris aquilina L. var. lanuginosa Fée, Mém. Fam. Foug. 9: 8 (1857), nomen nudum.—Although probably a synonym of Pteridium aquilinum var. Feei, this name cannot certainly be placed without an examination of the collection cited by Fée: “W. Schaffner, No 137 Orizaba (1854)."
Pteridium aquilinum longifolium, Am. F. Journ. 1: 88 (1910).—The publication of this name was an error. It was a new combination based on “Pteris aquilina longifolium Hook.", a name taken from sheet number 583 in the Herbarium of the American Fern Society. Dr. L. S. Hopkins informs me that the sheet is actually labeled Pteris aquilina lanuginosa Hook. and that lanuginosa was misread longifolium.
Pteris aquilina L. var. decipiens Lawson, Edinb. New Phil. Journ. n. s. 19: 110 (1864). Pteridium aquilinum (L.) Kuhn var. lanuginosum (Bong.) Fernald f. decipiens (Lawson) Fernald, Rhodora 37: 248 (1935).—Lawson's name cannot be definitely placed without an examination of the type, which is apparently lost. Lawson says he sent a specimen to D. C. Eaton, but an examination of Eaton's Herbarium at Yale University failed to reveal such a specimen. The name has been placed under var. pubescens and if this is correct would take precedence over it as an earlier varietal name. However, there is considerable doubt that the plant in question is var. pubescens and I am rejecting the name.
Although the plant was collected in the Gaspé, a likely place for var. pubescens to occur as a preglacial relie, and described as lanuginose, authentic material has never been collected in Gaspé Co., Quebec and the remainder of the description: "frond bipinnate, thin and membranous, . . . barren." indicates that the specimen was taken from a young plant. "Lanuginose" may apply to some part of the frond other than the lower surface between the margin and the midnerve. In a footnote Lawson himself says that: "Since the above was written, I have had an opportunity of studying the forms and development of Pteris aquilina [this would be var. latiusculum] and am quite satisfied that the doubtful plant [var. decipiens] 1з a state of that species, not old enough to be mature."
66 Rhodora [ FEBRUARY
Pteris aquilina L. var. glabra Hook. Sp. Fil. 2: 196 (1858). Pteridium aquilinum (L.) Kuhn var. glabrum (Hook.) Luerss. Rabenh. Kr. Fl. Ed. 2, 3: 107 (1889) —This name includes such a mixture that it cannot be definitely placed in any one variety Hooker placed the following names under it in synoymy: Pteris aquilina L. (= var. typicum), Pteris caudata Schkuhr (= var. latiusculum and var. pseudocaudatum), Pteris recurvata Wall., Pteris firma Wall. and Pteris excelsa Bl. (— var. Wightianum) and Pteris latiuscula Desv. (— var. latiusculum). His range- citations include the following localities: Europe, Cape of Good Hope (= var. typicum), Java (= var. Wightianum), Massa- chusetts (— var. latiusculum), New Orleans (— var. pseudo- caudatum) and Brazil (= var. arachnoideum).
EXPLANATION OF PLATES
Prate 650. Fig. 1, Segment of var. Wightianum, X 4, pubescence on one half not shown, on part of this half the sporangia and venation are not shown; Fig. 2, Segment of var. typicum, X 4, pubescence on one half not shown, on part of this half the sporangia and venation are not shown; Fig. 3, Segment of var. pubescens, X 4, pubescence on one half not shown, on part of this half the sporangia and venation and marginal pubescence are not shown; Fig. 4, Segment of var. Feet, X 4, pubescence on one half not shown, on part of this half the sporangia and venation and marginal pubescence are not shown; Fig. 5, Segment of var. decompositum, X 4, pubescence on one half not shown, on this half the sporangia and venation are not shown; Fig. 6, Segment of var. africanum, X 4, no pubescence re- moved, on part of the segment the sporangia and venation are not shown; Fig. 7, Segment of var. pseudocaudatum, X 4, no pubescence removed, on part of the segment the sporangia and venation are not shown; Fig. 8, Segment of var. latiusculum, X 4, no pubescence removed, except on part of the segment the marginal pubescence, sporangia and venation are not shown; Fig. 9, Segment of var. caudatum, X 4, on part of the segment the pubescence and sporangia are not shown and on another part the venation and farinaceous appearance are also not shown; Fig. 10, Seg- ment of var. arachnoideum, X 4, on part of the segment the sporangia and pubescence are not shown and on another part the venation and farinaceous appearance are not shown; Fig. 11, Sterile indusium of var. esculentum, about X 75; Fig. 12, Sterile indusium of var. caudatum, about x 75; Fig. 13, Farinaceous appearance of var. arachnoideum, about X 50, pubescence not shown; Fig. 14, Membranous wings along vein of var. arachnoideum, about X 40, pubescence and farinaceous appearance not shown; Fig. 15, Segment of var. esculentum, X 4, on part of the segment the pubescence and sporangia are not shown and on another part the venation and farinaceous appearance are not shown; Fig. 16, Segment of var. yarrabense, X 4, on one half the pubescence is not shown, and on part of it the sporangia and venation are not shown.
Prate 651. Fig. 1, Middle pinna of var. Feet, X %; Fig. 2, Upper pinna of var. decompositum, X №; Fig. 3, Tip of middle pinna of var. Wight- ianum, X %; Fig. 4, Upper half of middle pinna of var. typicum, X %.
19411 Raup,—An old Forest in Stonington, Connecticut 67
Prate 652. Fig. 1, Basal pinna of var. latiusculum, x %; Fig. 2, Basal pinna of var. pseudocaudatum, X Y2; Fig. 3, Next to basal pinnule of a basal pinna of var. africanum, X 6; Fig. 4, Frond of variant of var. latiusculum, X 1/6; Fig. 5, Next to basal pinna of var. pubescens, X %.
