Guide to Eastern Ferns [Reprint 2016 ed.] 9781512808568

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Guide to Eastern Ferns

Guide to Eastern Ferns By E D G A R T. W H E R R Y Professor of Botany, University of Pennsylvania Research Fellow, Academy of Natural Sciences of Philadelphia

Philadelphia UNIVERSITY OF PENNSYLVANIA 1948

PRESS

Copyright 1942 EDGAR T. WHERRY

Second Edition, Second Printing March 1948

CONTENTS PAGE

Introduction Preface to second edition Glossary of technical terms Classification Keys to families, genera, and species Illustrations and descriptions Fern cultivation Index to genera and species

iii

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1 3 5 18 21 40 236 244

GUIDE TO EASTERN FERNS T H I S guide has been prepared to fill a need for an aid in the identification of the ferns and fernallies of the region from Pennsylvania and New Jersey to Virginia, of convenient size for field use, and, though non-technical, free from errors and misidentifications such as occur in several current fern books. The data as to each species are presented under five headings: HISTORY ; FEATURES, especially those not brought out by the drawings; HABITAT, ascertained by extensive field work; RANGE, as known to date; REMARKS, comprising methods of distinguishing similar species, and corrections of certain errors often made in amateur writings on these plants, which, once published, get copied by one uninformed writer after another indefinitely. Every effort has been made to furnish in the illustrations and text reliable information on the ferns and their allies of this region, with special attention to making their identification easy. Since most of the species range over a far wider territory, the book should prove useful in adjacent regions as well. —1—

About 100 species of ferns and fern-allies are covered. The technical nomenclature is brought up to date, the names being somewhat simplified. Descriptive common names are used whenever practicable. Within each group the plants are taken up in an evolutionary sequence. To identify an unknown specimen, first place it in the proper order, family, and genus, and then check against the drawings and descriptions. Keys are being added in the second edition to facilitate this.

PREFACE TO THE SECOND EDITION The first edition of this Guide was published in November, 1937, and during the succeeding four years nearly 2000 copies were sold. In preparing a new edition, advantage has been taken of constructive criticism received from reviewers and users, especially as to how to make the book more useful to the beginning student of ferns. The changes made comprise: Addition of keys on pages 21 to 39 inclusive, in response to numerous requests; (none had been included in the first edition because the writer had observed that most people identify unknown plants by comparison with pictures). Remaking of 25 of the figures which proved not to bring out the features of the plants as well as could be desired. These were mostly drawn by Miss Inez Renningèr, who was engaged under two W. P. A. projects in preparing illustrations for a forthcoming Flora of Pennsylvania, and are published here by courtesy of the Department of Botany of the University of Pennsylvania. A few needed to complete the series were kindly contributed by W. H. Wagner, Jr. —

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Inclusion of several species and varieties discovered to grow in our area subsequent to the publication of the fh;st edition. Revision of the nomenclature and insertion of alternative names widely used in other works. Correction of data as to history and range. During the past four years new occurrences of several of the included species have been found, while, on the other hand, some previously accepted reports have proved to be erroneous. Rearrangement of certain genera to bring the sequence into better accord with present-day views as to their evolutionary relations, following, with minor modifications, the plan of C. Christensen as outlined in Verdoorn's Manual of Pteridology of 1938. And expanding the index so that it covers not only genera but also species. These changes will, it is hoped, make the Guide to Eastern Ferns more useful than before in the identification and study of these plants. A wide margin has been left at the bottom of the pages to enable notes to be written in by users of the book. The writer will appreciate the receipt of new data on any of the species.

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GLOSSARY OF TECHNICAL TERMS Acid: Descriptive of soils which show certain features recognized by chemists as connected with the presence of an excess of hydrion, or electrified hydrogen; opposed to alkaline. Acuminate: Tapering to a point. Acute: Ending in a sharp point or angle; opposed to obtuse. Aestival: Produced in Summer. Alkaline: Descriptive of soils containing an excess of hydroxylion, or electrified hydrogenoxygen groups; opposed to acid. Alluvial: Descriptive of areas along streams which sometimes overflow, spreading out sheets of gravel, sand, or mud. Annual: Lasting but one year. Appressed: Sloping upward and tending to lie parallel to a stalk or margin. Areole: The roundish space between veins which separate and then join together again. Attenuate: Tapering gradually to a long tip. Auricle: A more or less ear-shaped basal lobe. Axil: The angle between a branch, leaf, or frond and the supporting stalk. —

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Blade: The broad, flat portion of a leaf, frond, or division of «ither; also termed a lamina. Bog: An open, wet area, supporting mosses and low-growing vascular plants, often quivering when walked upon; soil usually intensely aeid. Bryophyta: The plant-group comprising Mosses and Liverworts. Calcareous: Descriptive of soils or rocks containing lime (calcium oxide) in relatively large amount, and in a form available to plants. Caudate: Tapering to a long tail-like tip. Chlorophyl: The green coloring matter of plants. Cilia: Slender hairs ranged along a margin. Circumboreal: Descriptive of plants which occur in lands lying adjacent to the arctic regions. Circumneutral: Descriptive of soils which are at most so slightly acid or alkaline that they may be regarded as essentially neutral. The reaction-range concerned is active acidity 10 to active alkalinity 10 (pH 6 to 8). Cm.: Abbreviation for centimeter, a metric unit about % inch in length. Compound: Regularly divided one or more times. Cone: A spike of sporophyls and sporangia. Cordate: Descriptive of a blade-base which is con—

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ventionally heart-shaped, that is, having two rounded lobes with a notch between. Cremate: Shallowly cut into rounded teeth. Crenulate: Finely or obscurely crenate. Cuneate: Wedge-shaped. Deciduous: Withering away at the close of a growing season, especially in Autumn. Deflexed: Bent abruptly downward. Deltoid: Shaped like a triangle lying on one side, which is usually shorter than the others. Dimorphic: Descriptive of fronds or of pinnae two sorts of which appear on a single plant. Division: The general term for the parts into which a leaf or frond may be divided; comprises pinnae, pinnules, segments, lobes, etc. Dorsal: Borne on the back of a leaf, frond, stem, or other structure. Eglandular: Descriptive of hairs which lack a gland at the tip. Ellipse: A curved, closed figure with one axis longer than the other, and widest midway between the ends. Ellipsoid: A curved solid, at least one section of which is an ellipse. —

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Elliptic: Descriptive of the outline of a frond or division which is widest in the middle. Entire: Descriptive of a margin which is uninterrupted by cutting, lobing, or toothing. Evergreen: Surviving the Winter without losing chlorophyl or disintegrating. Exindusiate: Lacking an indusium. F.: Abbreviation for forma. Falcate: Obliquely curved like a scythe-blade. Fern-ally: A vascular spore-bearing plant with various features unlike those of true ferns; comprises Clubmosses, Spikemosses, Quillworts, and Horsetails. Fertile: Bearing spores; opposed to sterile. Form (Latin forma, abbreviation f.) : A minor variant of a species, without special range. Free: Descriptive of veins which after separating do not unite again; opposed to areolate. Frond: The technical term frequently used for a fern leaf; it consists of a petiole or stipe supporting a blade. Funnelform: Circular in cross-section and flaring toward one end. Gametophyte: The stage in the life-cycle of a fern on which sexual organs are produced. —

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Glabrous: Smooth, free from hairs or scales. Gland: A globule of waxy nature, usually tipping a hair; in ferns often a source of fragrance. Glaucous: Covered with a thin bluish film. Hirsute: Bearing more or less stiff hairs. Humus: The amorphous organic matter of soils. Hybrid: A plant resulting from the fertilization of the female cell of one species by the male cell of another, usually combining the characters of the parents, but often anomalous in one or more respects, and frequently sterile. In ferns, hybridization may occur between members of different, though closely related, genera; hybrid plants are usually sterile, but fertility may develop in occasional individuals. Incised: Cut in a coarse, irregular fashion. Indusium (pi. -a) : A thin sheet of tissue covering a fern sorus, especially when young. It may be attached basally, laterally, or centrally. Inferior: Borne on the under side or lower part of a structure. Inframedial: Descriptive of sori which lie closer to the midvein than to the margin of the division on which they are borne. Internode: Part of a stem between two nodes. —

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Lanceolate: Descriptive of the outline of a leaf, frond, or division which is conventionally lance-shaped, that is, narrow at base, rapidly widening somewhat, and then gradually tapering to the tip; to be correctly so-termed, an object must be widest between the base and middle, and thus intermediate between deltoidelongate (wider at base) and narrow-elliptic (at middle). Grades by widening into ovate. Lateral: Having a position at one side. Lenticular: Shaped like a doubly convex lens or a sphere compressed in one direction. Linear: Long and narrow, with parallel sides; grades into elliptic, lanceolate, oblong, etc. Lobe: A small, more or less rounded division of a leaf, frond, or part thereof, bounded by narrow notches extending part way to the midvein. Lunate: Shaped like the quarter moon. Lycosphen: A new term for fern-ally, formed from the group-names Lyco- and Sphenopsida. Meadow: An open moist area supporting grasses and other small herbaceous vascular plants. Mediacid: A degree of acidity about as high as ever met with jn normal soils, comprising active acidity values 100 to 1000 (pH 5 to 4). —

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Medial: In the middle; descriptive of sori lying midway between midvein and margin of the division on which they are borne. Megaspore: A large, female, spore. Membranous: Relatively thin in texture. Meier: A metric unit of measurement, slightly under 40 inches or 3Ys feet in length. Micros-pore: A small, male, spore. Midrib: A heavy vein which extends up the middle of a simple leaf, frond, or division. Midvein: A relatively slender midrib. Mm.: Abbreviation for millimeter, a metric unit about 1/25 inch in length. Mucronate: Tipped by an abrupt point. Node: The place on a stem at which branches, leaves, fronds, etc., are borne. Oblanceolate: Inverted lanceolate, that is, Widened above instead of below the middle. Oblong: Somewhat longer than wide, with the long sides parallel; grades to linear or elliptic. Obovate: Inverted ovate, markedly widened above instead of below the middle. Obtuse: Blunt or rounded off at the tip; opposed to acute or acuminate. —

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Ovate: Descriptive of a blade or division which is conventionally egg-section shaped, that is, relatively narrow at base, then markedly widened, and finally tapering rapidly to the tip; grades into lanceolate (less widened), deltoid (widest at base), and elliptic (wide at middle). Palmate: Separated into radiating divisions. Panicle: A compound, open grouping of reproductive structures; in ferns, of sporangia. Peduncle: The stalk supporting a spike or cone. Persistent: Surviving unfavorable seasons without disintegrating. Petiole: A leaf-stalk; in ferns termed a stipe. Pilose: Covered with long soft hairs. Pinna (pi. -ae) : The primary division of a frond, set off by openings extending to the rachis. Pinnate: Divided into pinnae. 2-pinnate signifies that the pinnae are in turn divided into pinnules and 3-pinnate that the latter are again divided into second-order pinnules (divisions). The pinnateness diminishes upward. Pinnule: The secondary (or tertiary) division of a frond, set off by openings extending to a branch-rachis. This term is often mistakenly applied by amateurs to a segment or a lobe. —

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Prothallium: The gametophyte or sexual stage in the life-cycle of ferns and fern-allies. Pubescent: General term for hairy. Quadrate: Square in cross-section. Rachis: The axis of a pinnate frond, extending from the top of the stipe through the blade, ultimately becoming a midrib. .Reaction: The general term covering soil acidity and alkalinity. Receptacle: A support for reproductive organs. Reflexed: Bent abruptly downward or backward. Revolute: Turned under at a margin. Rhombic: Conventionally diamond-shaped, that is, 4-sided with the angles at the ends acute; grades into elliptic and ovate. Rootstock: An underground, often horizontal stem; in ferns the fronds may arise from it in a terminal cluster, or at intervals, in a row. The scales it bears are of diagnostic value. Segment: A division of a frond set off by wide openings; in the true ferns, these do not extend all the way to the midrib. Serrate: Cut, like a saw, into oblique acute teeth. Serrulate: Finely or obscurely serrate. — 13 —

Sessile: Lacking an individual supporting stalk. Sheath: A more or less tubular structure surrounding a stem or stalk. Simple: Undivided, or all in one piece. Soral: Pertaining to a sorus. Sorus (pi. -i): A patch of fertile tissue on the under side of a fern frond. Sphagnum: A genus of mosses growing characteristically in bogs or open swamps and producing an intensely 'acid humus soil known as peat. Spike: A close grouping of sporangia and/or sporophyls along an elongated axis. Spinulose: Bearing small pointed structures. Sporangium (pi. -a): The case in which spores are borne; usually a tiny compressed-spherical yellowish thin-walled capsule. Spore: A minute spherical, ovoid, or tetrahedral thick-walled cell containing dense protoplasm, capable of surviving long exposure to unfavorable environmental conditions, and, in ferns, germinating to form a prothallium. Spore-case: Alternative term for sporangium. Sporophyl: A leaf which bears or surrounds a sporangium; also often termed a fertile leaf. Sporophyte: The spore-producing stage in a life—14 —

cycle; in ferns and fern-allies, the ordinary plant. Alternates with a tiny gametophyte. Stalk: The general term for a rod-like supporting organ; comprising stem, petiole, stipe, etc. Sterile (as to a plant-part): Producing no spores; opposed to fertile. Sterile (as to soil): Poor in available plant-foods, especially soluble nitrogen compounds. Stipe: The preferred term for the stalk supporting the blade in a fern frond. Sub-: A prefix signifying incompletely, imperfectly, to a lesser degree, etc. Subacid: Descriptive of soils showing a moderate degree of acidity, comprising active acidity values 10 to 100 (pH 6 to 5). Subalkaline: Moderately alkaline, comprising active alkalinities 10 to 100 (pH 8 to 9). Superacid: An extreme degree of acidity, met with only exceptionally; active acidity 1000 or more (pH 3 or less). Superior: Borne on the upper side or part of a structure. Swamp: A wet area well grown up with large shrubs or trees; grading into a bog (with less woody growth) or a wet woods (with denser trees).

