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Les diagrammes suivants illustrent la mAthode. errata to ) pelure, on & D 32X 1 2 3 1 2 3 4 5 6 V w w OUR NATIVE FERNS AND THEIR ALLIES WITH Synoptical Descriptions of the American Pteridopbyta North of Mexico BY LUCIEN MARCUS UNDERWOOD SIXTH EDITION, REVISED NEW YORK HENRY HOLT AND COMPANY 1900 QV. SV- 5 , U i^ fc V^OO ' ( V » COPTRIGHT, 1888, 1900, IV HENBY HOLT & CXX BOBIRT DBUMMOND, PRnmR, SKW YORK. I \ PREFACE. When the writer issued this little book in 1880 as the honest effort of a novice to provide for the study of our ferns a roT^venient handbook by means of which they might be identified, he had no idea that the first edition would be exhausted within a year, and much less that a sixth edition would ever be called for. Though frequently urged to extend its scope, he has felt that if, with all the traces of its early imperfections of plan, there is still a demand for such a handbook, it is best to leave it in its original form, with only such changes as our changed conceptions of structures, relationships, and definition of species demand. Not only is this preservation of the original plan in harmony with the feeling of sentiment, but it seems the more desirable since the writer is preparing a monograph of all the North American Ferns (including those of the West Indies and the continent as far as the Isthmus), and in this more elaborate work he hopes from a study of a wider range of forms to include many more general matters that our own limited fern flora, though quite diverse, do not furnish a sufficient basis for inclusion here, and others still that would be out of place in an elementary manual. Changes in this edition are mostly verbal and such as arise from the modifications of nomenclature or the changed ideas of homologies and relations of structures. The chapter on nomenclature has been wholly rewritten and extended, particularly because the present edition more than any othei contributes to a modification of generic names. 148225 •v vi PREFACE. In the systematic part the sequence has also been modified, bringing the simpler eusporangiate forms first and introduc- ing the desirable distinction between orders and families which botanists have too long confused. The number of species is considerably increased, owing in part to new discoveries and in part to the seeming necessity of reestablishing the earlier and in many cases clearer views of the earlier writers on ferns, many of whose species have been reduced to synonymy by the English (Kew) school of fern writers whose dictum has hitherto been followed by American fern students. The number of genera has also been increased in accordance with the views of earlier and more scientific fern students. Columbia University, June, 1900. ■• ■*, TABLE OF CONTENTS. PAOB Introduction ix Chapter I. Haunts and Habits of Fbrns i II. The Organs of the Growing Fern 8 III. Fructification IN Ferns lo IV. Germination of Fern Spores 19 V. Fern Structure 24 VI. The Fern Allies ag VII. Classification AND Nomenclature 41 VIII. The Fern's Place in Nature 54 IX. Distribution in Time and Space 60 OUR NATIVE PTERIDOPHYTA. ORDER FILICALES. FAMILY!. OPHIOOLOSSACEiC 66 a. HYMENOPHYLLACEiE 74 3. SCHIZiGACEiE ^5 4. OSMUNDACEiC jj 5. CERATOPTERIDACEiC yg 6. POLYPODIACEiG yg 7. MARSILIACEiE fjj 8. SALVINIACEiC 125 ORDER EQUISETALES. Family I. EquisETACEiE 126 ORDER LYCOPODIALES. Family I. LYcoPODiACEiC , 2. Selaginellace^ J2- 3. ISCETACEA J, 2 Abbreviations j -o Glossary and Index jei vii INTRODUCTION. \ I In the entire vegetable world there are probably no forms of growth that attract more general notice than the Ferns. Deli- cate in foliage, they are sought for cultivation in conservatories and Wardian cases, and when dried and pressed add to the culture of many a domestic circle by serving as household deco- rations. They furnish to botanists a broad and inviting field for investigation, and he who examines their more minute struc- ture with the microscope will find deeper and still more myste- rious relations than those revealed to the unaided eye. Ferns thus appeal to the scientific element of man's nature as well as to the aesthetic, and while they highly gratify the taste, they furnish food for the intellect in a like degree. The Fern allies have also played their appointed part in the domestic and decorative economy of this and other generations. The scouring-rushes served our ancestors for keeping white their floors and wooden-ware in the days when carpets were a luxury. The trailing stems of various species of Lycopodium have long been valued for holiday decorations ; while their burning spores have flashed in triumphal processions, and have added their glow to the fervor of political campaigns. In olden time the obscure fructification of the common brake led to many superstitious ideas among the common people, and the older poets have woven these popular notions into our litera- ture. Butler tells in Hudibras of bugbears so often created by mankind : " That spring like fern, that infant weed, Equivocally without seed, And have no possible foundation Bnt merely in th' imagination.