PrarE 653. Fig. 1, Upper half of middle pinna of var. esculentum, X 1%; Fig. 2, Basal pinna of small plant of var. yarrabense, X %; Fig. 3, Upper pinna of var. arachnoideum, X !4; Fig. 4, Tip of frond of var. caudatum,
X 2%.
AN OLD FOREST IN STONINGTON, CONNECTICUT Носн M. RAUP
Ох November 18, 1939, the writer had occasion to visit, in com- pany with a group of students in Ecology, a piece of old woodland at the mouth of the Pawcatuck River in southeastern Connecticut. The area is of particular interest because it has been considered by some to have been not far removed from a primeval condi- tion. Like so many of our supposed or aetual remnants of the virgin forests in southern New England, it suffered great damage during the hurricane of 1938, and there now remains only a battered representation of the once handsome stand of trees.
The late Dr. G. E. Nichols, in deseribing the virgin forests in Connecticut wrote an account of the tract.! Since this account gives an excellent picture of the forest, and since such descriptions now take on rather more historic interest than they had before the hurricane, it seems worth while to quote Dr. Nichols in full.?
“Southeastern Connecticut, so far as ascertained, possesses only one possible fragment of original forest and, notwithstanding the owner’s assurance that the area has never been cut over, the writer must confess to some doubt as to the primeval nature of the tract. The area in question, some 40 acres in extent, occupies a low hill bordering the Sound at the mouth of the Paweatuck River in the town of Stonington. In contrast to the forests heretofore de- scribed there is a complete absence of hemlock, beech, sugar maple, yellow birch, pine, and even chestnut. The character trees are white oak and black oak (Quercus velutina), especially the former, associated with which are shagbark hickory and red maple. The stand is of a more open character than in any of the areas previously mentioned and in general aspect the forest re- sembles the climax oak-hickory type of the Chicago region.
1 Nichols, G. E. “The Vegetation of Connecticut” IT, Torreya 13: 214-215 (1913).
? No photograph of the Stonington tract was published by Dr. Nichols. The writer is indebted to Dr. H. J. Lutz of the School of Forestry at Yale for making a thorough, though unavailing, search in Dr. Nichols' files for any photograph that the latter may have had.
68 Rhodora [ FEBRUARY
Trees with a diameter of from 45 to 60 centimeters are common. The ground is not deeply shaded and the low, dense underbrush is quite xerophytic, being composed largely of Gaylussacia bac- cata, Vaccinium corymbosum, Vaccinium stamineum, and Corylus americana. It is of course not impossible that the xero-meso- phytic nature of the tract is due to its extremely exposed location and that it really represents a virgin forest. Moreover it must be borne in mind that in general the forests of eastern Connecticut are less mesophytic than are those in other parts of the state.”
At the time of the writer’s first visit, the loggers were actively engaged in clearing out the tangle of fallen stems and branches which resulted from the hurricane. Nearly all of the tract was destroyed, leaving only a few scattered trees on the north, or landward side. A considerable number of logs of fair quality were being taken out, leaving newly cut stumps upon which ring counts could be made. In the short time available only eleven stumps were counted, ranging in number of rings from 110 to 136. Two facts were apparent: first, the trees all showed relatively wide rings in their early period of growth; and second, their aver- age age was approximately 123 years which was the length of time between the hurricane of 1815 and that of 1938.1 Since they were among the largest and oldest trees in the stand there seemed some evidence that Dr. Nichols’ chariness about considering the forest to be entirely unmodified might be well founded. The spread in ages among the trees remained to be explained, however, and it was especially desirable to find some more conclusive evi- dence of an actual release which could be attributed to a cata- clysm similar to that of 1938.
With these problems in mind, the tract was visited again in May, 1940. The logging operations were then nearly completed, and many more stumps were available. Fifty-one were counted on this occasion, all of them solid, or nearly so, to the core, and cut from one to three feet above the base. Most of them may be divided roughly into two age groups. The smaller of these con- sists of five trees, all of which contained over 140 clearly visible rings. All of these showed a distinct release, indicated by a more or less sudden widening of the rings, immediately after 1815. The larger group, as indicated by the earlier observations, con-
1Channing, Walter (Editor). New England Hurricanes, 1635, 1815, 1938. Boston, 1939. Brooks, Charles F. Hurricanes into New England: Meteorology of the Storm of September 21, 1938. Geog. Rev. 29: 119—127 (1939).
1941] Raup,—An old Forest in Stonington, Connecticut 69
sisted of 41 trees in which, judging by field observations, no ob- vious release was evident near the center. In the case of those with ages under 123 years none could be expected if the 1815 hur- ricane were significant; but some 21 individuals showed ages ranging between 124 and 140, and might be expected to have experienced a release after 1815.
There is no way of knowing how many years must be added to the count for each tree to round out its full life, unless each could be dissected to the very base. If, however, a release in 1815 is accepted, then it is to be expected that seedlings and small trees existing at that date had previously been suppressed in greater or less degree depending upon their ages and local positions in the ancient forest. Counts were made on two small trees in the ad- vance growth under the forest which was blown down in 1938. Both of these were young white oaks, growing in the shrubby cover of Gaylussacia, Vaccinium, and Corylus. One of them, 2 feet high, had a stem only about six inches long and about one half an inch in diameter. The other was four feet tall, with a stem about two feet long, and a diameter of about five-eighths of an inch at the base. The first proved to be twenty-nine years old, and the second sixteen years. In both the rings were so close together and confused that microscopic sections had to be made before good counts were possible!; and both showed a sudden re- lease after 1938. That is, there was one unusually wide ring at the outside which constituted the growth of 1939.
Two suggestions are to be derived from these observations. One is that about half of the trees blown down in 1938 began as seedlings or sprouts immediately after 1815, or were present as suppressed advance growth in a forest which existed prior to 1815, and were released at that time. Second, the absence of a release in the 21 trees showing 123 to 140 rings may be due to a certain amount of clearing prior to 1815. At the margins of the tract there are some areas that have been pastured considerably, with grassy glades and a partial cover of blackberry bushes. Old white-oaks growing in these spots showed no release in 1815, in- dicating that similar conditions may have existed at that time, possibly more widespread than subsequently.