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Talus: Loose rocks at the base of a cliff, sloping down away from the rock face. Ternate: Separated into three more or less equal divisions, and correspondingly approaching a deltoid outline; grades into multiply pinnate. Truncate: Cut off abruptly or squarely. TJndulate: Descriptive of a margin which is uninterrupted by notches, but is wavy. Variety (abbreviation v.) : A variant of a species in which the differences are too slight to justify species independence, yet which exhibits more or less ecologic or geographic isolation. Some authors use the term subspecies for such plants instead. Variety names are here set off by parentheses, and may be ignored by any one interested primarily in the species. Vascular: Descriptive of a stem, leaf, etc., traversed by elongate conducting cells or veins; all ferns and fern-allies are vascular plants. Vernal: Appearing in Spring. Whorl: A grouping of branches or leaves in radiating fashion around a node on a stem. Woolly: Bearing long kinky matted hairs. —

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lobe pinnule rachis pinna

I

stipe l rootstock root

sca e

A Fern Frond, showing the Principal Parts

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CLASSIFICATION FERNS

Grapefern Order: Succulent plants with an erect stem forking into a leafy sterile segment and a fertile segment bearing paired or grouped globular sporangia. Genera, Botrychium (p. 41), with distinct sporangia and often compound spikes; and Ophioglossum (55), fused sporangia in simple spikes. Fern Order: Plants producing leaves known as fronds, which are sometimes entire but usually more or less divided according to a dominantly pinnate plan, and which bear variously disposed groupings of minute sporangia. Divided into: Flowering Fern Family, with panicles of globular sporangia; genus, Osmunda (59). Climbing Fern Family, with double rows of subglobular sporangia on narrow leaf-divisions; two genera, Lygodium (65), with large twining fronds, and Schizaea (67), tiny grass-like ones. Water-fern and Pond-fern Families (69), comprising a few introduced aquatics. Filmy-fern Family (71)", with clustered globular sporangia on marginal bristles; Trichomanes. —

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Fern Family, with stalked lenticular sporangia in sori on margins or backs of fronds. Grouping: Hay-scented Fern Group: sori marginal, globular; genus Dennstaedtia (73). Bracken Group: sori marginal, elongate, borne on heavy veins; genus Pteridium (75). Rockbrake Group: sori submarginal, linear; genera Adiantum (77), Cryptogramma (81), Cheilcmthes (83), and Pellaea (89). Beadfern Group: sori circular, elevated; indusium inferior; gen. Pteretis (93), Onoclea (95). Cliff-fern Group: sori circular, not elevated; indusium inferior; genus Woodsia (97). Brittle-fern Group: sori circular, with lateral hood-like indusium; genus Cystopteris (103). Chainfern Group: sori on inner netted veins; genera Woodwardia (109) and Lorinseria (111). Spleenwort Group : sori along diverging veins; Phyllitis (113), Asplenium (115), Asplenosorus (133), Camptosorus (135) and Athyrium (137). Woodfern Group: sori circular; indusia superior or rarely lacking; genera Dryopieris (145), Polystichum (165), Thelypteris (169), Phegopteris (175), and Curr'ania (179). Rockcap-fern Group: sori circular, large, without indusium; genus Polypodium (183). — 19 —

LYCOSPHENS or FERN-ALLIES These plants are grouped together f o r convenience, although actually not very closely related to ferns (being much more primitive), nor to one another. Many of them resemble mosses, b u t are distinguished by the presence of a vascular system and grouped sporangia. They fall into two groups differing in stem and spore characters. Clubmoss G r o u p : Moss-like plants with solid continuous often branched stems clothed with small overlapping leaves, or delicate sedge-like aquatics. Sporangia rounded, borne on or in the leaf-bases. Genera, Lycopodium (187), with fertile leaves in zones on stem, or in cylindric cones; Selaginella (211), with fertile leaves in small quadrate cones, and spores of two sizes; and Isoetes (215), sterile and fertile leaves in sedgelike rosettes, and two sizes of spores. Horsetail G r o u p : rush-like plants with porous jointed stems, lacking normal leaves but bearing sheaths at the joints, and often green branches. Fertile stems tipped by a cone of tiny mushroomshaped scales bearing pendent ellipsoidal sporangia. One genus, Equisetum (221). Anyone who p r e f e r s to identify plants by keys will find these on the following 19 pages. —

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KEY TO FAMILIES PLANTS bearing a few relatively large leafy segments or leaves (fronds) FERNS ROOTSTOCK sending up succulent stems bearing a lateral leafy segment and a terminal fertile one with spiked or panicled globular or ellipsoid sporangia. OPHIOGLOSSACEAE ROOTSTOCK sending up fronds which bear groups of globular to lenticular sporangia. SHAPE OF SPORANGIA more or less globular. SPORANGIA borne in compound clusters. OSMUNDACEAB SPORANGIA borne in double rows on narrow frond-divisions; plants rather delicate. SCHIZAEACEAE SHAPE OF SPORANGIA more or less lenticular. CLUSTERS OF SPORANGIA (sori) borne on marginal bristles; plants very delicate. HYMENOPHYLLACEAE CLUSTERS OF SPORANGIA (sori) borne on blades PQLYPODIACEAE (The genus Dennstaedtia, here placed in this family, has the sori strictly marginal, and so may really belong to a Tree-fern Family.) — 21 —

(Representatives of two families of water-ferns, MARSILEACEAE and SALVINIACEAE, are introduced in our region; for them see p. 69.) moss-like, bearing many small leaves; sedge-like, with medium-sized quill-shaped leaves; or rush-like, without normal leaves. LYCOSPHENS "FERN-ALLIES" STEMS solid, bearing green leaves; sporangia borne on or in the bases of sporophyls. SPOROPHYLS in zones on stems or in cones. SPOKES all alike; zones of sporophyls or cones cylindrical. LYCOPODIACEAE 1 SPORES of 2 kinds (J & $ ) ; cones quadrate; plants moss-like. SELAGINELLACEAE SPOROPHYLS in sedge-like rosettes; spores of 2 kinds & $) ISOETACEAE STEMS hollow, bearing sheaths of scales and sometimes green branches at nodes; sporangia borne under scales in cones; spores all alike EQUISETACEAE

PLANTS

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KEY TO GENERA AND SPECIES OPHIOGLOSSACEAE

(Adderstongue Fam.)

distinct, in compound spikes; sterile segment compound (or in B. simplex both at times simple) Botrychium (Grapeferns) PLANT aestival, evergreen; sterile segment ternate-pinnate; pinnae 1- to 3-divided. ULTIMATE DIVISIONS rounded; fruiting aestival ; green in winter B. multifidum. ULTIMATE DIVISIONS more or less elongate; fruiting autumnal; often bronzy in wint. MARGINS finely crenate to shallowly cut into broad teeth B. obliquum. MARGINS deeply cut into divergent oblong to linear teeth B. dissectum. PLANT vernal, deciduous in summer or autumn. SIZE relatively small. STERILE SEGMENT sessile, deltoid, its few pinnae sharp-toothed. B. lanceolatum. STERILE SEGMENT stalked. STALK short; blade oblong to ovate; teeth blunt B. matricariaefolium. STALK long; blade oblong to elliptic, pinnately cut into rounded obtuse lobes or rarely entire B. simplex.

SPORANGIA

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(OPHIOGLOSSACEAE: Botrychium, contd.) large; sterile segment sessile, deltoid, repeatedly pinnate with pointed ultimate divisions B. virginianum. SPORANGIA united, in a simple spike; sterile segment simple, elliptic or essentially so, entire. Ophioglossum (Adderstongues) STERILE SEGMENT obtuse, the veins rather uniformly areolate 0. vulgatum. STERILE SEGMENT mucronate; heavier veins forming large areoles within which fine veins form smaller areoles 0. engelmanni. SIZE

OSMUNDACEAE (Flowering-fern Family) One genus, Osmunda (Flowering-ferns) twice-pinnate, the pinnules well spaced; panicle of sporangia terminal 0. regalis. FROND once-pinnate, the lobed pinnae close-set. PANICLES OF SPORANGIA replacing few medial pinnae; axils of pinnae sparsely hairy; lobes oblong, obtusish 0. claytoniana. PANICLES OF SPORANGIA replacing all pinnae on a few early deciduous fronds; axils of pinnae densely hairy; lobes somewhat lunate, acutish 0. cinnamomea. — 24 — FROND

S C H I Z A E A C E A B (Climbing-fern Family) medium-sized, twining, pinnate: sterile pinnae of 2 palmately-lobed pinnules; fertile compd., terminal. Lygodium (Climbing-fern) One species, L. palmatum. FRONDS small, grass-like; fertile tipped by a tiny "comb" Schizaea (Curlygrass Fern) One species, 8. pusilla. FRONDS

M A R S I L E A C E A E and S A L Y I N I A C E A E (Water-ferns) (The introduced aquatic ferns, members of the genera Marsilea, Azolla, and Salvinia, belong near this point in the evolutionary sequence. For details as to their features see page 69.) HYMENOPHYLLACEAE

(Filmy-fern Fam.)

One genus, Trichomanes (Bristle-fern) One species, T. boschianum. P O L Y P O D I A C E A E (Fern Family) borne on marginal teeth Division A. borne on veins, marginal to dorsal. FROND-MARGINS reflexed over sori Div. B. FROND-MARGINS flat or sub reflexed Div. C.

SPORANGIA SPORANGIA

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Division A. Sori strictly marginal One genus, Dennstaedtia (Hay-scented Fern) One species, D. punctilobula. Division B. Sori submarginal borne on a heavy marginal vein extending around the lobes; fronds coarse, ternately 2- to 3-pinnate Pteridium (Bracken) One species, Pt. latiusculum. SORI borne near the tips of separate veins. MARGINS reflexed in short, spaced, lunate strips; stipe dark. Adiantum (Maidenhairs) RACHIS simple, zigzag A. capillus-veneris. RACHIS straight, then forked A. pedatum. MARGINS reflexed all along the divisions. FRONDS strongly dimorphic, delicate; stipe yellowish Cryptogramma (Rockbrake) One species, C. stelleri. FRONDS barely dimorphic, firm; stipe dark. SORAL BAND narrow, separated into short strips Cheilanthes (Lipferns) BLADE pubescent, especially on back. DIVISION 2-pinnate; pinnules lobed; pubescence hirsute C. lanosa. DIVISION 3-pinnate; pinnules obovate; pubescence woolly C. tomentosa.

SOKI

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BLADE glabrous; division 2-pinnate; southern Cheilanthes alabamensis. SORAL BAND broad, in long strips; blade glabrous Pellaea (Cliffbrakes) RACHIS rough-hairy P. atropurpúrea. RACHIS glabrous P. glabella. Division C. Sori dorsal SPORANGIA borne on elevated discoid receptacles; indusia inferior; fronds strongly dimorphic. STERILE FRONDS pinnate; pinnae lobed; fertile pinnae pod-like

Pteretis (Ostrich-fern) One species, Pt. nodulosa. STERILE FRONDS cut nearly to midrib into spaced opposite lobes; fertile pinnae bead-like. Onoclea (Beadfern). 1 sp., 0. sensibilis. SPORANGIA borne on low receptacles. SORI circular; indusia inferior or lateral. INDUSIA inferior Woodsia (Cliff-ferns) STIPE jointed above base, brownish, hairy; blade eglandular-hairy W. ilvensis. STIPE continuous, bearing scales. BLADE fine-gl'r; stipe pale W. obtusa. BLADE with narrow scales and long glandular hairs; stipe brown. W. scopulina.

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(Division C, sori dorsal, continued) (Subd. with circular sori on low receptacles) INDUSIA lateral ..Cystopteris (Bladder-ferns) STIPE brown ; blade oblong-lanceolate to ovate, 1- to 2-pinnate C. fragilis. STIPE pink; blade elongate-deltoid, pinnate ; pinnae lobed ; frond-backs bearing axillary bulblets C. bulbifera. SORI various; indusia superior or lacking. SORUS-BEARING veins heavy, near midribs. FRONDS barely dimorphic, pinnate; veins areolate near midribs Woodwardia. (Chainfern) 1 sp., W. virginica. FRONDS dimorphic ; lobes alternate, spaced, serrate; veins areolate Lorinseria. (Netvein Chainfern) 1 sp., L. areolata. SORUS-BEARING veins fine, diverging. SORUS-SHAPE linear, straight or hooked. TISSUE evergreen; sori straight; mostly rather small rock ferns. VEINS bearing paired sori in regular rows; blade oblong Phyllitis. (Hartstongue) 1 species, P. scolopendrium. VEINS bearing single sori, or when paired not in regular rows. —

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BLADE compound or lobed; veins ± free. -AspZentitm (Spleenworts) STIPE, and often rachis, dark. BACHIS dark throughout. FRONDS uniform; pinnae opp. RACHIS-COLOR b r o w n ; pin.

cuneate A. trichomanes. RACHIS-COLOR black;

pin.

truncate A. resiliens. FRONDS dimorphic; rachis brown; pinnae truncate, many alt....A. platyneuron. RACHIS dark only $ its length • margin serrate. A. bradleyi. STIPE green toward top. CUTTING 1- to 3-pinnate.

PINN. obov. A.

rutamuraria.

PINN. ovate A. montanum. CUTTING pinnate bel. to lobed. LOWER BLADE-DIVISIONS

der stalked

A.

slen-

trudelli.

LOWER BLADE-DIVISIONS b r o a d

stalked A. pinnatifidum. (Hybrids combining the features of pairs of the above species are occasional.)

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(Division C, sori dorsal, continued.) (Subd. with sporangia on low receptacles, indusia superior, sorus-bearing veins fine, diverging, sori linear, and tissue evergreen.) BLADE lobed to entire; veins areol. MARGIN irregularly lobed. Asplenosoras. (Walking-spleen wort) 1 species, A. ebenoides. MARGIN essentially entire; blade deltoid-attenuate, rooting at t i p ; stipe green Camptosorus. (Walking-fern) 1 species, C. rhizophyllus. TISSUE deciduous • sori curved; large woods plants. Athyrium (Lady-ferns) CURVING OF SORI slight; blade pinnate. PINNAE entire A. pycnocarpon. PINNAE lobed A. thelypteroides. CURVING OF SORI marked; blade 2-pin. BLADE widest near middle; stipe with many dark scales; spores yellow, granular A. angustum. BLADE widest near base; stipe with f e w light scales; spores blackish, wrinkly A. asplenioides.

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SORUS-SHAPE

circular.

firm; sori large; tissue more or less persistent; stipe bases and rootstock stout. INDUSIUM kidney-shaped Dn/opfens. (Woodferns and Swampferns) BLADE 2- to 3-pinnate; pinnules and lobes spinulose-serrate. FRONDS evergreen; indusia and upper rachis glandular; inner pinnules at blade-base shorter than next outward ones (rarely equal) D. intermedia. FRONDS subevergreen to decid. STERILE FRONDS evergreen; indusia and rachis glabrous; inner pinnules at blade-base longer than next outward ones D. spinulosa. STERILE FRONDS, like fertile, deciduous ; indusia glabrous to sparse-glandular; inner pinnules at blade-base much larger than the next outward ones D campyloptera.