** IN TROD UCTION, Shakespeare only reflects a prevalent belief of his time when he says : •* We luiT* the receipt of fern seed ; we walk inviiible." Others allude to the falling of the seed on the anniver- sary night of the birth of John the Baptist. The old simplers with their lively imagination were impressed by the fancied resemblances of some parts of fern growth to various organs of the human body, and introduced them into their system of specifics. Traces of their influence still remain in the names of some of our common ferns, as spleenwort and maidenhair. To form a correct understanding of ferns we must study the ferns themselves as well as the text-book, as it is only by direct contact with nature that we gain definite and satisfactory informa- tion. The text-book is useful only io giving directions how to investigate. To understand thoroughly an animal we must study its habits in its native haunts. To know its structure and posi- tion in the animal kingdom we must carefully dissect a large number of specimens, and study the development of the individ- ual from its beginning. In like manner, to understand fully a fern we must search where nature has planted it, watch it as it un- coils from the bud, matures, produces its fruit, and finally returns to the earth ; examine it with needles and lenses, and discover its minute structure and its life-history. These pages, which aim to give an outline of the forms of fern growth, the methods of fruiting, the germination or growth from the spore, and finally the more minute structure of the entire plant, can only be thoroughly understood by taking the ferns in hand and studying them in connection with the text. Let no one imagine that the study of ferns will be an easy one. Patient application and careful observation are essential to success, yet he who becomes once interested in the work will find a subject that deepens in interest with every step, and even becomes enchanting as he seeks to determine the mysteri- ous processes of fern development and the marvels of fern structure. OUR NATIVE FERNS AND THEIR ALLIES. CHAPTER I. HAUNTS AND HABITS OF FERNS. i Our outward life requires them not, — Then wherefore had they birth ? To minister delight to man, To beautify the earth. —Mary Howitt. 1 . 0«n«ral Characters.— Our native ferns comprise plants varying in height from less than an inch to six or seven feet, or even more. Some are stout and fleshy, others are delicate and even filmy, but most are herbaceous, resembling ordinary flowering plants in the texture of their foliage. While most would be recognized as ferns by even a novice, a few differ so widely from the ordinary typical forms that to an unskilled ob- server they would scarcely be considered as bearing any resem- blance to ferns whatever. The fronds of one of our Florida species resemble narrow blades of grass, and the fertile spilces of another from New Jersey might be mistaken for a diminutive species of sedge. A third from Alabama would, perhaps, be called a moss by the inexperienced, while the " Hartford fern," found from New England to Kentucky, has a climbing stem and broad palmate leaves. When we add to these peculiar forms of our own country those of foreign lands, and include the immense tree-ferns of tropical regions, we find our early conception of a fern inade- quate to cover this diversity of forms. Without attempting an accurate definition of a fern, let it be regarded for present pur- I V, 2 OUX NATIVE I^EHNS AND THEIR ALLIES. poses as a flowerless plant, producing spores instead of seeds, possessing more or less woody tissue, and having its leaves coiled in the bud from apex to base. After the necessary study of the structure of some of our common ferns, we will be able to comprehend the more technical definition found later in the work. 2. Mode of Growth. — Ferns vary greatly in their method of growth, yet each species has a plan which, within certain limits, is fixed and definite. Some, like the common brake, have their fronds rising from more or less distant portions of the creeping rootstock. Others, like Aspienium trichomanes, are tufted, many fronds rising irregularly in a cluster ; while still others, like the ostrich-fern {Matteuccid), and many shield- ferns {Dryopteris), grow in crowns or circles, the later fronds continually rising within the older ones. In the grape-ferns {Botrychiutn) the rootstocks usually produce a single frond each season, the bud for the succeeding year growing within the base of the common stalk. 