The five older trees, which showed a definite release, had at-
1 The writer is indebted to Mr. F. €. Barghoorn for cutting these sections.
70 Rhodora [ FEBRUARY
tained such size in 1815 as to have reached up into better light and to have begun to put on more wood with more clearly defined rings each year. The oldest of the five showed about 180 rings on the stump to which must be added a considerable number for the years required to reach stump height. The rings nearest the center were, as would be expected, very small, but there followed & period of gradual increase to 1815 when the tree was about six inches in diameter. After the sudden release of 1815 it began to put on wood much faster. All of these five old trees had minute rings near the center, clearly indicating suppression. "There seemed to be no close relationship between the diameter of the trees and their age. Some that were 12 to 15 inches in diameter were fully as old as those two feet or more in diameter.
Another observation which bears out the theory that the 1815 hurricane was responsible for a great amount of damage is that the older trees counted, that is, those which had attained some size by 1815 and were reaching toward the canopy, were all on the landward side of the tract where they would not be subject to quite so much wind as those nearer the shore line. It has al- ready been noted that the remnants of the last hurricane are for the most part on the landward side.
This study indicates that the old oak-hickory wood at Stoning- ton was far from unmodified prior to the hurricane of 1938; that it was seriously damaged in 1815, presumably by the hurricane of that year; and that its canopy had been partially opened before 1815. "There is also the suggestion that in spite of the early clear- ing, and in spite of devastating destruction in 1815, it did not change much in its hardwood composition. The existence of ad- vanced growth of white oak, black oak, hickory, and red maple, suppressed because of a dense canopy prior to 1815, suggests that the trees in that canopy were not far different from those which were destroyed recently. Ап alternative hypothesis would be that prior to 1815 there had been some very old second growth or primeval forest of other facies which might have created a habi- tat similar to that which now appears in our old field white pine or red cedar stands. Such a primeval forest is not consistent, however, with the early accounts of the coastal vegetation in this
19411 Wherry,—Remarks on the Name Phlox nivalis 71
region, which describe forests of oaks, hickories, and chestnuts. Furthermore, it does not seem at all likely that in a growing colony along the coast, farm-land could have been cleared, used, and abandoned so long prior to 1815 as to give rise to old-field stands which could have ereated such a habitat.
Between 1815 and 1938 there is no evidence of release cutting. There are no sudden breaks in the development of wood except for occasional periods of very slow growth, probably due to dry or cold seasons. One such period occurred about 1890, and lasted for three or four years.
ARNOLD ARBORETUM Harvard University.
REMARKS ON THE NAME Рнгох nivatis.—In Ruopona xlii. 476 a question is raised as to the validity of the name Phlox nivalis as applied to the subulate-leaved species of the southeastern Coastal Plain and Piedmont. While this name, when proposed by Loddiges, was not accompanied by an adequate description, it was validated by Sweet in Brit. Flow. Gard. ii. no. 185 only four years later. The latter author published a more faithful representation of the habit of the plant than had been given by Loddiges’ crude plate, and also an accurate drawing of the floral parts. His text included a detailed english description and a latin diagnosis, so that all requirements are fulfilled.
Nuttall’s name Р. Hentzii was not published until seven years later, and his diagnosis was not so full as that of Sweet. A sheet in the herbarium of the Academy of Natural Sciences of Phila- delphia bears an annotation by Nuttall which shows that he was familiar with the earlier name, but he failed to mention this when he proposed his new one. Under our present rules of nomenclature Nuttall’s name is to be rejected as superfluous when published, and the plant should be known as Phlox nivalis Lodd. ex Sweet —Epcar T. Wuerry, University of Pennsylvania.
1 For reviews of early descriptions of southern New England Forests see Bromley, Stanley W. The original Forest Types of Southern New England, Ecol. Monog. 5: 61-89; and Raup, H. M. Recent Changes of Climate and Vegetation in Southern New England and Adjacent New York, Jour. Arn. Arb. 18: 79-117.
72 Rhodora [FEBRUARY
REAPPEARANCE OF RARE NEW ENGLAND MARINE ALGAE
WM. RANDOLPH TAYLOR
Sporapic occurrences of marine algae are often more puzzling to botanists than would have been the case had the plants be- longed to the land flora. It is notably more difficult to picture comprehensively the distribution of marine species, to locate small isolated colonies and, from lack of detailed knowledge of the sea bottom to differentiate physiographic from climatic limitations of range, particularly if the plants are rare, or at least rarely reported. The writer, having rather intensively observed a considerable area about Woods Hole, Massachusetts for over twenty years, has seen a number of reappearances of forms but once, or seldom, reported previously. It seems im- probable that the plants have been abundantly present during the intervening years but remained undetected; they may have persisted as scattered individuals vegetatively propagated, or they may have been reintroduced from outside the area.
The current season has provided two notable examples. On landing at Gay Head, Marthas Vineyard, on July 10th, 1940, the writer almost immediately noticed on the sand an alga not previously found by him, but recognized as Platoma Ваай (Farlow) Kuckuck. With the help of his class, there collecting algae for study, he secured several specimens and returned a week later for more. Altogether, many score specimens were brought back and preserved. Farlow (1875, p. 372) described! the plant as Nemastoma ? Bairdii on one small tetrasporic piece, and it has not been found in America in the interval; the next significant report? is that of Kuckuck (1912, p. 189) who made the present combination after finding more ample material off Helgoland and studying the asexual and sexual material thor- oughly. The writer has not noticed sexual plants in the new Gay Head material, but it is richly tetrasporiferous. Here, then, is а species but once seen, reported again in great abundance after nearly 70 years of probable absence from the type locality
1 Farlow, W. С. 1875. List of the Marine Algae of the United States with notes of New or Imperfectly Known Species. Proc. American Acad. Arts and Sci., 10(11) : 351—380.