INDUSIUM-TEXTURE

— 31 —

(Division C, sori dorsal, continued) (Subd. with sporangia on low receptacles, indusia superior, sorus-bearing veins fine, diverging, sori circular, indusium firm, kidney-shape.) to 2-pinnate; pinnules entire to serrate (spinulose only in D. boottii). BLADE-OUTLINE oblong to ovate. SOEl near margins; tissue evergreen ; lower pinnae elongate deltoid; pinnule-margins entire to lobed D. marginalia. SORI near midribs; tissue deciduous; lower pinnae oblong lanceolate, undulate - margined; pinnule-margins appressed serrate ..D. goldiana. BLADE-OUTLINE oblong to lanceolate or oblanceolate; evergr. PINNAE long-acuminate; lowest long-deltoid, pinnate below, not undulate; sori near midribs to med D. clintoniana. (Hybrids combining the features of various pairs of Dryopteris species are not infrequent.) BLADE 1 -

— 32 —

short-acuminate. serrate; lowest pinnae deltoid, lobed; indusia glabrous D. cristata. MARGINS spinulose; lowest pinnae long-deltoid, pinnate ; glandr. ...D. boottii. INDUS. round Polystichum (Holly-f.) BLADE 2-pinnate; margins serrate; many pinnae fertile P. braunii. BLADE 1-pinnate; margins fine to coarse-serrate; small terminal pinnae fertile P. acrostichoides. INDUSIUM-TEXTURE delicate, or indusium lacking; stipes and rootstock slender. SORI small; tissue deciduous; pinnate. BLADE widest above base; indusia pres Thelypteris (Marshferns) PINNAE

MARGINS

GREATEST FROND-WIDTH b e l .

mid.

broadest at base; sterile veins forked; sori crowded T. palustris. LOWER PINNAE broad at mid.; veins simple T. simulata. GREATEST FROND-WIDTH upward; veins simp. T. noveboracensis. LOWER

PINNAE

— 33 —

(Division C, sori dorsal, continued) (Subd. with sporangia on low receptacles, indusia superior, sorus-bearing veins fine, diverging, sori circular and small, tissue deciduous, and stipes and rootstock slender.) BLADE widest at base; indusia none. CUTTING 2-pinnately lobed or pinnate at base; blades hairy. Phegopteris (Beechferns) MIDRIBS bearing pointed hairs and dark scales; blade narrow P. connectilis. MIDRIBS bearing both pointed and glandular hairs; blade broad P. hexagonoptera. CUTTING 2-pinnate, the lower pinna-stalks jointed at the rachis Currania (Oakferns) LOWER P I N N A E relatively large; stalks glabr C. dryopteris. LOWER P I N N A E relatively small; stalks gland. .. C. robertiana. SORI large, exindusiate; fronds evergreen Polypodium (Rockcapferns) BLADES glabrous P. virginianum. BLADES scaly P. polypodioides. — 34 —

LYCOSPHENS or "FERN ALLIES" LYCOPODIACEAE (Clubmoss Family) (One genus, Lycopodium (Clubmosses)) borne in groups on stems, alternating with groups of sterile leaves. LEAVES elongate-deltoid, entire or nearly so; sporophyls slightly shorter; spores about 0.035 mm. in diameter L. selago. LEAVES oblanceolate, more or less serrate; sporophyls decidedly shorter; spores about 0.025 mm. in diameter L. lucidulum. SPOROPHYLS borne in terminal spikes (cones). STEM creeping or arching, sending up simple peduncles bearing a terminal cone. CONE AND PEDUNCLE densely clothed with similar leaves; stem leaves uniform. LEAVES AND SPOROPHYLS appressed; peduncle 3 to 5 and cone 5 to 8 mm. thick; stem rooting all along L. adpressum LEAVES AND SPOROPHYLS spreading; stem often arched, rooting at the tip. PEDUNCLE 5 to 10 and cone 10 to 15 mm. thick; sporophyls broad-lanceolate, entire or few-toothed L. inundatum.

BPOROPHYLS

— 35 —

( L Y C O S P H E N S , continued) (Genu? Lycopodium, continued) (Group with sporophyls borne in terminal spikes, stem sending up simple peduncles with terminal cone, densely clothed with similar spreading leaves, stem often arched and rooting at tip.) PEDUNCLE 10 to 15 and cone 15 to 20 mm. thick; sporophyls narrow-lanceolate, several-toothed L. alopecuroides. CONE AND PEDUNCLE bearing respectively many broad and few narrow leaves; lateral stem-leaves broad L. carolinianum. STEM creeping on or in soil, Sending up more or less compound branches. BRANCHES forking into a few relatively thick branchlets. CONE solitary, sessile; branches with annual constrictions; leaves firm, with short bristle-tips L. annotinum. CONES grouped (rarely solitary) on a peduncle; branches unconstricted; leaves soft, long-bristled L. clavatum. BRANCHES forking into numerous relatively slender branchlets. HABIT tree-like; leaves spaced; cone single ; peduncle short, leafy. L. obscurum. — 36 —

fan-like; branchlet-leaves coalescing ; cones grouped on scaly peduncles. BRANCHLETS bluish green, cord-like, the leaves on lower side similar to the lateral ones; annual constrictions present; stem deep L. tristachyum. BRANCHLETS yellow-green, narrow-ribbon like; leaves on lower side smaller than the lateral ones. STEM buried in soil; branchlets irregularly disposed, with annual constrictions; peduncles and 1 to 3 cones short L. complanatum. STEM merely in litter; branchlets disposed in regular fans, not constricted ; peduncles and 3 to 4 cones elongate L. flabelliforme.

HABIT

S E L A G I N E L L A C E A E (Spikemoss Family) One genus, Selaginella (Spikemosses) all alike, linear, bristle-tipped; cones definitely 4-angled 8. rupestris. LEAVES of two kinds, the lateral larger, ovate, none bristle-tipped; cones obscurely 4-angled. S. apoda. LEAVES

— 37 —

(LYCOSPHENS, continued) I S O E T A C E A E (Quillwort Family) One genus, Isoeies (Quillworts) (The markings on the megaspores as shown in the plates will serve to distinguish the members of this genus. The two common ones in our region are the emersed I. engelmanni, with the spores covered by prominent angular-areolate ridges; and the submerged I. braunii, with the spores densely spinulose.) E Q U I S E T A C E A E (Horsetail Family) One genus, Equisetum (Horsetails, etc.) evergreen; stem simple or with unequal appressed branches; cone sharp-pointed. STEMS large and coarse; teeth of sheaths more or less deciduous. SURFACE markedly rough; sheaths short, cylindric, soon becoming dark-banded and splitting up E. prealtum. SURFACE moderately rough; sheaths long, expanding upward, darkening or splitting only when very old E. laevigatum. STEMS small and delicate; teeth of sheaths persistent ; sheaths funnel-shaped, light and dark-banded E. variegatum.

TISSUE

— 38 —

TISSUE deciduous; stem simple or often bearing whorls of branches, smoothish; cone blunt. STEMS all green, often tall, stout, and manyridged, varying to slender; sheaths cylindric, tight E. fluviatile. STEMS of two types: fertile at first simple and not green; sterile branched, green; sheaths rather loose and flaring. FERTILE STEMS becoming green and branched after the spores are shed. TEETH OF LOWER SHEATHS long, red-brown,

grouped; branches regularly compound, drooping at tips TEETH

OF LOWER

E. SHEATHS

sylvaticum. short,

light

brown to greenish, separate; branches simple, spreading, the lowest joints short; branch-sheaths with 3 deltoid teeth; northern; rare E. pratense. FERTILE STEMS withering after spores are shed; teeth of sterile stem-sheaths dark brown • branches simple or irregularly compound, ascending, their lowest joints rather long; branch-sheaths with 3 or 4 sharp-pointed teeth; common E. arvense. (For the remaining three species of Equisetum, which may enter our region, see the figures.)

— 39 —

Botrychium multifidum (2 varieties) LEATHER

GEAPEFERN

H I S T O R Y : In 1 7 6 6 Gmelin applied the binomial Osmunda multifida to a Russian species; this was changed to Botrychium multifidum by Ruprecht in 1859. In the region covered by this book two varieties (sometimes classed as independent species, or as subspecies) are now known to occur. The larger one was named v. intermedium by Farwell in 1916, and the smaller v. typicum by the writer in 1942. F E A T U R E S : Plant 8 to 4 8 cm. tall, aestival and evergreen, not becoming bronzy. Sterile segment ternate-pinnate, with the pinnae repeatedly cutlobed, their ultimate divisions round and obtusish. The two varieties differ chiefly in size. H A B I T A T : Cool, mossy meadows, moist thickets, and open woods; soil decidedly acid. R A N G E : A circumboreal fern, ranging from Alaska and the northern Rocky mountains to Newfoundland. Yar. typicum extends south to Wayne and v. intermedium to Berks Co., Penna. R E M A R K S : Fruits in late Summer, often a month earlier than the next-following species; grades into the latter through v. oneidense. — 41 —

Botrychium oiliquum (v. typicum) x l / 3 pinnae of v. oneidense and v. tenuifolium to right — 42 —

Botrychium obliquum (v. typicum) (B. dissectum v. obliquum) COARSE-LOBE GKAPEFERN HISTORY : Discovered in Lancaster County, Pennsylvania, by Muhlenberg, whose description of it was published by "Willdenow in 1810. The varietal name was proposed by the writer, 1942. F E A T U R E S : Plant 1 0 to 4 0 cm. tall, autumnal and evergreen, often becoming bronzy; sterile segment ternate-pinnate; lower pinnae twice and upper once-pinnately lobed, with the ultimate divisions obliquely ovate to oblong-lanceolate, the terminal ones especially long, mostly acutish; margins minutely to moderately toothed. HABITAT : Moist open woods, thickets, and meadows, also invading pastures; soil subacid. RANGE : Centering on the Appalachians, but extending south to upper Florida, and northward to Michigan and Nova Scotia; common practically throughout our region, except in pine-barrens. REMARKS : A rather striking variant, with the tissues tending to remain green in winter, and having shorter and more obtuse lobes, is known as B. obliquum v. oneidense; one with few, littledivided lobes, in lowlands, as v. tenuifolium. — 43 —

Botrychium

dissectum x y s

— 44 —

Botrychium dissectum (v. typicum) LACELEAF GKAPEFERN HISTORY : This fern was discovered in Virginia and sent to Sprengel for identification, being named by him as an independent species in 1804. The varietal name was assigned by Clausen, 1938. FEATURES: Plant 10 to 40 cm. tall, autumnal and evergreen, and often bronzed; sterile segment ternate-pinnate and repeatedly lobed, the lobes divided into divergent linear teeth. HABITAT : Open woods and moist thickets, sometimes invading pastures; frequently mingling with the next-preceding. Soil usually subacid. RANGE: Appalachians of South Carolina and northward to the lower Great Lake region; widely distributed in our area except toward the coast. REMARKS: The next-preceding species has a form with moderately toothed lobes, and as this sometimes approaches the present plant, some confusion as to their interrelationships has arisen, leading to a complex system of nomenclature. The writer prefers to maintain two independent species. The range of B. dissectum is more restricted than that of B. obliquum, especially on the south.

— 45 —

Botrychium lanceolatum (v. angustisegmentum) TRIANGLE GRAPEFERN HISTORY: This is another of the species of grapeferns discovered by Gmelin in Russia, and assigned by him in 1768 to the genus Osmunda; its transfer to Botrychium was made by Angstrom in 1854. The temperate-American plant was considered identical with the European one for many years, but in 1906 Pease and Moore pointed out its varietal distinctness. It was made a subspecies by Clausen in 1938. FEATURES : Plant 5 to 2 5 cm. tall, vernal, often persisting through the growing season; sterile segment high on stem, sessile or nearly so, triangular, ternate-pinnate, the pinnae sharp-toothed. HABITAT : Humus-rich soil in cool woods, hummocks in swamps, and rarely meadows; soil mediacid or subacid. RANGE : The typical representative of the species occurs in all northern lands, the present variety from the Great Lakes to Newfoundland and south to Gloucester Co., New Jersey, Pocahontas Co., West Virginia, and Giles Co., Virginia. R E M A R K S : The herbage is dark green. — 47 —

— 48 —

Botrychium matricariaefolium DAISYLEAF GRAPEFERN H I S T O R Y : The above species name was first suggested by A. Braun in 1843, and taken up by Koch two years later, German material being referred to. Under the impression that the similar American plant is an independent species, "Wood named it B. neglectum in 1847, but our present view is that the two are not sufficiently different for even varietal segregation. F E A T U R E S : Plant 5 to 2 5 or rarely 3 5 cm. tall, vernal, and often withering by midsummer; sterile segment about half-way up the stem, stalked, oblong to narrow-deltoid, pinnate, with the pinnae cut into obtusish lobes. HABITAT : Humus-rich soil in woods and thickets often mingling with the next-preceding species; soil subacid or circumneutral. R A N G E : A rather northern species, in Europe and from Nova Scotia to Washington, extending south in our region to central New Jersey and in the mountains to Rockingham Co., Virginia. R E M A R K S : The herbage is paler green than in the next-preceding species.

— 49 —

Botrychium simplex (3 varieties) D W A R F GRAPEFERN H I S T O R Y : Most circumboreal species have been named in Europe first, but the present one was overlooked there by the early workers, and described from Massachusetts by Hitchcock in 1823. Three varieties enter our region; the original, known as v. typicum; v. tenebrosum, described as a separate species in 1899; and v. laxifolium, only distinguished by Clausen in 1937. F E A T U R E S : Plant 3 to 1 5 cm. tall, vernal, and early withering; sterile segment low on stalk in typicum, higher in others; elliptic and simple to round-lobed or pinnate; short and broad in v. typicum, longer but narrower in v. tenebrosum, and large all around in v. laxifolium; fertile segment simple or barely compound. H A B I T A T : Cool woods, swamps, and open sterile north-facing slopes; soil usually subacid. R A N G E : Circumboreal. The v. typicum extends down to Northampton and laxifolium to Philadelphia Co., Pennsylvania; tenebrosum also to Philadelphia and Howard Co., Maryland. R E M A R K S : Reports further south are unconfirmed. Inconspicuous and easily overlooked.

— 51 —

Botrychium virginianum (v. typicum) RATTLESNAKE PERN

HISTORY: Although this species occurs in Europe and Asia, it was first named on specimens from Virginia by Linnaeus in 1753. His name f o r it was Osmunda virginiana, but it was transferred to the genus Botrychium by Swartz in 1 8 0 1 . The varietal name was proposed b y the writer in 1942. FEATURES : Plant usually 20 to 60 cm. tall, vernal, the leaf membranous and lasting through the growing season; sterile segment large, high on the stalk, ternate, the divisions pinnate with sharply cut-lobed pinnae, resembling the true ferns more than do the other Botrychiums. HABITAT : Woods, swamps, and moist thickets; soil circumneutral to subacid. RANGE: Common nearly throughout the eastern United States and southern Canada, with varieties f u r t h e r north, and in the Old World. The most abundant Grapefern in our region. REMARKS : Sometimes assigned to a distinct genus, Osmundopteris, on the basis of the leaftexture and the development of the bud only partly within the stalk-base.