3. In many there is a tendency to dimorphism, the fertile or fruit-bearing fronds differing to a greater or less extent from the sterile ones. In a few species, like the sensitive-fern {Ono- cled) and some others, this is carried so far that the sterile and fertile fronds bear no resemblance to each other, and in one instance have been mistaken for different species, and so de- scribed. Osmunda cinnamomea, Woodwardia areolata, our two species of Cryptogramma, and Strut hiopter is* ofiex further ex- amples of this principle of growth. 4. Variation. — The same species will often present wide differences in the size of the fronds. This depends to some ex- tent on the character of the soil and the ordinary climatic con- ditions. For example, the lady-fern {Aspienium fiUx-fcemind), which in ordinary locations grows from two to four feet high, in mountainous regions is sometimes reduced to from three to six inches, when it forms the var. exile. In like manner the marginal shield-fern {Dryopteris marginalis), usually two or three feet high, is reduced to five inches when growing on rocky cliffs, and yet regularly produces fruit.* * Cf. Bulletin Torrey Botanical Club, vi. 966 (Oct. 1878). HAUNTS AND HABITS OF FERNS, 5. In some cases there is a tendency to variation in size that cannot be referred to soil or climatic influences. The com- mon grape-fern {Botrychium Virginianwn) will be found in some localities to vary from six inches to two feet in height, all well fruited and matured, and with the extreme sizes growing within a pace of each other in the same soil and with the same environment. The other species of the same genus present similar variations, and judging from size and external appear- ance alone, a regular gradation of forms might be arranged from the most diminutive undivided forms of B. simplex to the larg- est of B, Virginianum. 6. Another tendency to variation is noticed in the forking of fronds either at the summit or at the ends of the branches. The hart's-tongue {Phyllitis) is frequently forked at the sum- mit, the walking-leaf {Camptosorus) less commonly, while the same tendency is noticed in various compound forms, as Aspie- ntum angustifolium, Cheilanthes lanosa^ Gymnopteris hispida^ Dennstcedtia, Pellcea atropurpurea, and others. Some of the species of Botrychium show the same tendency, especially in their fertile segments. It is probable that all our species will be found to fork under certain conditions. More definite in- formation is desirable with regard to many species that show this tendency, as it doubtless involves the question of ancestry of existing ferns. 7. In those species whose sterile and fertile fronds are un- like, forms often appear that are intermediate between the ster- ile and fertile fronds, and sometimes even form a graded series from one to the other. This is especially true of the sensitive- fern {Onoclea) and the cinnamon-fern {Osmunda cinnamomed), and has frequently been the source of so-called "varieties." Whether this variation arises from some peculiarity of environ- ment, or from some inherent tendency to reversion toward an older form, will require more extenwcd observation to deter- mine. One of the varieties of Botrychium obliquum seems to have been founded on a condition which is intermediate in structure between the sterile and fertile segments. 8. In a few forms there is an apparent mimicry, one species imitating another in foliage or method of fruiting. In the cin- namon-fern just alluded to, which has a cinnamon-colored OUR NATIVE FERNS AND THEIR ALLIES. sterile frond totally unlike the fertile, sterile fronds will some- times be found which are fertile at the apex — ^the normal method of fruiting in the royal flowering-fern (Osmunda regalis) ; and in turn the royal flowering-fern is sometimes fertile in the middle, in imitation of Osmunda Claytoniana. 9. Time of Fruiting. — The time of maturing fruit is dif- ferent among different species, and also varies with geographi- cal location and proximity to tropical climates. In the Northern States some species produce their fruit as early as May (Osmunda cinnatnomed), and others as late as September {Lygodiutn), but the gp-eater number are best studied in July and August. In the Northeastern States, where the two species of Filix abound on limestone rocks, F.fragilis matures its spores and withers in June or July, while F. bulbifera reaches its maturity only in August or September. In semi-tropical climates, like Southern California and the Gulf States, the time of fruiting is often earlier, sometimes occurring in February or March. Some fronds are killed by the early frosts, while others, like the Christ- mas-fern, are evergreen, and may be gathered in midwinter. 