2 Kuckuck, P. 1912. Uber Platoma Bairdii (Farlow) Kuckuck. Wiss. Meeresuntersuch., N. F., Abt. Helgoland, 5(3): 189-210.
1941] Taylor,—Rare New England Marine Algae 73
in an area inspected annually (for more than two decades) by dredging and shore collecting.
On the same day the Platoma came to light, one of the party brought to the writer in the field a small brown specimen which was easily distinguished as Tilopteris Mertensii (J. E. Sm.) Kütz., whieh he? had found in dredging off the same shore in 1931 (Taylor in Lewis and Taylor 1933, p. 151). On the first occasion two or three small pieces were found; on this there were several good specimens, all sporangial as before. There were none to be had at the same place a week later. Both Platoma and Tilopteris were washed ashore, but their normal habitat is on rocks in relatively shallow water. No plants were secured in 1940 when dredging nearby. With them was another species, rare in the region, which appeared in some abundance, namely Gloiosiphonia capillaris (Huds.) Carm. This the writer has only found on one or two earlier occasions.
A few words regarding other plants recognized as recently new to the district, or rare, may be pertinent. The Phaeophycean Acrothrix novae-angliae Taylor, first seen in 1925 and described? in 1928 (Taylor 1928, p. 577) has persisted and become more common, particularly during the current season. In 1934 the writer found? the tropical Sargassum fluitans Borg. washed ashore on Nonamesset Island (Taylor 1937, p. 211). The better known S. natans (L.) J. Meyen has frequently been reported, probably blown out of its usual path in the current of the Gulf Stream. We may add to these as a rare visitor to the coast S. Hystrix var. buxifolium (Ch.) J. Ag., which the writer found on the south side of Nantucket Island on the 17th of July 1938, associated with both of the above mentioned species. Among Rhodophyceae T'railiella intricata (J. Ag.) Batt. (Drew and Hof in Lewis and Taylor 1928, p. 196) has not continued as abundant as it was about 1930, but from 1927 to the present has turned up each summer, sometimes rare but generally in moderate amount. Asparagopsis hamifera (Har.) Okam. has continued in the flora,
? Lewis, I. F. and Taylor, W. R. 1933. Notes from the Woods Hole Laboratory, 1932. Кноровл 35: 147-154.
4 Taylor, W. К. 1928. A Species of Acrothrix on the Massachusetts Coast. Amer. Jour. Bot., 15: 577—583.
5'Taylor, W. R. 1937. Marine Algae of the Northeastern Coast of North America. vii + 427 pp., 60 pl. Ann Arbor.
74 Rhodora [ FEBRUARY
with less fluctuation (Taylor in Lewis and Taylor 1928, р. 197).* Plumaria sericea (Harv.) Rupr. (Taylor 1937, p. 330)? and Phycodrys rubens (Huds.) Batt. (Taylor 1937, p. 351)? have oc- casional seasons of relative frequency, but are generally very scarce. Lomentaria orcadensis (Harv.) Coll. (Taylor 1937, p. 309) was rarely reported, but for two or three years about 1931 it was very frequent in shallow water at several stations; it now has not been seen for some years. Scinaia furcellata (Turn.) Biv., considered not rare by Davis about 1911 at suitable places, is only found at rare intervals in small pieces. Among Myxophyceae Brachytrichia Quoyii (C. Ag.) Born. & Flah. also is sporadie, having been seen here three times in over twenty years, tend- ing to return in the old localities.
UNIVERSITY OF MICHIGAN.
NYMPHAEA TETRAGONA IN SOMERSET County, MarNE.— While on Waterfowl Survey work for the Maine Department of Inland Fisheries and Game, the writer with Virgil S. Pratt found Nymphaea tetragona Georgi in three localities in Somerset County. Previous reports of this plant for the State are by Wayne E. Manning, who found it in Chase Brook at Portage Lake!, and by Olof O. Nylander, who reported it at Salmon Brook Lake Bog in Perham? and in Mosquito Brook at Portage Lake? These stations are all in Aroostook County.
In Attean Pond in the town of Attean near the mouth of Moose River the plant was found scattered over several acres of open water, associated with Potamogeton natans L. and Poly- gonum natans A. Eaton. 'The water was about four feet deep over a bottom of firm mud; it had a pH of 7.0. Specimens of this collection will be deposited in the herbarium of the University of Maine and in the herbarium of the New England Botanical Club. In addition seattered plants were found in Dennistown. These were in Branch Stream, which flows into Little Big Wood
6 Lewis, I. F. and Taylor, W. R. 1928. Notes from the Woods Hole Laboratory, 1928. Ruopora, 30: 193-198.
1 Manning, Кноровл 38: 375, 1936.
2 Nylander, Contribution to Free Booters Club of Knowledge, Caribou, Maine, January,
1938. 3 Nylander, Presque Isle Star-Herald, November 21, 1940.
19411 Weatherby,—Deam’s Flora of Indiana 75
Pond. A few plants were also seen in Holeb Pond in the town of Holeb.
Dr. Е. С. Ogden of the University of Maine verified the writer's identification.—J. S. GASHWILER, Department of Inland Fisheries and Game, Augusta, Maine.
DkEAM's FLona or IxpiANA.!—Dr. Deam's Flora of Indiana has been long awaited. Its preparation has involved forty years of collecting and observation throughout the state, the accumulation of one of the largest and best private herbaria ever got together in this country and of a library of like scope and excellence, and unremitting vigilance in check- ing detail and in keeping abreast of current taxonomic developments. Those who knew this and were familiar with the author’s previous pub- lications had formed high expectations of the Flora. They will not be disappointed.