— 53 —

v. typicum and v. pycnostichum — 54 —

Ophioglossum vulgatum (2 varieties) ADDERSTONGUE F E R N HISTORY : This species was named by Linnaeus in 1753; two of its geographic varieties occur in our region: pycnostichum, described by Pernald in 1939, and typicum, named by the writer in 1942 (incl. v. pseudopodum, Farwell, 1916). F E A T U R E S : Plants 1 0 to 4 0 cm. tall, vernal. Sterile segment short-stalked, obtuse; veins simply areolate; v. typicum tends to have an elliptic sterile blade and globular sporangia; v. pycnostichum an ovate blade and ellipsoidal horizontally elongate sporangia; intermediates between them are, however, common. HABITAT : The v. typicum grows on dry grassy slopes, meadows, and swamps, in acid soils; v. pycnostichum in circumneutral moist woods. R A N G E : The northern v. typicum extends to central New Jersey, southern Pennsylvania, and the Appalachians; v. pycnostichum ranges from coastal New England southward, and was first recognized in lowland Virginia. R E M A R K S : Looks so unlike most ferns that although common it is rarely noticed. A slender New Jersey form has been named f. arenarium. — 55 —

Ophioglossum engelmanni LIMESTONE ADDERSTONGUE H I S T O R Y : Sent from Missouri to Prantl by Engelmann, and named for the latter in 1883. FEATURES : Plant 5 to 20 cm. tall, vernal, shortlived and soon withering, but sometimes sending up new shoots in summer, following heavy rains; sterile segment sessile, elliptic, mucronate; veins of two series, the heavier forming large areoles, within which lie several more or less interlacing finer veins. H A B I T A T : Clayey and gravelly depressions between outcrops of limestone and other calcareous rocks, and thinly wooded alluvial flats, sometimes invading pastures and old fields; soil circumneutral or subalkaline. R A N G E : A southwestern species, ranging from Lower California to Texas and extending eastward and northward along limestone outcrops, to Kansas, Ohio, and Florida; also migrating along the Shenandoah Valley, and found to date up to Frederick and Warren Counties, Virginia. REMARKS : The habitat and the mucronate leaves with two series of veins are distinctive.

— 57 —

C

reLOMAN

Osmunda regalis (v. spectabilis) x % — 58 —

Osmunda regalis (v. spectabilis) ROYAL F E R N H I S T O R Y : Linnaeus in 1 7 5 3 defined Osmunda regalis as comprising both European and Virginian plants; he designated the latter subspecies 3, but Willdenow made it a species, 0. spectabilis, in 1810. That they are too similar to justify this was pointed out by Gray in 1856, when he transferred Willdenow's name to varietal status. F E A T U R E S : Plant usually 50 to 150 cm. tall, the fronds clustered; fronds elliptic-oblong, twice pinnate, the sterile pinnules widely separated; fertile pinnules terminal, bearing dense clusters of sporangia, which become light brown. H A B I T A T : Swamps, bog-margins, moist woods, and springy slopes, or rarely wet cliffs; soil most frequently mediacid, rarely circumneutral. R A N G E : This species is widespread, occurring in many parts of the world. The American variety ranges from Saskatchewan to Newfoundland, Florida, and Texas, also through the West Indies and far down into South America. R E M A R K S : The European variety differs from ours in having the fertile portion black-hairy.

— 59 —

\ Osmunda claytoniana (v. vera) —

60

—

Osmunda claytoniana (v. vera) INTERRUPTED F E R N HISTORY: Clayton collected this fern in Virginia in the early 1700's, and it was named in his honor by Linnaeus in 1753. This original variety was designated vera by the -writer, 1937. FEATURES: Plant usually 7 5 to 1 5 0 cm. tall, the fronds clustered, their stalks woolly when young; fronds elliptic-oblong, pinnate, lacking wool at pinna-bases; sterile pinnae deeply cut into oblong obtuse lobes, lasting through the growing season; fertile pinnae borne in 1 to 5 pairs near middle of frond, short, bearing dense clusters of sporangia, which change from dark green to brown and wither away early, leaving the fronds "interrupted." HABITAT: Moist wooded slopes, swamp-margins, and open thickets; soil circumneutral or subacid. RANGE : This species appears in one variety in eastern North America and another in eastern Asia; ours is an upland plant, ranging from Manitoba to Greenland, northernmost Arkansas and Georgia; common except on coastal sands. REMARKS : Medial fertile pinnae unique. —

61

—

Osraunda cinnamomea (v. typica) CINNAMON F E R N H I S T O R Y : Specimens of this f e r n were sent to Linnaeus from Maryland and named by him in 1753. Fernald termed the original v. typica in 1930. F E A T U R E S : P l a n t usually 7 5 to 1 5 0 cm. tall, the fronds clustered, their stalks brown-woolly; fertile fronds vernal, wand-like, bearing compound groups of green sporangia which become cinnamon-brown and soon wither away; sterile fronds oblong-lanceolate, pinnate, with a t u f t of wool at the base of each p i n n a ; pinnae deeply cut into somewhat lunate though obtusish lobes, lasting through the growing season. H A M T A T : Swamps, bog-margins, stream-banks, damp depressions in woods, springy slopes, and occasionally wet rock-ledges; soil often mediacid, rarely low in acidity. K A N C E : The typical variety ranges throughout eastern United States and Canada, and is common in our region; another is tropical. R E M A R K S : The woo) at the pinna-bases, and the subcrescentic segments, distinguish the sterile f r o n d s from other similar ferns.

— 63 —

C.mOMAM

Lygodium palmatum x y¿ — 64 —

Lygodium palmatum CLIMBINO F E R N H I S T O R Y : This fern was first found by William Bartram in Georgia, but lie failed to {rive it a valid name. Material from New Jersey or Pennsylvania sent in by Muhlenberg was named by Willdenow Ilydroglossum, and by Bernhardi Gisopteris palmata in 1801. Swartz transferred the species to his genus Lygodium five years later. F E A T U R E S : Rootstock slender, sending up scattered fronds usually 50 to 100 cm. tall, their stipe and rachis twining around shrub stems; fronds pinnate, the sterile pinnae with two palmate pinnules; fertile pinnae terminal, smaller than the sterile, repeatedly pinnate. H A B I T A T : Wet sphagnous thickets, moist open woods, and springy banks; soil intensely acid. R A N G E : Scattered through the eastern states, Florida to central Tennessee, southern Ohio, eastern New York, & southern New Hampshire. In our region rare southward, but frequent in the northeastern Pennsylvania mountains, and across New Jersey, from Camden to Monmouth counties. R E M A R K S : The climbing habit is unique. — 65 —

Schizaea pusilla about natural size

—

66

—

Schizaea pusilla CURLY-GRASS F E R N H I S T O R Y : Discovered at Quaker Bridge, Burlington Co., New Jersey, and named by Pursh in 1814. F E A T U R E S : The least fern-like of all our species. Rootstock slender, sending up a clump of tiny fronds, 2 to 15 cm. tall; sterile fronds linear, curled and twisted, subevergreen; fertile fronds autumnal, overtopping the sterile, with a slender kinked stipe and a pinnate blade a few mm long, forming a comb-like structure. H A B I T A T : Moist, sterile, intensely acid soil on hummocks of humus or of sphagnum moss, and in wet grassy depressions; in New Jersey usually under white-cedar trees. R A N G E : The genus Schizaea is widespread over the southern hemisphere and the tropics; at some past geological period this single species migrated northward and developed hardiness. This evidently occurred on a land-mass lying east of our present coast, from which it spread westward into New Jersey, Nova Scotia, and Newfoundland, the land ultimately sinking beneath the sea. R E M A R K S : Our most un-fernlike fern. — 67 —

Marsilea quadri/olia ^oUa caroliniana * isalvinia rotundijoha —

68

—

Marsilea quadrifolia WATERCLOVER F E R N

HISTORY : Linnaeus named this plant from native material in 1753. In some unknown manner it became introduced into America, being first found in Connecticut in 1862. FEATURES : Plant aquatic, the rootstock slender, sending to the water-surface a row of fronds; blade divided into 4 nearly equal obovate pinnae ; fertile segment short-stalked, bearing 2 ellipsoidal hollow structures with sori inside. HABITAT : Ponds and lakes; soil circumneutral. RANGE: Introduced in a few ponds in Maryland, New Jersey, Pennsylvania, etc. REMARKS: TWO other aquatic ferns have been introduced locally in our region; these float free on the surface of the water in pools, ponds, and ditches, and are known as Pond-ferns. Carolina Pond-fern, Azolla caroliniana, forms groups of tiny elliptic leaves; South American Pond-fern, Salviiiia rotundifolia, has larger, more separated leaves, with coarse hairs on the upper surface. They are subtropical plants and are not likely to persist here long. — 69 —

VW Trichomanes boschianum x 1/2 Hymenophyllum gametophyte to left x — 70 —

Trichomanes boschianum APPALACHIAN BRISTLE-FERN HISTORY : When discovered in Alabama in 1 8 5 2 regarded as identical with the subtropical T. radicans, but named as distinct by Sturm, 1861. F E A T U R E S : Rootstock slender, black, creeping over rock surfaces; f r o n d s oblong-lanceolate, 10 to 20 cm. long, often pendent, only one cell thick, deeply cut into pinna-like segments which in t u r n are divided in a lacy p a t t e r n ; spores borne in clusters on marginal bristles, surrounded by a funnel-shaped sheath. HABITAT : On vertical or overhanging surfaces in damp shaded recesses in sandstone and similar rocks; soil strongly acid. RANGE: Alabama (or Florida?) to southern Illinois and Ohio, j u s t entering our area in W a y n e Co., West Virginia, where found in 1938. R E M A R K S : F r o m the mountains of Virginia west to Ohio and Kentucky there occurs in cavernous sandstone the gametophyte of another filmy-fern, consisting of branching ribbons with marginal bud-like structures, by which it reproduces. I t has not yet been observed to form a sporophyte, b u t may be a Hymenophyllum.

— 71 —

Dennstaedtia punctilobula HAY-SCENTED F E R N HISTORY : Michaux discovered this fern in Canada and named it Ncphrodium punctilobulum in 1803. It was later mistakenly transferred to the tropical genus Dicksonia, an assignment accepted by American botanists for many years. In 1857, however, Moore showed it to belong in Bernhardi's genus Dennstaedtia, and this view is now coming to be accepted. FEATURES : Rootstock slender, bearing a row of fronds usually 40 to 80 cm. tall; stipe lustrous brown, rachis pale green; fronds thin-textured, lasting through summer, lanceolate, twice-pinnate, the pinnules sharp-toothed; covered with fine hairs, many of them tipped with glands containing fragrant wax; sori small, borne on teeth. H A B I T A T : Dry, open woods, rocky slopes, and rarely swamps; unpalatable to animals and becoming abundant in sterile pasture-land; soil acid. R A N G E : Southern uplands, Georgia to Arkansas, north to Minnesota and Nova Scotia. Common in sterile situations throughout our region. R E M A R K S : Pubescence and sori distinctive. — 73 —

C TEL DM AN Pteridium latiusculum (v. verum) x % a second-order pinnule of v. pseudocaudatum above to left. — 74 —

Pteridium latiusculum (2 varieties) (Pt. aquilinum v. latiusculum) EASTERN BRACKEN HISTORY: Linnaeus named the European Bracken Pteris aquilina in 1753. Seven years later Scopoli placed it in a genus Pteridium, on the basis of the presence of a special indusium. The east-American relative was named Pteris latiuscula by Desvaux in 1827, and Hieronymus, made the above combination in 1914. The original v. was named by the writer verum in 1937. F E A T U R E S : Rootstock stoutish, producing a row of firm but deciduous fronds usually 40 to 80 cm. tall; stipe brown at base; blade coarse, ternately 2-pinnate, the ultimate divisions oblong; sori forming continuous marginal strips, sparse-hairy. H A B I T A T : Open woods, dry slopes, gravel banks, and rarely bog-margins. Soil highly sterile, but varying in reaction; most often strongly acid but locally, in the north, circumneutral. R A N G E : Newfoundland to "Wyoming, Arizona, and Florida. Common over our region in barrens. R E M A R K S : A glabrous phase, with linear terminal pinnule-divisions, growing chiefly at low altitudes, is known as v. pseudocaudatum. — 75 —

«HSntr.

Adiantum

capillus-veneris x

— 76 —

Adiantum capillus-veneris SOUTHERN MAIDENHAIR HISTORY : Linnaeus obtained this fern in southern Europe and named it in 1753. FEATURES : Rootstock slender, producing a row of glabrous drooping subevergreen fronds usually 15 to 45 cm. long; stipe lustrous blackish brown, passing into a zigzag rachis; blade ovatelanceolate, 2 to 3-pinnate below and 1-pinnate toward the tip; pinnules rhombic-obovate, irregularly jagged-lobed; some of the lobe-tips reflexed to form indusia over the lunate sori. HABITAT: Moist limestone cliffs, cataracts of limy water, walls of sinks, and spring basins; soil circumneutral to subalkaline. RANGE : A subtropical fern which extends northward locally on all continents. In America sporadically to British Columbia, Missouri, and Wythe Co., Virginia. Not found in the last state recently, but should be sought on river cliffs. Also apparently escaped from cultivation locally in Maryland, Pennsylvania, and New York. REMARKS : Distinguished from the next-following species by the habitat and the simple rachis.

— 77 —

Adiantum pedatum (v. originarium) NORTHERN MAIDENHAIR H I S T O R Y : Sent to Linnaeus f r o m Canada and Virginia, and named by him in 1753; the widespread variety termed originarium by the writer. F E A T U R E S : Rootstock slender, producing a row of glabrous deciduous f r o n d s usually 25 to 50 cm. t a l l ; stipe elongate, lustrous blackish brown, forking into two subequal rachises, and yielding a fan-shaped blade; each rachis bearing 5 to 9 spirally a r r a n g e d pinnae, which are pinnately divided; pinnules obliquely oblong, lobed on the u p p e r m a r g i n ; sori marginal, covered by indusia formed by reflexed lobe-tips. HABITAT: Wooded slopes, in humus-rich soils, or rarely on rocks; r a t h e r indifferent to soil reaction, but most l u x u r i a n t at low acidities. RANGE: Minnesota to Nova Scotia, Georgia, Louisiana and Oklahoma. Common over our region, except in pine-barrens. Also in Asia. REMARKS: One of our most beautiful ferns, readily cultivated. Various flowering plants, when not in bloom, may be mistaken for it, but they lack the dark-colored, terminally forked stalk. — 79 —

—

80

—

Cryptogramma stelleri FRAGILE ROCK-BRAKE H I S T O R Y : First named Pteris stelleri from Siberian material by Gmelin in 1768. Without knowing of this, Michaux named identical Canadian plants Pteris gracilis in 1803. Then in 1823 R. Brown proposed a genus Cryptogramma, and in 1882 Prantl placed the present species in this. F E A T U R E S : Rootstock slender, producing a short row of delicate deciduous fronds usually 8 to 15 cm. long; stipe brown at base; blade deltoid, 1- to 2-pinnate; sterile fronds with rounded ultimate divisions, the margins crenate; fertile ones taller, their divisions with undulate margins; sori in nearly continuous marginal strips. H A B I T A T : Moist shaded ledges of limestone or calcareous phases of other rocks; soil circumneutral to subalkaline, always cool. R A N G E : Circumboreal, in all northern lands; in America, Alaska to Labrador and Newfoundland, and south to Colorado, Iowa, and Bergen Co., New Jersey; also known in several deep gorges from Lycoming to Wayne Cos., Pennsylvania, and isolated in Randolph Co., W. Va. R E M A R K S : Inconspicuous, often overlooked. — 81

—

Cheilanthes lanosa —

82

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Cheilanthes lanosa HAIRY LIPFERN H I S T O R Y : Discovered by Michaux in the Tennessee and North Carolina mountains and named Nephrodium lanosum in 1803; assigned to the correct genus, Cheilanthes, by Eaton in 1858. F E A T U R E S : Rootstock short, producing a tuft of subevergreen fronds, usually 15 to 30 cm. long, hirsute, with rusty, jointed hairs; stipe and rachis deep brown; blade oblong-lanceolate, acuminate, twice-pinnate; pinnules cut into round oblong lobes, the margins reflexed to form indusia. H A B I T A T : Dry rocky slopes and ledges, in our region mostly non-calcareous, but from central Virginia southwestward often calcareous, the soil correspondingly subacid to circumneutral. R A N G E : This species extends further northeast than any other member of the genus, ranging from Texas to northwestern "Wisconsin and southern Connecticut. Occasional at moderate altitudes in our region, but absent from the higher mountains and the coastal lowlands. R E M A R K S : Fronds which shrivel up during long droughts may become normal again after a rain.