It has about everything yet thought of which a good local flora should have—an introduction giving physiographic information; an abundantly annotated list, arranged according to the Engler & Prantl system, of the 2530 species and varieties admitted and the 707 rejected records; a glossary, statistical summary, bibliography, ete. Its taxonomy in a few eritical groups has been contributed by specialists and in others rests on their determinations, but is enriched by an occasional original study or eritical comment out of the author's own experience; and its keys are largely original and made up from Indiana material. Less usual, but equally commendable, features are: a list of obsolete place-names, and one of collectors who have worked in Indiana, giving dates of birth and death when available, areas in which the collections of each were chiefly made, herbaria in which specimens are preserved, and the number of each collector’s specimens seen, even when that number is only one.
The general plan of the work is familiar enough; its distinction lies, not in any innovation, but in the high quality of its execution. There is constant and successful striving after clarity; all terms and methods used are explained with extreme care. There is, as Prof. Coulter points out in his foreword, an unusual wealth of ecological data for all species, as to soils, types of forest, associated species, and the like. And this and a great mass of other detail are set forth clearly and systematically.
If, with all this care, functioning in one minor matter, the orthography of English names, is not perfect; if Dr. Deam has failed to achieve con- sistency therein? and has occasionally admitted such orthographie and grammatical monstrosities as “ Smallflower Sweetbrier," he is no worse off than many others of us. Perhaps no one but the present reviewer will notice these minutiae anyway. Nevertheless, it would have been
1 Deam, Charles С. Flora of Indiana. Indiana Department of Conservation, Indianapolis. June, 1940. 1236 pp., 1 pl. 2247 maps. $3.50, at the State Library, Indianapolis.
? For instance: Curly Wildginger (p. 403), Blue Wild-indigo (591), Trailing Wild Bean (622); Crested Woodfern (48), Violet Wood Sorrel (627); Green Adder’s Mouth (349), Green Adder’s-mouth (1167); Few-flowered Spikerush (204), Large-flower Sensitive Plant (587).
76 Rhodora [FEBRUARY
gratifying if, as one detail of a major enterprise, he could have worked out a better balanced system, more consonant with the best existing usage.
In publishing the Flora, the Department of Conservation of Indiana has happily not limited Dr. Deam as to space and has turned out a neatly bound volume, with handsome typography. Some rather egregious printer's errors, such as repeating the line containing the name Dryop- teris Goldiana in the place where D. cristata ought to be, have, however, slipped by and seem particularly glaring in a work so meticulously careful in nearly every detail under the author's control.
There can be no question of the lasting importance of this Flora in its field. Few have been so well done. It will long serve students of floris- ties and distribution as an abundant source of reliable information; and (like Brendel's Flora Peoriana, which Dr. Deam once told me he had taken as his own model) it sets a standard of excellence for authors of future local floras to live up to—if they can.—C. A. WEATHERBY.
A New Form or Bromus INERMIS.—Hungarian or smooth brome-grass is distributed throughout Minnesota as a cultivated crop and as an intruder along roadsides.
In the fall of 1938 a specimen of Bromus inermis Leyss. which exhibited a peculiar bulbiferous state was submitted to me by Mr. Clemens Kaufman. This appears to be a new form and is being proposed as such at this time.
BROMUS INERMIS Leyss. forma bulbiferus, forma nova. A forma normali differt inflorescentiis bulbillos gerentibus.
In this form some of the florets are replaced by bulbils. This state is similar to that well known condition in Poa alpina L.!
The TYPE was collected on Cleveland Avenue four miles north of the campus of Minnesota College of Agriculture, Ramsey County, Minnesota, October 11, 1938. The type is deposited in the University of Minnesota Herbarium.—Jouw W. Moors, University of Minnesota.
1 Роа alpina L. forma vivipara (Willd.) Scribner & Merrill. Contrib. U. S. Nat. Herb. 13: 68, June 8, 1910.
Volume 43, no. 505, including pages 1-36 and plates 650-652, was issued 15 January, 1941.
Hovova
JOURNAL OF THE
NEW ENGLAND BOTANICAL CLUB
Conducted and published for the Club, by MERRITT LYNDON FERNALD, Editor-in-Chief
CHARLES ALFRED WEATHERBY LUDLOW GRISCOM Associate Editors STUART KIMBALL HARRIS
Vol. 43. March, 1941. No. 507. CONTENTS:
Additions to Flora of Fishers Island, New York. Harold St. John 77
Pubescent Form of Ceanothus ovatus. J. H. Soper ............. 82
Sparganium glomeratum in Minnesota. Olga Lakela ............ 83
Тегаїо1ор1с урһа ато Оп ИНИ 85
Lewis and Clark: Linguistic Pioneers (Review).
George CINE опет о О н eI 92 Napaea dioica in New England. John P. Brown ................ 94 Moss Flora of North America (Review). D. L. Ordway ........ 95 Research Fellowship for отеп (Notice "ur C T 96
The New England Botanical Club, Ine. 8 and 10 West King St., Lancaster, Pa. Room 1001, 53 State St., Boston, Mass.
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Rhodora
JOURNAL OF
THE NEW ENGLAND BOTANICAL CLUB
Vol. 43. March, 1941.
No. 507.
ADDITIONS TO THE FLORA OF FISHERS ISLAND, NEW YORK
HAROLD ST. JOHN
И
ABOUT two miles off the coast of Connecticut, near New London, is a long, wooded island, Fishers Island. Though remote geo- graphically, it is within the political boundaries of Suffolk County, New York. The island is seven miles in length, with an extreme width of one and one half miles and an average width of one half a mile, and its highest hills are 110 and 120 feet in height. There are salt ponds, salt marshes, sandy or gravelly beaches, several fresh ponds, and the rolling, hilly uplands which have a dense but wind-swept serub or forest.
Published accounts of the botany of Fishers Island are few. Those known to the author are the following, arranged chrono- logically:
Graves, C. B. Notes from Plum Island and Fisher's Island, М. Ү. Bull. Torrey Bot. Club xxiii. 59 (1896).
Evans, А. W. A New Station for Coelopleurum actaeifolium. Torreya xvii. 103 (1917).
Evans, А. W. The Hepaticae of Fisher’s Island. Torreya хху1. 85-86 (1926).
Hanmer, C. C. Plants of Fishers Island. Torreya xl. 65-81 (1940).
The last publication gives the common name, the scientific name, and often the abundance of about 500 species of vascular plants. These are the records of thirty years of collecting by Mr. Hanmer who has long maintained a summer home on the island.
78 Rhodora | Marcu
He refers to the disappearance of numerous species due to drain- ing. It is obvious that the recent development of numerous es- tates, summer homes, hotels, country clubs, ete., have much restricted the native flora.
The writer here presents a supplement to Hanmer's list of the vascular flora, based upon his own collections and upon published records. It is probable that other records may be found in the C. B. Graves herbarium at Connectieut College, New London, which the writer has not had an opportunity to examine.
In 1920 from August 10th to the 13th the author collected on the western end of the island while visiting at the army post, Fort Н. С. Wright. This 1920 collection was made in behalf of the Gray Herbarium, so was left there when the collector resigned to accept a position at the State College of Washington. C. A. Weatherby kindly determined this collection. His determina- tions are used here, except when modified owing to more recent nomenclatorial changes. On September 21, 1939, the author had three hours on the island with an auto for transportation. Тһе collections made on these two short visits total 238 species. Of these 75 are not included by Hanmer. They are listed below. Various other species common to Hanmer's and the author's col- lections have been distributed with different specific names. In general, he followed the names in Gray's Manual, ed. 7, while the author has endeavored to use all corrections or later changes that are valid. These nomenclatorial divergencies are not in- cluded here, as the initiated can readily check them. The author's collections are deposited in the Gray Herbarium, with duplieates in several other eastern herbaria. In the following list is given the scientific name of each species and the author's collection number which represents it.
OSMUNDACEAE Osmunda regalis L. var. spectabilis (Willd.) Gray, 2523.
EQUISETACEAE
Equisetum arvense L. forma nemorosum Braun, 2525. Forma decumbens (G. F. W. Meyer) Klinge, 2526.
LYCOPODIACEAE Lycopodium obscurum L., 20058.
19411 St. John,—Additions to Flora of Fisher’s Island 79
PINACEAE
Juniperus virginiana L. var. crebra Fern. & Grise., 2535. It is probable that the collections recorded by Hanmer as the species belong to this recently described northern variety.
NAJ ADACEAE
Zostera marina L., 2545.
ALISMATACEAE
Sagittaria Engelmanniana J. G. Sm., 2547.
(GRAMINEAE
Agropyron repens (L.) Beauv. f. pilosum (Seribn.) Fern., 2550.
Agrostis perennans (Walt.) Tuckerm., 2551.
Andropogon scoparius Michx. var. frequens Hubb., 20039; 20097.
Var. septentrionalis Fern. & Grise., 20061.
Glyceria striata (Lam.) Hitehe., 2560.
Panicum meridionale Ashe var. albemarlense (Ashe) Fern., 2569; 20041.
P. mierocarpon Muhl., 2571.
P. virgatum L. var. spissum Linder, 2568; 20080.
CYPERACEAE Cyperus filiculmis Vahl var. macilentus Fern., 2595. Eleocharis parvula (R. & S.) Link, 2610. ARACEAE Symplocarpus foetidus (L.) Nutt., 2619.
J UNCACEAE Juncus effusus L. var. solutus Fern. & Wieg., 2628. J. marginatus Rostk., 2629. J. macer S. F. Gray, 2635.
LiLIACEAE Smilax glauca Walt. var. leurophylla Blake, 2645.
SALICACEAE
Populus alba L., 2663, introduced. P. grandidentata Michx., 2662.
P. tremuloides Michx., 2664.
"alix discolor Muhl., 2670.
80 Rhodora | Marci
MYRICACEAE
Comptonia peregrina (L.) Coult. var. asplenifolia (L.) Fern., 2674. JUGLANDACEAE
Carya glabra (Mill.) Spach, 2678.
BETULACEAE Alnus incana (L.) Moench., 2683. Betula populifolia Marsh., 2687. FAGACEAE Quercus velutina Lam., 2693.
POLYGONACEAE Polygonum Convolvulus L., 2701, introduced. P. Hydropiper L. var. projectum Stanf., 20092. P. pensylvanicum L. var. laevigatum Fern., 20071. CHENOPODIACEAE Salsola Kali L. var. caroliniana (Walt.) Nutt., 2710.
CARYOPHYLLACEAE Arenaria peploides L. var. robusta Fern., 20066. CRUCIFERAE Brassica kaber (DC.) L. C. Wheeler, 20085, introduced. DROSERACEAE Drosera rotundifolia L., 2732.
ROSACEAE
Agrimonia striata Michx., 2737.
Potentilla canadensis L. var. typica Fern., 2743. P. pacifica Howell, 20074.
P. simplex Michx. var. calvescens Fern., 2742. Rubus ostryifolius Rydb., 2756.
LEGUMINOSAE Lathyrus japonicus Willd. var. pellitus Fern., 2773.
EUPHORBIACEAE
Euphorbia glyptosperma Engelm. fide Graves, C. B. (Bull. Torrey Bot. Club xxiii. 59, 1896), and Gray, A. (Man. Bot. ed. 7, 546, 1908).
19411 St. John,—Additions to Flora of Fisher's Island 81
ANACARDIACEAE
Rhus copallina L. var. latifolia Engler, 2790.
AQUIFOLIACEAE Ilex verticillata (L.) Gray var. tenuifolia (Torr.) Wats., 2793.
VITACEAE Vitis labrusca L., 2811. GUTTIFERAE Hypericum mutilum L. var. parviflorum (Willd.) Fern., 20099.