— 83 —

Cheilanthes tomentosa WOOLLY LIPFERN H I S T O R Y : Named by Link in 1833 on material supposed to have come from Mexico (or Texas). F E A T U R E S : Rootstock densely scaly, producing a cluster of firm, drought-resisting, but mostly deciduous fronds usually 20 to 50 cm. long; stipe and rachis stoutish, lustrous dark brown, densely chaffy and hairy; blade oblong, 3-pinnate, the tiny ultimate divisions obovate, densely covered on the back with matted whitish hairs, sparsely hairy on top; margins reflexed to form indusia. H A B I T A T : Dry ledges and crevices in cliffs of various rocks; soil circumneutral to subacid. R A N G E : Another southwestern plant which has spread into our region, though very much rarer than the preceding one: from Mexico and Arizona to Arkansas, Georgia, and Jefferson Co., West Virginia. In the last-named state it grows on schist, in Virginia also on shale, gneiss, and even on limestone at one locality. R E M A R K S : Even more of a " resurrection-plant'' than the preceding, curling up into a woolly mass in dry weather, but wholly restored by moisture. Budding fronds are folded instead of coiled. — 85 —

(Iheïlanthes

alabamensis

—

86

—

x ]/>

Cheilanthes alabamensis SMOOTH LIPFERN H I S T O R Y : Discovered by Buckley in Alabama and named Pteris alabamensis in 1843; correctly assigned to the genus Cheilanthes by Kunze four years later. F E A T U R E S : Rootstock scaly, producing a short row of subevergreen fronds usually 8 to 20 cm. long; scales limited to stipe-base; stipe and rachis blackish brown, lustrous; blade ovate-oblong, 2pinnate, glabrous; pinnules oblong to deltoid, somewhat auriculate and round-lobed; sorus a continuous marginal strip; indusium prominent, pale and firm. H A B I T A T : Sheltered crevices in limestone and other calcareous rocks; soil circumneutral to subalkaline. R A N G E : Mexico and Arizona to Missouri, Alabama, and to Giles Co., Virginia. Rare in our region, but probably occurs on occasional limestone cliffs in the Great Valley of southwestern Virginia. R E M A R K S : Readily distinguished from our other members of the genus by the absence of pubescence. — 87 —

Pellaea

atropurpúrea —

88

—

x y»

Pellaea atropurpúrea HAIRY CLIFFBRAKE H I S T O R Y : Discovered by Clayton at "Point Lookout" on the Rappahannock or James River, Virginia, and described by Gronovius about 1743; named by Linnaeus Pteris atropurpúrea ten years later. When Link founded the genus Pellaea in 1841, he included this species there. F E A T U R E S : Rootstock-tip densely covered with whitish hairs which turn brown in age; fronds in a loose cluster, thinnish but firm and evergreen, usually 10 to 50 cm. long; stipe and rachis blackish brown, rough-hairy; blade gray-green, oblong, openly 1- to 2-pinnate; pinnule? oblong, the fertile ones narrowly so; sori marginal. H A B I T A T : Cliffs, ledges, and talus slopes of limestone or of various more or less calcareous rocks; soil circumneutral or subalkaline. R A N G E : Arizona to northwestern Florida, northern Vermont, upper Michigan, and western South Dakota. Common over our region, except at the highest and lowest elevations. R E M A R K S : Frequently invades masonry, obtaining lime from the mortar.

— 89 —

Pellaea glabella

— 90 —

x

Pellaea glabella SMOOTH CLIFFBRAKE HISTORY : F o r many years this species was mistaken for depauperate material of the preceding one. Specimens sent from Kimmswick, Missouri, to Mettenius, were recognized by hun to be distinct, however, and his suggested name, Pellaea glabella, was published by K u h n in 1869. The plant was also named P. atropurpúrea v. bushii by Mackenzie in 1902. FEATURES : Rootstock-tip densely covered with deep red scales; f r o n d s clustered, firm and evergreen, usually 5 to 25 cm. long; stipe and rachis dark brown, glabrous; blade pale bluish green, oblong, compact-pinnate; lower pinnae sometimes divided into 3 (rarely 5) pinnules, or all simple; sori marginal; fertile divisions narrow. H A B I T A T : Bare crevices in cliffs and ledges, d r y or exposed to moist air, of limestone or rarely calcareous sandstone; soil subalkaline. R A N G E : British Columbia to northern Vermont, Tennessee, and northern Arkansas. I n our region rare, but scattered over limestone valleys. REMARKS : Can invade masonry even when solid and but little weathered.

— 91 —

Pteretis nodulosa OSTRICH F E R N H I S T O R Y : In 1 8 0 3 Michaux described Onoclea nodulosa, mistakenly ascribing it to Carolina. The identity of this remained a mystery for some time, but his type specimen proved to represent the American Ostrich Fern, and to have come from Montreal. Some botanists have assigned it to Mattcucia and Struthiopteris, but in 1916 Nieuwland placed it in Rafinesque's Pteretis. F E A T U R E S : Rootstock stout, producing a cluster of fronds 50 to 200 cm. tall; stipes brown, scaly; sterile fronds rather coarse but withering by late Summer, very narrow at base, widening gradually, and then rapidly contracted to the tip; pinnate, the linear pinnae cut into sub-lunate segments; fertile fronds autumnal, plumy. HABITAT : Swamps, moist humus-rich talus slopes, and wooded alluvial flats; soil circumneutral. R A N G E : Alaska to Newfoundland and south in our region at sea-level to northern Virginia, and in the mountains to Greenbrier Co., W. Va. R E M A R K S : The outline of both the sterile and the fertile fronds are highly characteristic. — 93 —

Onoclea sensibilis (v. genuina) BEAD-FEKN H I S T O R Y : Clayton found this fern in Virginia in the early 1700's, and Linnaeus named it in 1753. The American variety was named by the writer, 1937. FEATURES: Rootstock rather stout, sending up a row of fronds mostly 30 to 75 em. tall; sterile fronds coarse, withering early in Fall, ovate, openly cut nearly to the midrib; segments opposite, elliptic, entire, undulate, or coarsely lobed; fertile fronds autumnal, firm and long-persistent, twice pinnate, with the pinnules rolled into beadlike globules. HABITAT: Swamps, meadows, springy slopes, and wet muddy depressions; thriving in circumneutral soil, but tolerating moderate acidity. RANGE : Widespread in eastern North America, from Saskatchewan to Newfoundland, northern Florida, and Texas. Common over our region, except in acid pinelands. Has a relative in Asia. REMARKS: A very weedy plant, often termed ' ' sensitive fern,'' but as this is inappropriate, the common name bead-fern, in reference to the beadlike fertile pinnules, is here preferred.

— 95 —

Woodsia ilvensis x y2

— 96 —

Woodsia ilvensis RUSTY CLIFF-FERN HISTORY : In 1 7 5 3 Linnaeus termed Acrostichum ilvense a fern occurring in various parts of Europe (the species name referring to the Island of Elba). In 1810 Robert Brown recognized this to belong to an independent genus, which he named Woodia, correcting it to Woodsia 3 years later. Material from America proved so similar to the European that the same names are applied here. F E A T U R E S : Rootstock short; fronds usually 5 to 15 cm. tall, deciduous; stipe jointed, brownish ;. rachis brownish green; backs of fronds covered with copious rusty scales; indusium fringy. H A B I T A T : Cool dry cliffs, ledges, and rocky slopes, usually in subacid soil. RANGE : A circumboreal plant, occurring in all northern lands; rare in our region, but extending south near sea-level as far as Chester Co., Pennsylvania, and at higher altitudes along the mountains to northwesternmost North Carolina. R E M A R K S : Readily told by the jointed stipes and resulting uniform stubble, and the scaliness, yet frequently misidentified.

— 97 —

Woodsia obtusa — 98 —

Woodsia obtusa BLUNT-LOBE CLIFF-FERN H I S T O R Y : Discovered by Muhlenberg in Lancaster Co., Pennsylvania, and named by Sprengel in 1804 Poly podium obtusum, which was changed by Torrey in 1840 to Woodsia obtusa. F E A T U R E S : Rootstoek very short; fronds usually 20 to 40 cm. tall; tissue delicate, but sterile fronds evergreen; scales extending up stipe and rachis, mingling with minute glands which cover the herbage; stipe and rachis pale green to yellowish; lobes obtuse, but their margins in part reflexed so that they appear acutish; sori in irregular rows, toward margin; indusium lobed. H A B I T A T : Well-drained shady situations, crevices in cliffs, talus slopes, and sandy banks, often invading masonry; soil circumneutral. RANGE : Common and widespread nearly throughout the eastern states, except on the Coastal Plain; also in southeastern Canada, and in Alaska. R E M A R K S : -The fronds are often said to be nonevergreen and not scaly; actually the presence of scales forms an excellent means of distinguishing this Cliff-fern from Cystopteris.

— 99 —

Woodsia scopulina x y 2

— 100

—

Woodsia scopulina MOUNTAIN C L I F F - F E R N H I S T O R Y : Although occurring at several eastern localities, this fern was named on Rocky Mountain material by D. C. Eaton in 1865. It was first found in the Appalachians in 1897. F E A T U R E S : Rootstock short; fronds usually 1 5 to 30 cm. tall, deciduous; scales extending up stipe and mingling with long white minutely glandular hairs; stipe brown and rachis yellow, lustrous. H A B I T A T : Crevices in cliffs, ledges, and talus slopes; soil circumneutral to subacid, usually decidedly cool. R A N G E : Scattered over upland North America in several areas: Alaska to California and western Oklahoma; Minnesota to Arkansas; Quebec, Ontario and North Carolina. In our region rare, occurring on a few shale cliffs in southeastern West Virginia, and Bath to Smyth Cos., Ya. R E M A R K S : Most readily distinguished from the widespread Bluntlobe Cliff-fern by the presence of long hairs, and by the shining rachis, which suggests varnished wood. Our eastern plant may well represent a distinct variety. — 101 —

Cystopteris fragilis (v. mackayi) x y 2 indusium of v. genuina below to right -102

—

Cystopteris fragilis (2 varieties) U P L A N D BRITTLE F E R N HISTORY : The species here concerned was named by Linnaeus Polypodium fragile in 1753, and referred to a new genus, Cystopteris, by Bernhardi in 1806. The original was named v. genuina by Bernouilli in 1857, and the common east-American one v. mackayii by Lawson, 1889. F E A T U R E S : Rootstock short or elongated and covered with old stipe-bases, bearing fronds clustered at the scaly tip, usually 15 to 30 cm. tall; stipe brown and rachis green, glabrous; fronds lanceolate-oblong, 1- to 2-pinnate, with the divisions deeply cut. In v. genuina the pinnules are ovate and the narrow indusium is fringytipped; in v. mackayii, the pinnules elliptic and broad indusium entire at truncate tip. H A B I T A T : Moist recesses in cliffs, mossy slopes and sheltered ledges; soil circumneutral. RANGE : Widespread in uplands. V. genuina is circumboreal, and is sporadic in northern and high-altitude localities of our region; mackayii is our common representative. R E M A R K S : Intergrades are frequent between the above two varieties and the third (page 105).

— 103 —

Cystopteris fragilis (v. protrusa) LOWLAND B R I T T L E F E R N H I S T O R Y : The third variety of Cystopteris fragilis represented in our region was not separated until 1935, when C. A. Weatherby named it variety protrusa. F E A T U R E S : Rootstock elongated, nearly bare of stipe-bases, bearing fresh fronds in a row 2-4 cm. back from the bare tip, these usually 20 to 40 cm. tall; stipe brown and rachis green, glabrous; fronds lanceolate-oblong, 1- to 2-pinnate, with the pinnules deltoid to lanceolate, distinctly stalked, acutish, deeply lobed; indusium entire. H A B I T A T : Alluvial flats, wooded slopes, and humus-rich rock-crevices; soil circumneutral. R A N G E : Although most varieties of this species are northern, the present one centers in the lower Appalachians and adjacent lowland provinces, ranging from Georgia to Arkansas, southern Minnesota, and southeastern New York. In our region widespread, except in acid-soil areas. R E M A R K S : The long slender rootstock and the stalked pinnules would suggest specific independence were there not occasional intermediates.

— 105 —

CfetOriAN

Cystopteris

bulbifera

— 106

—

Cystopteris bulbifera BULBLET FERN H I S T O R Y : This North American fern was collected in Canada and sent to Linnaeus, who named it Polypodium bulbifcrum in 1753; it was included in the new genus Cystopteris by Bernhardi in 1806. F E A T U R E S : Rootstock short; fronds clustered, 20 to 40 or rarely 100 cm. long, often reclining; stipe and lower rachis pinkish; fronds long-deltoid, often elongate-acuminate, pinnate, with the pinnae pinnately divided into sharp-toothed lobes; axils of some pinnae or lobes bearing bulblets on under side of frond. H A B I T A T : Shaded ledges and talus slopes of limestone, calcareous shale, and occasionally other rocks, also hummocks in limy swamps; soil circumneutral or subalkaline. R A N G E : A cool-climate plant, extending from Manitoba to Newfoundland, Arizona and northern Georgia. In our region it is fairly common in calcareous districts, but absent from the Coastal Plain. R E M A R K S : The fronds are minutely glandular, a distinction from the other species.