VIOLACEAE Viola pallens (Banks) Brainerd, 2813.
ONAGRACEAE Cireaea latifolia Hill, 2825. Ludwigia palustris (L.) Ell. var. americana (DC.) Fern. & Guse 20101. UMBELLIFERAE
Cieuta bulbifera L., 2832.
Coelopleurum lucidum (L.) Fern. Reported (as C. actaci- folium) by Evans, A. W. (Torreya xvii. 103, 1917).
Hydrocotyle sp. A sterile collection, 20059, proved inadequate for specifie identification,
"ium suave Walt., 2837.
ERICACEAE
Gaylussacia frondosa (L.) T. & G., 2849.
Kalmia angustifolia L., 2853.
Rhododendron viscosum (L.) Torr., 2863.
PRIMULACEAE x Lysimachia producta (Gray) Fern., 2870. CONVOLVULACEAE Cuscuta pentagona Engelm., 20090.
LABIATAE
Monarda didyma L., 2890. Prunella vulgaris L. var. lanceolata (Barton) Fern., 2891.
SOLANACEAE Datura Stramonium L., 20076, introduced.
82 Rhodora [Marcu
PLANTAGINACEAE
Plantago elongata Pursh was reported by Graves, С. В. (Bull. Torrey Bot. Club, xxiii. 59, 1896).
RUBIACEAE Mitchella repens L., 2920.
COMPOSITAE
Ambrosia artemisiifolia L. var. elatior (L.) Descourtils, 20073.
Aster paniculatus Lam. var. simplex (Willd.) Burgess, 20047.
A. pilosus Willd., 20083.
Bidens connata Muhl. var. petiolata (Nutt.) Farw., 20098.
Erechtites megalocarpa Fern., 20053. "This is a range exten- sion, westwards from Buzzards Bay, Mass.
Helianthus annuus L., 20044, introduced.
Lactuca canadensis L. var. latifolia Kuntze, 2956.
Xanthium italicum Mor., 20050, introduced.
OSBORN BOTANICAL LABORATORY, YALE UNIVERSITY, New Haven, Connecticut
THE PUBESCENT FORM OF CEANOTHUS OVATUS J. Н. SoPER
WHILE determining and mapping the distributions of some plants in my Southwestern Ontario collections of 1940, I came upon a set of plants which were clearly the pubescent extreme of Ceanothus ovatus Desf., first described by Torrey and Gray.! This has been passing in most Floras and Manuals as var. pubescens Torr. and Gray, but a study of the nomenclature shows that the correct varietal epithet is var. pubescens Watson. The facts disclosed by the investigation are here briefly summarized.
Unfortunately when Torrey and Gray described the pubescent phase of Ceanothus ovalis Bigel.? they did not give it a name, thus rendering the publication incomplete. In 1862 Engelmann took up the Torrey and Gray plant as distinct, for in his Plants of the Upper Missouri there is listed a Ceanothus ovalis, var. pubescens? Although no reference is given to the Torrey and
1 ЕЁ]. N. Amer. 1: 265 (1838)—''B.? leaves (especially on the veins), young branches, and peduncles pubescent.”
2 Fl. Bost. ed. 2, 92 (1824). This name was later replaced by the earlier C. ovatus Desf. Arb. 2: 381 (1809).
з Trans. Amer. Phil. Soc. n. s. 12: 187 (1862).
194341 Lakela,—Sparganium glomeratum in Minnesota 83
Gray description, there is little doubt that the plant in question was the same and that Engelmann was aware of the description. As far as I have been able to determine, the first writer to bring together both the description and the name is Sereno Watson." He cited the Torrey and Gray reference and also the Engelmann reference, and in addition placed the variety under the correct binomial С. ovatus Desf. For this reason the correct name of the pubescent extreme of Ceanothus ovatus, when given varietal rank, should be var. pubescens Watson.
Distribution-maps of both C. ovatus and the pubescent extreme were made from the representative material in the Gray Her- barium. These showed that the pubescent phase, which has gen- erally been considered as having a more western distribution, occurs within the range of the species even as far east as the Great Lakes and northeastern Massachusetts. Since this pubes- cent extreme does not have definite claim to rank as a geographic variety, I am considering it as merely a pubescent form. The essential bibliography is as follows:
CranorHts ovatus Desf., forma pubescens (Wats.), stat. nov. C. ovalis Bigel., 8.? Torrey and Gray, Fl. N. Amer. 1: 265 (1838). C. ovalis, var. pubescens Engelmann in Pl. Upp. Miss. 187 (1862), nomen nudum. С. ovatus, var. pubescens Watson, Bibl. Ind. 166 (1878); Trelease in Proc. Calif. Acad. Sci. 2nd. ser. 1: 108 (1889); and later authors, wrongly ascribed to Torr. & Gray. C. pubescens (T. & б”) Rydberg in Small, Fl. Se. U. S. 751, 1334 (1908); not Ruiz & Pavon, Fl. Peruv. 3: 6, pl. 228 (1802).
Cray HERBARIUM.
SPARGANIUM GLOMERATUM IN MINNESOTA.—A colony of Spar- ganium glomeratum Laest. occurs in Duluth, Minnesota. It is located on Minnesota Point in Sec. 19, in a shallow bog formerly connected with Superior Bay. Cushioned in Sphagna in a Calla- Acorus association the plants grow in water from a few to several inches deep, depending on the fluctuating bay level.
The Duluth colony occasioned much interest to the author whose collections Nos. 1627 and 1703 were distributed as А. fluctuans. Dr. John B. Moyle’s annotation of the sheets in the University of Minnesota Herbarium as S. fluitans Fries, the
* Bibl. Ind. 166 (1878).
84 Rhodora [Marcu
occurrence of which in America was unknown to the author at that time, led to further study of the material in living and dried states.