— 107 —

Woodwardia virginica (Anchistea virginica) EASTERN CHAINFERN H I S T O R Y : Although collected in Virginia in colonial days, this fern did not come to Linnaeus' attention in time to be included in his Species Plantarum, but he named it Blechnum virginicum in a Supplement in 1771. J. E. Smith placed it in the genus Woodwardia in 1793, but Presl founded Anchistea on it in 1851. F E A T U R E S : Rootstock stout, producing a row of firm but deciduous fronds usually 75 to 125 cm. tall; stipe lustrous brown below; blade oblong, open-pinnate; pinnae elongate, deeply cut into obliquely deltoid lobes; veins forming a double chain of areoles along midvein, but free beyond that; fertile fronds aestival, the sori borne in chain-like rows of areoles. HABITAT : Bogs, swamps, and moist thickets, in moderately or often intensely acid soil. R A N G E : Florida to Texas and northward, chiefly along the coast, to Nova Scotia, also sporadically inland to Michigan. In our region common only in the Coastal Plain, rare inland. R E M A R K S : Resembles the Cinnamon Fern. — 109 —

Lorinseria areolata (Woodwardia areolata) NETVEIN C H A I N F E R N H I S T O R Y : This fern reached Linnaeus from Maryland and Virginia, and he named it Acrostichum areolatum in 1753. Presl founded the genus Lorinseria on it in 1851. F E A T U R E S : Rootstock slender, producing a row of dimorphic fronds; sterile ones deciduous, usually 30 to 75 cm. tall; stipe blackish at base; blade oblong, deeply cut into remote alternate segments; margins undulate, serrulate, or rarely lobed; veins areolate throughout; fertile fronds autumnal, persistent, taller but narrower than sterile ones; stipe and lower rachis blackish; blade pinnate with linear pinnae bearing chainlike rows of large sori. HABITAT : Swamps, wet woods, bog-margins, and springy slopes, in intensely acid soil. R A N G E : Chiefly near the coast, Florida to Texas, Arkansas, and Nova Scotia; also rare inland to Missouri and Michigan. In our region frequent in lowlands, but very rare elsewhere. R E M A R K S : Resembles the Bead-fern in aspect, but differs in having alternate segments. — Ill —

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112

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Phyllitis scolopendrium (2 varieties) HARTSTONGUE F E R N H I S T O R Y : The European Harts-tongue was known to the ancients as Scolopendrium, and Linnaeus made this the species name under Asplenium in 1753. Another early name, Phyllitis, was suggested for the genus by Ludwig, and combined with that species name by Newman in 1844. Pursh found an American variety of the species in Onondaga Co., New York, in 1805; this was named v. americana by Fernald, 1935. FEATURES : Fronds clustered, evergreen, usually 15.to 45 cm. tall; blade oblong, auricled, undulate. The American variety differs from the European in having the stipe-scales all narrow and slender vein-tips well back from margin. H A B I T A T : Sheltered rocks. The European variety is indifferent as to sort of rock and tolerant of high temperatures; the American thrives only on continually cool limestone talus. R A N G E : The American variety does not occur in our region, but the European lias been introduced in a ram-pit in Ilarford Co., Maryland. R E M A R K S : Very difficult to cultivate.

— 113 —

Asplenium,

trichomanes

— 114 —

Asplenium trichomanes MAIDENHAIR S P L E E N W O R T H I S T O R Y : Named by Linnaeus on European specimens in 1753. Subsequently found to be widespread in North America, the material here showing no essential differences from the original. F E A T U R E S : Fronds all alike, tufted, dark green, delicate but evergreen, usually 5 to 20 cm. tall; short stipe and long rachis lustrous dark brown; blade linear, pinnate, the pinnae small, numerous and subopposite, obliquely oval to roundoblong, often crenate along upper margin; sori few, short, medial. H A B I T A T : Moist sheltered crevices and cavernous places in rocks of all sorts, and on dry ledges of limestone; soil circumneutral to subacid. R A N G E : Alaska to Nova Scotia, central Georgia, and Arizona; common in our uplands, though not known on the Coastal Plain. R E M A R K S : One of our most beautiful ferns, forming charming rosettes against rock surfaces. The pinnae show considerable variation in shape, but are scarcely ever auricled; their margins vary from subentire to deeply cut-lobed.

— 115 —

—

116

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A s p l e n i u m resiliens BLACKSTEM SPLEENWORT

HISTORY : Discovered in Mexico and named by Martens and Galeotti A. parvulum in 1842. This being preoccupied, renamed by Kunze in 1844. FEATURES: Fronde dark green, clustered, uniform, firm in texture and evergreen, usually 10 to 20 cm. tall; stipe and rachis shining black; blade linear-elliptic, pinnate, the pinnae mostly opposite, oblong, auricled at base on upper (rarely also lower) side, obtuse; margins entire, crenate or rarely cut-lobed; sori few, medial. HABITAT: Crevices in cliffs of limestone and other calcareous rocks, usually in the shade, the soil correspondingly neutral to minimalkaline. RANGE: Western South America to Arizona, central peninsular Florida (and Jamaica), northward along limestone valleys to Missouri, and in our region from southwestern Virginia to Franklin Co., Pennsylvania, where known on a single rock. REMARKS: The common name here adopted refers to the feature which differentiates this from the two related species; it is sometimes inaptly termed Small Spleenwort.

— 117 —

pinna of v. incisum below to right.. —

118

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Asplenium platyneuron BROWNSTEM SPLEENWORT H I S T O R Y : Linnaeus in 1 7 5 3 proposed the name Acrostichum platyneuros for material from Virginia, comprising this plant and a Polypodium. Oakes combined the Linnaean species name with the correct genus, as recorded by Eaton in 1878. F E A T U R E S : Fronds dimorphic, the tuft including short prostrate evergreen sterile and erect subevergreen fertile ones usually 30 to 60 cm. tall; short stipe and long rachis lustrous dark brown; blades linear-elliptic, narrow at base, pinnate ; pinnae mostly alternate, the fertile deltoidoblong, auricled at base on upper side, crenate to jagged-serrate; sori near midrib. H A B I T A T : Hummocks of humus in swamps, sandy or clayey banks in open woods and thickets, talus slopes and rock ledges; indifferent to reaction. R A N G E : Central Florida to Texas, Colorado, and Maine; common over our region. R E M A R K S : A much used book-name, Ebony Spleenwort, is inapt since the stem is not black. Giant forms with much-cut margins are frequent southward, and have been named v. incisum. — 119 —

Asplenium bradleyi CLIFF

SPLEENWORT

H I S T O R Y : First collected near Coal Creek in eastern Tennessee by Bradley, in whose honor it was named by D. C. Eaton in 1873. F E A T U R E S : Fronds clustered, firm, evergreen, usually 5 to 20 cm. long; stipe and lower half of rachis lustrous brown; blade oblong, acuminate, pinnate; pinnae obliquely deltoid, deeply lobed, the segments serrate-margined; sori medial, dark brown. H A B I T A T : Crevices in bare cliffs of sandstone, gneiss, and other non-calcareous rocks, the record of its occurrence on limestone repeatedly copied by non-critical compilers of data being erroneous. Soil intensely acid. R A N G E : Sporadic, eastern Oklahoma to central Georgia, northeastern Kentucky, and latitude 42° in eastern New York. In our region rare, but known both in lower mountain and piedmont areas. R E M A R K S : The habitat of this fern is so often mis-stated that a reaffirmation of the situation is called for: Cliff Spleenwort does not grow on limestone. — 121

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Asplenium ruta-muraria (v. cryptolepis) pinnule of v. ohionis above to right

—

122

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Asplenium ruta-muraria (2 varieties) AMERICAN WALL-RUE SPLEENWORT H I S T O R Y : The European Wall-rue Spleenwort was named by Linnaeus Asplenium ruta-muraria in 1753. The American was long considered identical, but Pernald named it A. cryptolepis in 1928, and it was made a variety in Edition 1. The acutish-lobed variant was named v. ohionis by Fernald and transferred by the writer, 1942. F E A T U R E S : Fronds clustered, thickish, evergreen, usually 5 to 10 cm. long; stipe green except at the very base; blade ovate, 2- to 3-pinnate; pinnules rather wide-spaced, stalked, obovate, acutish to obtuse, coarsely to finely toothed at tip; sori few, linear, crowded. H A B I T A T : Sheltered cliffs and ledges of limestone and calcareous shale, sometimes invading masonry; soil circumneutral or subalkaline. R A N G E : New York and northwestern Vermont to central Alabama and northern Arkansas; also isolated on cliffs around the upper Great Lakes. In our region frequent in the limestone valleys. R E M A R K S : Differs from the European plant in obscure rootstock scales with thick-walled cells. The v. ohionts is occasional in Pennsylvania. — 123 —

— 124 —

Asplenium montanum MOUNTAIN SPLEEN WORT H I S T O R Y : Sent to Willdenow from the North Carolina mountains and named by him in 1810. F E A T U R E S : Fronds clustered, thinnish, evergreen, usually 5 to 10 or rarely 20 cm. long; stipe brown below, the upper part and rachis green; blade deltoid-oblong, 1 or 2-pinnate; pinnae and pinnules short-stalked, rhombic-elliptic, fewtoothed, acutish; sori few, short, somewhat separated. HABITAT : Moist sheltered crevices in sandstone, gneiss, and other non-calcarous rocks; soil mediacid or subacid. RANGE : Central Georgia and Alabama to Ohio and southeastern New York, and locally to Connecticut and western Massachusetts. In our region occasional in the mountains, except those of northern Pennsylvania, and rare in the Piedmont, reaching its lowest altitude in Lancaster Co., Pennsylvania. R E M A R K S : Sometimes confused with the nextpreceding species, but differs markedly in habitat and in cutting. Shows gradation toward the following one, but specifically distinct.

— 125 —

Oitt**--

Asplenium

—

trudelli

126

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Asplenium trudelli PINNATE

SPLEENWOBT

HISTORY : Although known for many years and even figured (as a form of the Lobed Spleenwort), this was not definitely distinguished until 1925, when named by the present writer on material from Lancaster Co., Pennsylvania. FEATURES: Fronds clustered, thickish, evergreen, usually 5 to 15 cm. long; stipe brown below, upper part and rachis green; blade deltoid-acuminate, somewhat caudate, pinnate at the base; mature fronds having 3 to 5 pairs of pinnae on definite slender stalks; upper part of blade, and pinnae, often deeply lobed; margins serrately coarse-toothed; sori short, about medial. HABITAT: Shaded crevices in sandstone and other firm, non-calcareous rocks, usually near colonies of the related species; soil mediacid. RANGE: Sporadic from northern Alabama to eastern Ohio and northwestern New Jersey. Once found in the last state near Blairstown, Warren Co., but apparently now extinct there. REMARKS: Presumably a hybrid between the Mountain and Lobed Spleenworts, into which it grades.

— 127 —

Asplenium pinnatifidum x % acuminate-lobed form below to right — 128 —

Asplenium pinnatifidum LOBED SPLEENWORT HISTORY : Discovered by Muhlenberg in Lancaster Co., Pennsylvania, and named A. rhizophyllum B, pinnatifidum in 1813; raised to species rank by Nuttall five years later.. F E A T U R E S : Fronds clustered, thickish, evergreen, usually 5 to 15, but exceptionally 30 cm. long; stipe brown below, green above; blade oblong-acuminate or lanceolate, caudate, variously lobed; lower lobes sometimes almost pinnae, but connected to the midrib by broad stalks; lobes obtuse to acuminate, their margins sometimes few-toothed; frond-tip rarely taking root; veins free or forming a few areoles; sori short, about medial. HABITAT : Shaded, moderately dry crevices in sandstone, gneiss, and other firm non-calcareous rocks; soil mediacid or rarely subacid. R A N G E : Bibb Co., Georgia, to Latimer Co., Oklahoma, Fountain Co., Indiana, and Berks Co., Pennsylvania. Reports from further northeast are apparently erroneous. REMARKS : Forms an intermediate between the genera Asplenium and Camptosorus. — 129 —

— 130 —

Asplenium hybrids Beside Asplenium, trudelli, already discussed, unmistakable hybrids between several pairs of species appear occasionally. A. Asplenium platyneuron x pinnatifidum (A. stotleri). Differs from presumed parents i n having round-lobed pinnae, but otherwise intermediate. On one cliff in Jefferson Co., West Virginia. B. Asplenium platyneuron x bradleyi. Shows close resemblance to both parents. In York and Lancaster counties, Pennsylvania. C. Asplenium montanum x bradleyi. Well intermediate between parents. Known in Lancaster Co., Pennsylvania, and Warren Co., New Jersey only. D. Asplenium pinnatifidum x bradleyi {A. gravesii). Combines parental characters. Stipe brown, rachis barely so; sori dark. Upshur Co., West Virginia, and Lancaster and York Cos., Pennsylvania. E. Asplenium platyneuron x trichomanes (A. virginicum). Fairfax Co., Virginia, and Monroe Co., Pennsylvania. — 131 —

— 132 —

x Asplenosorus ebenoides (Asplenium ebenoides) WALKING SPLEENWORT HISTORY : Discovered by R. R. Scott along the Schuylkill River in Montgomery Co., Pennsylvania, and named as an Asplenium, in 1865. F E A T U R E S : Fronds clustered, thinnish, evergreen, usually 8 to 25 cm. long; stipe and lower half of midrib shining brown; blade oblongacuminate, caudate, often pinnate at the base and lobed above; lobes irregular and highly variable in shape, usually acutish ; sori medial. HABITAT : Crevices in shaded rocks, in association with the parent species; soil reaction circumneutral or subacid. R A N G E : Vermont to Missouri and Hale Co., Alabama. Occasional in uplands. R E M A R K S : The genus name (E. T. W., 1937) refers to this being a hybrid between Asplenium, platyneuron and Camptosorus rhizophyllus. It has been produced by artificial crossing of the parents. In Alabama and sporadically further north it produces viable spores which yield normal plants, so is to be regarded as having attained species standing. — 133 —

c.rrLOMAM

Camptosorus rhizophyllus x

— 134 —

Camptosorus rhizophyllus WALKING F E R N H I S T O R Y : Linnaeus received this fern from Virginia and Canada and named it Asplenium rhizophylla in 1753, the ending a being a misprint for um. Link founded a genus Camptosorus to comprise this species and a related Asiatic one in 1833. F E A T U R E S : Fronds clustered, thinnish but evergreen, usually 10 to 30 cm. long; stipe brown at very base; blade simple, elongate-deltoid, usually cordate-auriculate at base, long caudate, the tip often rooting and forming a new plant; margins undulate; veins freely areolate. H A B I T A T : Sheltered ledges, cliffs, and talus slopes, rarely on hummocks of humus or even tree trunks, most luxuriant on limestone, but common also on other rocks; soil circumneutral. R A N G E : Minnesota to southern Quebec, central Georgia, and eastern Oklahoma. Frequent in our region, except on coastal sands. R E M A R K S : Camptosorus is little more than a subgenus of Asplenium, and this species indeed hybridizes with one member of that genus, yielding Asplenosorus.