The Duluth plants have a sparse and delicate root develop- ment with slender rhizomes, apparently different from European plants which are more robust with thicker roots and rhizomes. Moreover, the fruiting heads in the Duluth plants are usually four, sometimes three, whereas in the plants from Sweden and Finland they are usually five, rarely three or six. However, a careful comparative study of flowers and fruits was conclusive as to identity of the Duluth collections.
Regarding the status of this species Professor M. L. Fernald kindly furnished the following critical statement: “5. glomeratum Laest. (1852) is regularly kept up by the European authors. It was maintained by Graebner, is also maintained in Holmberg’s edition of Hartman’s Handbok i. 78 (1922) and in Lindman’s Svensk Famerogamflora 44 (1918), both Lindman and Holm- berg stating that S. fluitans Fries was a mixture or nomen con- fusum, standing primarily upon S. affine Schnitzl. (1845) which, as I have repeatedly shown, is S. angustifolium Michx. (1803).” Furthermore, Prof. Fernald concludes: “Since the Scandinavian authors so generally reject S. fluitans as a nomen confusum and cling to S. glomeratum, it seems to me wiser to follow their inter- pretation."
Pertaining to the occurrence of S. glomeratum in America, according to Prof. Fernald the species was reported by Dr. Har- rison F. Lewis from the Natashaquan River region of Saguenay County, Quebec in Canadian Field Naturalist xlv. 178 (October, 1931). Meinhausen's statement in part, “Aus Nord-Ameriea . . ." in Mel. Biol. Acad. St. Petersbourg XIII. (1893) 389, is based upon S. simpler Huds. var. fluitans Engelm., synonymous with S. fluctuans (Morong) Robinson.
In addition to the Duluth specimens there is a single sheet in the University of Minnesota Herbarium labeled as follows: "Lake Ithasea, Minn., July 1893. A. B. Aiton." Obviously the locality is Lake Itasca where the plants have not been dis- covered since.
The author is indebted to Dr. Н. A. Gleason, New York Botanical Gardens, and to Dr. C. O. Rosendahl, University of
1941] St. John, —Teratologie Typha 85
Minnesota for herbarium and library facilities; to Dr. F. J. Hermann, Bureau of Plant Industry for cheeking distribution data; to Prof. M. L. Fernald for the needed data on the specific status and records of distribution. —OLGa LAKELA, State Teachers College, Duluth.
TERATOLOGIC TYPHA HAROLD 8r. JOHN
THE occasional finding of an abnormal specimen of some well- known plant is an incident of decided interest to the collecting botanist. From that time on, he watehes with keener eyes for a repetition of similar monstrosities within that particular species, and soon he ean add it to his list of those species whieh commonly produce teratologie individuals. All of us have such a list of a score or more names, often it is only a mental list, but it is nevertheless available for quick and ready reference. A record of some of these observations may prove of great value to those active in tracing the phylogeny of the species.
During the latter part of August and early September, 1917, Miss Grace M. Bryant noticed and collected some Cat-tails that were decidedly out of the ordinary. Nearly all of these are shown in the accompanying photograph. On September 14th Miss Bryant and the author visited the locality together. It is along both banks of Alewife Brook between Massachusetts Avenue and the first bridge, a railroad bridge, a quarter of a mile distant to the south. The stream here forms the boundary between Cambridge and Arlington. Both Cat-tails, Typha latifolia L. and T. angustifolia L., occur here along the banks of the sluggish stream, but the latter predominates often form- ing a continuous fringe.
Three abnormal plants of T. latifolia were observed. One is shown to the far right in the photograph. The staminate part of the spike has fallen. Below the broken tip of the stem is the normal pistillate part, while five inches below this and partially sheathed in the axils of the two uppermost leaves is an additional pistillate part. Such specimens with two superposed pistillate parts of the inflorescence have been found before in North America as is indicated by a specimen in the Cray Her- barium and one in the Herbarium of Yale University. The
86 Rhodora [ Marcu
Fig. 1. From the left, five plants of Typna ANGUsTIFOLIA. L. with juxtaposed pistillate spikes; at the right one TypHa ТАТІЕОІЛА L. with super- posed pistillate spikes.
1941] St. John, —Teratologie Typha 87
former is without any statement of locality; it was collected by William Boott, July 25, 1869. The latter was found at New Haven, Connecticut, in 1858 by Prof. D. C. Eaton. Dr. С. Е. Koch deseribed! a somewhat similar condition in a plant with three superposed segments of the spike, the lowest being pistillate, the uppermost being staminate, while the median contained both staminate and pistillate flowers. Similar plants of T. latifolia were described and illustrated? by Dr. R. v. Soo. The other two abnormal specimens of T. latifolia observed along Alewife Brook also had two instead of one pistillate part of the spike, but on these plants the two pistillate parts were juxtaposed or twinned instead of being superposed. On their nearer faces each has a line which seems to represent a line of sterile tissue below and originally the line of cohesion of the two now separate parts. A single stem bears these two but as the fruits and their attendant bristles develop they exert a pressure upon each other causing the splitting of the stem both above and below. The upper part usually gives way utterly, thus reducing the tension on the lower part. In the Herbarium of the New England Botanical Club is just such a specimen as these, MASSACHUSETTS: bog on Day Farm, Scituate, Aug. 13, 1899, Emile F. Williams. Also, Mr. К. A. Ware writes me that he found a specimen of just this type in Provincetown, Massachusetts. American authors have occasionally recorded this sort of monstrosity, “The? same col- lector [name not given] has from Carlstadt, N. J., Typha latifolia, in which there are several small spikes arising side by side from the top of the culm, and which appear to be due to fission.” In the same volume is another record, “Mr. Bicknell displayed
specimens of Typha latifolia, L., showing fission of the spike." Half a dozen years later it was again recorded,’ “David F. Day stated . . . that double fertile spikes are not uncommon in T. latifolia.” European observers, as V. Borbas,® L. Ducamp,” and J.-B. Géze.^