— 135 —

otó Athyrium

pycnocarponx — 136 —

1/5

Athyrium pycnocarpon (Diplazium pycnocarpon) GLADE F E R N H I S T O R Y : Discovered by Michaux "on the banks of the Ohio River" and named by him Asplenium angustifolium in 1803. This name was already in use for a tropical fern, however, so is inacceptable for the present plant. Sprengel proposed for this Asplenium pycnocarpon in 1804, and Tidestrom made the combination here accepted in 1906. F E A T U R E S : Rootstock stoutish, producing a cluster of rather delicate deciduous fronds usually 50 to 100 cm. tall; stipe brown and scaly below; blade oblong-lanceolate, pinnate; pinnae acuminate, entire or crenulate; fertile fronds few, aestival, their blade and pinnae relatively narrow; sori numerous, elongate, in regular rows. H A B I T A T : Cool moist woods, shaded alluvial flats, and talus slopes; soil circumneutral. RANGE : Wisconsin to Maine, northern Georgia, Louisiana and Kansas. Occasional in uplands. R E M A R K S : Sometimes placed in the Asiatic Diplazium, again in the monotypic Homalosorus. Often inaptly termed Narrowleaf Spleenwort. — 137 —

Athyrium thelypterioides (Diplazium acrostichoides) SILVERY GLADEFERN HISTORY : First named Asplenium acrostichoides by Swartz in 1801, and two years later A. thelypterioides by Michaux, with reference to occurrences in Virginia and North Carolina. The combination of the earliest species name with Athyrium having been used otherwise, Michaux's is to be taken up, as done by Desvaux, 1827. FEATURES : Rootstock slender, producing a short row of firm but deciduous fronds usually 50 to 100 cm. tall; stipe yellowish, scaly below; blade oblong-lanceolate, markedly narrowed below, pinnate; pinnae deeply cut into oblong obtuse lobes, the lowest sub-auriculate; margins finely or sometimes coarsely toothed; indusium lustrous, the lobes appearing silver-striped. HABITAT : Woods, stream-banks, shaded slopes, and open thickets; soil most often subacid. R A N G E : Minnesota to Nova Scotia, and Missouri ; common in our region, except in the pinebarrens. A variety occurs in Asia. R E M A R K S : Related to Diplazium in its paired sori. Formerly called Silvery Spleenwort. — 139 —

Athyrium angustum (3 varieties) NORTHERN LADY-FERN H I S T O R Y : Linnaeus had named the Lady-fern of Europe Polypadium filix femina, and Michaux altered this to Nephrodium f. f. for a related Canadian plant. Other material from there was named by Willdenow Aspidium angustum in 1810, and his species was made an Athyrium by Presl in 1825. The division into varieties was worked out by Butters in 1917. FEATURES : Rootstock short, horizontal, covered by old stipe-bases, producing a short row of deciduous fronds usually 40 to 80 cm. tall; stipe yellowish to reddish; blade elliptic, widest at about the 5th pinna-pair above base, twice pinnate; pinnules deltoid-oblong, doubly serrate; fertile fronds narrowed; sori crowded, short, and hooked; indusia ciliate; spores yellow-brown. HABITAT: Moist acid woods and thickets. R A N G E : Manitoba to Newfoundland, Virginia, and Missouri; rare south of Pennsylvania. REMARKS -. V . ruiellum, with fertile and sterile fronds alike and sori spaced, is our common one. The small v. typicum, and the larger v. elatius, both have dimorphic fronds and confluent sori.

— 141 —

Athyrium asplenioides SOUTHERN

LADY-PERN

H I S T O R Y : Michaux was the first to observe this fern, but when naming it Nephrodium asplenioides in 1803 he mistakenly gave its range as in "New England and Canada." It was suggested to belong to Athyrium by Amos Baton as early as 1817, and formally placed there by Desvaux 10 years later. F E A T U R E S : Rootstock elongate, retaining few old stipe-bases, producing a short row of deciduous fronds usually 40 to 80 cm. tall; stipe yellowish or reddish; blade ovate, broadest at about the 2nd pinna-pair above base, 1- or 2-pinnate; pinnules or lobes obliquely deltoid-oblong, coarsely double serrate; sori medial, short, hooked; indusia bearing gland-tipped cilia; spores blackish. H A B I T A T : Swamps, moist woods, wet thickets, and brook banks; soil subacid or mediacid. R A N G E : Texas to Florida, Massachusetts and Missouri. Common over our region. R E M A R K S : Moderately variable. One striking form known as subtripinnatum, so deeply cut asto be nearly 3-pinnate, occurs in the mountains from North Carolina to southern Pennsylvania. — 143 —

Dryopteris

intermedia

— 144 —

Dryopteris intermedia (D. spinulosa v. intermedia) EVERGREEN WOODFERN H I S T O R Y : Another of the ferns discovered by Muhlenberg in Lancaster Co., Pennsylvania. His manuscript name for it, Aspidium intermedium, was published by Willdenow in 1810, and transferred to Dryopteris by Gray in 1848. F E A T U R E S : Rootstock producing a cluster of evergreen fronds usually 40 to 80 cm. tall; stipe bearing light or centrally dark brown scales; blade oblong-ovate, twice pinnate, with the pinnae almost perpendicular to the rachis; pinnules pinnately lobed, the lobes cut into sharp divergent teeth; inner pinnules of basal row usually shorter than the next outer ones, but longer in certain varieties; young herbage and indusia covered with tiny gland-tipped hairs. H A B I T A T : Woods, shaded rocky slopes, and hummocks in swamps; soil circumneutral or subacid. R A N G E : Wisconsin to Newfoundland, northern Alabama and Missouri; fairly common. R E M A R K S : Known commercially as Fancy Fern. V. fructuosa combines the features of this and the next, and is probably their hybrid. — 145 —

Dryopteris

spinulosa x

— 146 —

Dryopteris spinulosa TOOTHED WOODFERN H I S T O R Y : First named in Europe Polypodium spinulosum by 0 . F. Mueller in 1777, and placed in the genus Dryopteris by Watt in 1867. F E A T U R E S : Rootstock producing a short row of fronds usually 35 to 70 cm. tall, the sterile evergreen but the fertile withering by Fall; stipe bearing pale brown scales; blade ovate, twice pinnate below, once above, the pinnae ascending; pinnules and lobes marginally cut into oppressed teeth; inner pinnules of basal row longer than next outer ones; herbage and indusia glabrous. H A B I T A T : Swamps, wet woods, springy banks, and moist rocky slopes; soil most often subacid. R A N G E : A circumboreal plant, growing in Europe and Asia, and in this country from Idaho to Newfoundland, south to Missouri and Virginia. Common in our region, except in pinebarrens. R E M A R K S : Often stated to be evergreen, but actually only incompletely so. Some workers hold this and the next-preceding to be only varietally distinct, but the differences seem significant.

— 147 —

Dryopteris campyloptera (D. spinulosa v. americana) MOUNTAIN WOODFERN H I S T O R Y : The earliest name for this fern was Aspidium spinulosum americanum, suggested by Fischer and published by Kunze in 1848, together with the substitute A. campylopterum. Transferred to Dryopteris by Clarkson in 1930. F E A T U R E S : Rootstock bearing a cluster of wholly deciduous fronds, up to 100 cm. tall; stipe bearing dark brown-centered scales; blade deltoid-oblong, twice pinnate below, with the pinnules cut into lobes with divergent teeth; inner pinnules of basal row conspicuously elongated and as wide as two pinnules in the upper row; indusium glabrous or sparsely glandular. H A B I T A T : Cool rocky woods and ravine slopes, at high elevations southward; soil decidedly acid. R A N G E : Western Ontario to Labrador, Newfoundland, and sporadically southward to North Carolina. In our region, limited to a few widely separated mountains. R E M A R K S : Often confused with the European and west-American Dryopteris dilatata, but seems distinct. Grades somewhat toward D. spinulosa.

— 149 —

Dryopteris marginalis x y 5 pinna of forma elegans below to right — 150 —

Dryopteris marginalis MARGINAL WOODFERN H I S T O R Y : Sent to Linjiaeus from Canada and named by him Polypodium marginale in 1753; assigned to Dryopteris by Gray in 1848. FEATURES : Rootstock bearing a cluster of thick evergreen fronds usually 30 to 75 cm. tall; stipe bearing numerous light brown lustrous scales; blade ovate-oblong, acuminate, pinnate; pinnae deeply lobed, the lobes oblong, obtuse, subentire, crenate, or sometimes lobed; sori near lobe-margins or notch-bases. H A B I T A T : Shaded ledges, talus slopes, rocky woods, and rarely hummocks in swamps; indifferent to reaction, but often in subacid soil. R A N G E : Minnesota to Nova Scotia, northern Alabama and Oklahoma, also in British Columbia. One of the commonest ferns in our region, though rare on the coastal lowlands. REMARKS : Also' known as the Leather Woodfern. Occasionally develops an attractive variant, termed f. elegans; this exhibits broadening of the pinnae and greater cutting of the lobes, with the sori distinctly back from the margins.

— 151 —

Dryopteris goldiana GIANT WOODFERN H I S T O R Y : First collected by Goldie at Montreal, Canada, this fern was named Aspidium goldianum by Hooker in 1822, and transferred to Dryopteris by Gray in 1848. F E A T U R E S : Rootstock-tip covered with dark brown lustrous scales, bearing a cluster of few tardily deciduous fronds often 50 to 100 cm. tall; stipe bearing below dark, and above light brown scales; blade broadly ovate-oblong, with close-set pinnae, abruptly pointed; pinnae cut into oblong lobes of varying lengths, the basal pinnae narrow at the rachis, their margins decidedly undulate farther out; lobes appressed-serrate; sori near the midvein. H A B I T A T : Shaded rocky slopes, alluvial banks, and cool moist woods, in humus-rich and usually circumneutral soil. R A N G E : Mountains of South Carolina to Minnesota and New Brunswick; scattered over our region, but very rare in the coastal lowlands. R E M A R K S : Often termed Goldie's Fern; here named in reference to its being our largest Woodfern. — 153 —

Dryopteris clintoniana x % segment of v. genuina above v. australis — 154 —

Dryopteris clintoniana (2 varieties) (Dryopteris cristata v. clintoniana) BROAD S W A M P - F E R N H I S T O R Y : Named on northeastern material by D. C. Eaton Aspidium cristutum clintonianum in 1867. Transferred to -Dryopteris, though still as a variety, by Underwood in 1893, and raised to species rank by Dowell in 1906. The writer has separated varieties australis and genuina, 1937. FEATURES : Rootstock-tip covered with centrally dark brown scales, bearing a cluster of fronds usually 60 to 120 cm. tall; blade oblongacuminate, pinnate; pinnae deltoid-oblong, the basal ones broadly so, all long-acuminate, deeply lobed, the lobes oblong, obtusish, appressed-serrate; sterile fronds shorter, evergreen; fertile ones withering in Fall; sori near the midvein in v. genuina, medial in v. australis. HABITAT : Swamps, wet woods, and thickets on stream banks; soil usually subacid. R A N G E : V. australis occurs at wide intervals from the Gulf coast to northwestern and eastern Pennsylvania; v. genuina from the southern mountains to Wisconsin and Quebec. REMARKS : Sometimes approaches D. cristata.

— 155 —

— 156 —

Dryopteris cristata NARROW S W A M P - F E R N H I S T O R Y : This species was named from European specimens Polypodium cristatum by Linnaeus in 1753, and made a Dryopteris by Gray in 1848. The American plant is identical with the European. F E A T U R E S : Rootstock-tip covered with pale brown scales, producing a cluster of subevergreen fronds 35 to 70 or rarely 100 cm. tall; stipe sparsely scaly; blade linear-oblanceolate, pinnate; lower pinnae deltoid, upper deltoidoblong, mostly obtusish, deeply cut into deltoidoblong obtuse appressed-serrate lobes; sterile fronds shorter, evergreen; fertile ones withering early; sori medial; indusia glabrous. H A B I T A T : Marshes, bogs, swampy thickets, wet meadows, and springy slopes; soil sub-acid. R A N G E : Newfoundland to Idaho, Arkansas, and upland North Carolina; one of the few plants with such a range which grows also in Europe but not in Asia. Common nearly throughout. R E M A R K S : The narrow fronds, with the pinnae often turned to a horizontal position, give this species a characteristic aspect. — 157 —

Dryopteris boottii GLANDULAR S W A M P - F E R N H I S T O R Y : Discovered in Massachusetts and named by Tuckerman Aspidium booltii in 1843; transferred to Dryopteris by Underwood 50 years later. A F E A T U R E S : Roostock producing a cluster of mostly evergreen fronds up to 80 cm. tall; stipe moderately scaly; blade oblong or essentially so, twice pinnate below, once above; pinnules oblong, acutish, conspicuously serrate; sori near the midvein; indusiuin densely glandular. H A B I T A T : Moist thickets and swamps, most often in subacid soil. R A N G E : Minnesota to Nova Scotia and northern Virginia; occasional through the more northern and upland portions of our region. R E M A R K S : The most widespread and best known of the hybrids between Dryopteris species. That D. cristata is one ancestor is shown by the elongate-oblong outline, while the bipinnate cutting, serrate margins, and glandular indusia indicate D. intermedia to be the other. From both of these it differs, however, in the position of the sori near the midvein, and deserves species rank.

— 159 —

Three Dryopteris hybrids — 160 —

Dryopteris hybrids All of the species of Dryopteris seem to be capable of hybridization with one another. "When the parents are closely related, the hybrids exhibit intermediate characters, and are readily recognizable. Two pages are here devoted to hybrids between distantly related species; they are sometimes found remote from either parent. Dryopteris intermedia x marginalis. "Wooded slopes. A t first sight this suggests a phase of D. marginalis f. elegans, but the sori lie well in from the margin and the indusia are conspicuously glandular. D. spinulosa x marginalis (v. pittsfordensis) is non-glandular. Dryopteris intermedia x goldiana (separabilis Small). Moist woods. Rather unlike the parents in aspect, in that the frond is bipinnate, with the pinnules wide-spaced; indusia glandular. The corresponding spinulosa hybrid is non-glandular. Dryopteris marginalis x goldiana (leedsii W.) Rocky woods. Looks like a narrow D. goldiana or wide D. marginalis, but the sori are medial, and the rootstock-scales have dark centers. — 161

—

Dryopteris hybrids, continued Dryopteris marginalisxcristata (slossonae W.) In swamps, rather common. Resembles a narrow phase of D. marginalis, but told by the sori being medial. The first of these hybrids to be recognized as such, by Davenport in 1897. D. marginalis x clintoniana is similar but broader. Dryopteris goldiana x. clintoniana (D. celsa). Hummocks in swamps. Has the frond-outline of D. clintoniana, but the basal pinnae of D. goldiana. First recognized in the Dismal Swamp of Virginia, but now known to occur up to northeastern Pennsylvania. Dryopteris goldiana x cristata (atropalustris Small). Wet depressions in swamps. Widest near the base; lower pinnae resembling those of D. goldiana, but highly variable. Relationship with D. cristata is shown by the pale rootstockscales and the medial sori. Known only in Kent Co., Delaware, the Dismal Swamp of Virginia, and southeastern South Carolina. Others recorded comprise D. spinulosa x cristata and x clintoniana. — 163 —

Polystichum braunii (v. purshii) EASTERN HOLLY F E R N H I S T O R Y : The species here concerned was discovered in Europe, and named Aspidium braunii by Spenner in 1825, then transferred to the genus Polystichum by Fée about 1850. The American material differs somewhat from the European and was named v. purshii by Fernald in 1928. F E A T U R E S : Rootstock stout, chaffy, producing a cluster of subevergreen fronds, usually 35 to 70 cm. tall ; stipe short, brown-scaly ; blade elliptic, twice-pinnate, the pinnules cut into bristly appressed teeth ; fertile and sterile pinnules alike; sori large, toward the midvein. HABITAT : Cool shaded talus slopes, rocky woods, humus-rich pockets in rock ledges, and southward in sheltered ravines near watercourses ; soil usually circumneutral. R A N G E : Newfoundland to northern Wisconsin (also isolated in Alaska), extending southward into the uplands of New England and New York ; entering our region only in northeastern Pennsylvania, in Luzerne, Sullivan, and Wayne cos. R E M A R K S : Our most beautiful northern fern.

— 165 —

Polystichum acrostichoides CHRISTMAS F E R N H I S T O R Y : Michaux observed this fern during his explorations in '' Pennsylvania, Carolina, and Tennessee," and named it Nephrodium acrostichoides in 1803. It was transferred to Polystichum by Schott in 1834. F E A T U R E S : Rootstock stout, chaffy, producing a cluster of evergreen fronds 30 to 75 cm. tall; stipe and rachis very scaly; blade lanceolate, pinnate, the pinnae lanceolate and often subfalcate, with an upward-pointing basal auricle, the margins bristle-toothed, serrate, or sharp-lobed; sterile fronds relatively short; fertile bearing crowded sori on several reduced terminal pinnae. H A B I T A T : Wooded slopes, liumus-rich crevices in rocks, hummocks in swamps and open thickets; practically indifferent to soil reaction. RANGE : Southeastern Wisconsin to Nova Scotia, central Florida, and eastern Texas; one of the commonest ferns on shaded slopes in our region. R E M A R K S : Uniform in some colonies but in others strikingly variable in size, frond-outline, cutting, and distribution of sori.

— 167 —

Thelypteris palustris —

168

(v. pubescens) —

Thelypteris palustris (v. pubescens) (Dryopteris thelypteris) NORTHEASTERN M A R S H - F E R N H I S T O R Y : Linnaeus in 1 7 5 3 gave the name Acrostichum thelypteris to a European fern, and 9 years later Schmidel made Thelypteris a genus. Schott proposed T. palustris in 1834. Then in 1848 Gray made the combination Dryopteris thelypteris. In 1929 Fernald showed the northeast-American plant to be distinct from the European, as v. pubescens of Lawson, 1864. F E A T U R E S : Rootstock rather slender, producing a row of grayish green deciduous fronds usually 40 to 80 cm. tall; stipe bronzy below; blade lanceolate, pinnate, often fine-pubescent; pinnae deeply cut into oblong obtusish lobes; veins of sterile lobes forked; fertile fronds aestival, their lobes reflex-margined, their veins in part simple; sori medial, crowded. H A B I T A T : Marshes, swamps, wet thickets, and bog margins; thrives best in soil of low acidity. R A N G E : Manitoba to Newfoundland, Georgia and Oklahoma; common nearly throughout our region. Also in northeastern Asia. R E M A R K S : Veins often wrongly described.

— 169 —

— 170 —

Thelypteris simulata (Dryopteris simulata) BOG-FERN H I S T O R Y : First collected by Dodge at Seabrook, New Hampshire, in 1880, and named by Davenport Aspidium simulatum.in 1894. Transferred to Thelypteris by Nieuwland in 1910. F E A T U R E S : Rootstock slender; fronds in a row, usually 30 to 60 cm. tall, yellowish green; stipe brown below; blade elliptic-lanceolate, pinnate; pinnae narrowly elliptic, obtusish, lobed, the basal pair relatively short, somewhat narrowed where they join the rachis; veins all simple; fertile fronds aestival, their lobes flat; sori submarginal, well-spaced. H A B I T A T : Shaded margins of sphagnum bogs and moist woods, in intensely acid, sterile, soil. R A N G E : As yet imperfectly known; mountains of northeastern Alabama to Prince Edward Island, Canada. In our region sporadic, in New Kent Co., Virginia, and Randolph Co., West Virginia, New Jersey, and eastern Pennsylvania. R E M A R K S : Readily confused with other ferns, as the species name implies. Sometimes termed Massachusetts Fern, but this has no significance. — 171 —

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ZULOHAH

Thelypteris noveboracensis x % — 172 —

Thelypteris noveboracensis (Dryopteris noveboracensis) TAPERING F E R N H I S T O R Y : Sent to Linnaeus from "Canada" —probably the northeastern United States—and named by him Polypodium noveboracense in 1753. Made a Thelypteris by Nieuwland in 1910. F E A T U R E S : Rootstoek slender, producing a short row of yellowish green thin deciduous fronds usually 30 to 60 cm. tall; stipe short, pale green, barely scaly; blade elliptic and longacuminate at both ends, pinnate, hairy on the heavier veins beneath; lower pinnae deltoid, upper oblong-acuminate, cut into oblong obtuse lobes; veins mostly simple; fertile fronds aestival, the sori submarginal; indusia finely glandular. H A B I T A T : Moist woods, thickets, and swamps, usually in rather acid soil. R A N G E : Minnesota to southern Newfoundland, Georgia and Arkansas. Common over our region. R E M A R K S : A translation of the species name, New York Fern, is often used. This and the two preceding species are frequently placed in Dryopteris, but differ so from its typical representatives that generic distinctness seems reasonable. — 173 —

Phegoptcris

conncctilis — 174 —

x

Phegopteris connectilis (Phegopteris polypodioides) (Dryopteris or Thelypteris phegopteris) NORTHERN B E E C H F E R N HISTORY : Named by Linnaeus Polypodium phegopteris in 1753, and by Micliaux P. connectile in 1803. Foe called it Phegopteris polypodioieles in 1853, while "Watt made the combination here adopted in 1867. F E A T U R E S : Rootstock slender, sending up a row of thin, deciduous fronds usually 15 to 30 cm. tall, hairy throughout and bearing brown scales on under side along rachi.s and main veins; blade narrowly deltoid, witli one basal pair of pinnae, above which the rachis is bare, and pinnalike segments above; segments cut into obtuse lobes; sori small, toward the margin, exindusiate. HABITAT : Woods, wet thickets, and moist rockcrevices, southward only in cool situations; soil varying in reaction, most often subacid. R A N G E : A circumboreal plant, occurring in afll northern lands; Alaska to Newfoundland and south at low altitudes to southeastern Pennsylvania, and in the mountains to North Carolina. R E M A R K S : Rather rare in our region.

— 175 —

Phegopteris

hexagonoptera — 176 —

x ys

Phegopteris hexagonoptera (Dryopteris or Thelypteris hexagonoptera) SOUTHERN B E E C H F E R N H I S T O R Y : Though early collected in Virginia and figured by Plukenet in 1691, this was confused with the preceding by Linnaeus. Michaux named it Polypodium hexagonopterum in 1803, Fée placed it in his genus Phegopteris in 1853. F E A T U R E S : Rootstock slender, sending up a row of deciduous fronds usually 20 to 40 cm. tall, minutely pubescent, with some glandular hairs; blade broadly deltoid, acuminate, divided into pinna-like segments, which are cut into oblong obtusish undulate-margined lobes; sori small, toward margin, exindusiate. HABITAT : Wooded slopes and open rocky thickets ; soil usually subacid. R A N G E : Eastern Texas to northernmost Florida, Quebec, and Minnesota. Rather common nearly throughout our region. R E M A R K S : The broadly deltoid outline is characteristic. This with the next-preceding species is here retained in the minor genus Phegopteris because of the combination of slender rootstock, deltoid blade, lack of indusium, etc.

— 177 —

Currania dryopteris (Phegopteris dryopteris) WOODLAND OAKFEBN H I S T O R Y : Known to Linnaeus in Europe, and named by him Polypodium dryopteris in 1753; included in Gymnocarpium by Newman in 1751 and Phegopteris by Fee two years later. Has also been placed in six other genera, but assigned to still another, Currania, by the writer, 1942. FEATURES : Rootstock slender; fronds in a row, thin, deciduous, usually 20 to 40 cm. tall, glabrous, or the rachis fine-pubescent; stipe pale yellow, becoming greenish brown; blade deltoid, ternately pinnate; stalks of basal pinnae jointed at the rachis; pinnules deeply obtuse-lobed; sori small, toward margin, exindusiate. HABITAT : Cool rocky woods, shaded talus slopes, and swamp margins; soil often sub-acid. R A N G E : Circumboreal, in all northern countries; Alaska to Labrador and Newfoundland, and southward; rather rare in our region, but extending at low altitudes to southeastern Pennsylvania and in mountains to Page Co., Virginia and Randolph Co., West Virginia. REMARKS : The jointed pinna-stalks are a unique feature of this minor genus. — 179 —

W-WACNL.-R.

Currania robertiana x — 180 —

Currania robertiana (Phegopteris or Dryopteris robertiana) LIMESTONE OAKFERN H I S T O R Y : First named Polypodium robertianum in Europe by Hoffmann in 1795; placed in various other genera by subsequent workers, and assigned to Currania by the writer in 1942. F E A T U R E S : Rootstock slender; fronds spaced, deciduous, to 50 cm. tall, glandular-pubescent; stipe yellowish; blade deltoid, pinnate, subternate; basal pinna-stalks jointed to rachis; pinnules obtuse-lobed; sori small, exindusiate. H A B I T A T : Moist shaded limestone cliffs and talus and cool calcareous swamps. R A N G E : Circumboreal but rarer than the nextpreceding species. Alaska to Newfoundland and the Great Lakes—St. Lawrence valley region. Recently discovered in Blair Co., Pennsylvania, 450 miles from the nearest colony in Douglas Co., Michigan, and 750 miles from the southwesternmost in New Brunswick, Canada. REMARKS : Best distinguished from the Woodland Oakfern by the longer frond and glandular pubescence. A remarkable disjunct occurrence. — 181 —

Polypodium virginianum x V2 aeuminate-lobed form to right — 182

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Polypodium virginianum (P. vulgare v. virginianum) ROCKCAP F E R N H I S T O R Y : Linnaeus in 1 7 5 3 recognized a European Polypodium vulgare and an American P. virginianum-, for many years American writers have used the former name for both, but their distinctness is now coming to be accepted. F E A T U R E S : Rootstock densely brown-scaly, producing a row of evergreen fronds, usually oblong and 10 to 30 cm. long, cut nearly to the midrib into entire obtusish, linear segments; in variants the outline may be deltoid, and the segments may be lobed or auricled, and terminally acute or acuminate; sori large, exindusiate. H A B I T A T : Dry rocks, often capping boulders, also cliffs, ledges, and talus slopes, sometimes wooded banks or humus on tree-trunks; soil varying in reaction, most often subacid. R A N G E : Eastern Alberta to Newfoundland, northern Georgia and Arkansas; common. R E M A R K S : Develops a considerable number of forms, at least 10 of which have received names; the most striking is f. cambricoides, which has the segments cut-lobed in a lacy pattern. — 183 —

Polypodium

polypodioides

— 184 —

v.

michauxianum

Polypodium polypodioides v. michauxianum RESURRECTION F E R N H I S T O R Y : This fern reached Linnaeus from Virginia and Jamaica, and was named Acrostichum polypodioides in 1753. It was transferred to the genus Polypodium by Watt in 1866. The variety was named by Weatherby in 1938. Some workers prefer to place it in Marginaria, to which it was referred by Tidestrom in 1905. F E A T U R E S : Rootstock fine-scaly, producing a row of evergreen fronds usually 5 to 15 cm. long, densely scaly on the under side; outline deltoidoblong, pinnately divided into linear obtuse segments ; sori toward the margin, exindusiate. HABITAT : High or sometimes low on treetrunks, mossy cliffs, and rarely in hummocks of humus; seemingly indifferent to soil reaction. R A N G E : Tropical America northward to Iowa, Montgomery Co., Maryland, and Sussex Co., Delaware; rare in our region. R E M A R K S : Sometimes ascribed by amateurs to New Jersey or Pennsylvania, but not thus far known in these states. The common name refers to the fact that in dry weather the fronds curl up and appear dead, but revive following a rain. — 185 —

Lycopodium selago x 1 / 2 v. typicum and v. patens

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186

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Lycopodium selago (2 varieties) FIR

CLUBMOSS

HISTORY : This plant was well-known to Linnaeus and was named in his Species Plantarum in 1753. In the first edition of this Guide the original was designated v. appressum, but this was changed to v. typicum by the writer in 1942. Its relative, named Plananthus patens by Beauvais in 1805, was reduced to L. selago v. patens by Desvaux in 1827. F E A T U R E S : Plant evergreen, 5 to 1 5 cm. high; horizontal stem short, with clustered upcurving branches; leaves appressed in typicum, spreading in patens; sporophyls in zones on stems. H A B I T A T : V. typicum, on cool shaded ledges, rocky barrens, and bog-margins, in mediacid soil; patens, cliffs and rock-slides, also acid. R A N G E : Widespread in northern lands, extending south along mountain ranges. In our region r a r e : typicum known only in Monroe Co., Pennsylvania and Page-Madison cos., Virginia; patens, Pike Co., Penna. and eastern \V. Va. Both occur at high altitudes in the Carolinas. R E M A R K S : Variants of v. patens (or even typicum) are sometimes termed Lycopodium porqphilum; this is not now considered valid. — 187 —

Lycopodium lucidulum

— 188

(v. verum) x '/Ó

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Lycopodium lucidulum (•. verum) S H I N I N G CLUBMOSS H I S T O R Y : In the course of his travels, Michaux observed this plant from ' ' Canada to the Carolina mountains," and named it in 1803. The writer has designated the original variety verum, 1937. F E A T U R E S : Horizontal stem well developed; upright stems evergreen, usually 10 to 30 cm. tall and 2 or 3 times forked; leaves wide-spreading, deflexed in age; sterile ones oblanceolate, terminally sharply serrate (or in v. occidentale entire), the fertile ones shorter, deltoid-oblong, and subentire; cones lacking. H A B I T A T : Wet woods, shaded stream-banks, and moist rocky slopes, in acid humus-rich soil. R A N G E : Widespread from central Canada south to Missouri and northern Alabama, and common over much of our region, except in the warm lowlands. V. occidentale scarcely enters our region. R E M A R K S : Grades through v. occidentale into L. selago patens, but the spores of all lucidulum varieties are less than 0.03 mm. in diameter while in selago varieties they are over this. The supposed species L. porophilum included both. — 189 —

Lycopodium inundatum, x y