1 0. Local Distribution. — Ferns are largely dependent for successful growth on the amount of warmth, moisture, and shade to which they are subjected, and we would naturally ex- pect to And them reaching a maximum in size and abundance in warm swamps or shady marshes. While this is in general true, we nevertheless find many species thriving only in rocky places, thrusting their roots into the crevices of the rocks with little earth for their nourishment, and many times exposed to the scorching rays of the sun. Of necessity, such species are of comparatively small size, and likely to be protected in some way against the heat of the sun, and provided with means to retain their moisture in times of drought. Others still are found in wet, rocky ravines, often where moistened by the spray of cascades or waterfalls, and consequently have no such pro- vision against the heat of an extended summer. Certain others thrive in open fields that are comparatively dry aiid unshaded. One species of Southern Florida is aquatic, having the sterile fronds floating in shallow water. A few species are epiphytic, or grow on other plants, some being found on tree-trunks to the height of 1 50 or 200 feet ! HAUNTS AND HABITS OF FERNS, So, while moisture, warmth, and shade in abundance are the climatic conditions essential to promote luxuriant fern growth, it can and does continue when any or all these conditions are reduced to a minimum. 1 1 . Ferns may then be sought in any of the following situa- tions, and it will be seen that each situation has its charac- teristic species : A. Wet swamps or marshes with or without abundant shade. B. Rich woods, more or less moist. C. Uncultivated open places and dry hillsides. D. Moist, rocky ravines or rocky places not subject to sum- mer drought. E. Exposed rocky cliffs. F. Standing water. G. Growing on other plants. (Epiph3rtic.) 1 2. In the first location mentioned above, we may find the chain-ferns ( Woodwardid), many of the spleenworts {Aspleniutn\ a few of the shield-ferns {Dryopterts),x\it flowering- ferns (C>j;w«/»- dd), as well as the genera Acrostichunit Onoclea, etc. These in- clude some of our largest and coarsest ferns. A few more deli- cate in structure are also found here, notably the dainty Phegop- teris dryopterts. 1 3. In the second we find a few spleenworts, most of the shield-ferns, the beech-ferns {Phegopterii), most of the grape- ferns {Botrychtum), the maidenhair {Adtantum), Dtnnstadiia, and some others. In this situation we find the finest develop- ment of foliage and the greatest artistic finish among all the ferns. 14. In uncultivated places and on rocky hillsides we often find the common bracken or brake {Pteridtum), and also the lady-fern {Aspienium filtx-fcemtnd), though these are by no means confined to these locations, the latter growing quite fre- quently in moist woods, and even in cold, wet swamps. Many other ferns are found occasionally in openings of the forest or recent clearings, whbre they maintain a sickly existence, some- times for a series of years. In such locations ferns often be- come contracted and abnormal in growth, and take on a faded yellow hue from their exposure to the open sunshine. 1 6. In moist ravines and on rocky banks the bladder-ferns {Filix) may be found, with the peculiar walking-leaf {Camp' OUR NATIVE FERNS AND THEIR ALLIES. toscrus), the rare hart's-tongue (P Ay //tit's)* and many of the smaller spleenworts. The long, pendent fronds of our Ft'/t'x lmU>ifera add greatly to the beauty of our natural ravines, and often serve to conceal the uncouth rocks, or at least draw the attention to that which is more delicate and artistic. On dripping rocks, or where the sides of ravines are kept con- tinually moist by the spray of waterfalls, such delicate pellucid ferns as the filmy-fern {Trichomanes) and one Cryptcgratnma may be sought. There seems to be a direct connection between the environment and the texture of the fern. The last two mentioned g^ow in very damp situations, and are pellucid and almost membranous. The Fi/ix in somewhat drier situations is thinly herbaceous, while Asp/enium trichomanes and Camp^ iosorus, requiring less moisture, are more firm, and form the transition to the next group. 1 6. On dry cliffs we may look for the various si>ecies of Woodsta, the cloak-ferns {Notho/and), the lip-ferns {Cheilanthes), and the cliff-brakes (Pe//aa), Many of these are firm and even leathery in texture, and others are thickly covered on one or both sides with tangled hair or scales, fitting them to survive long periods of drought. 1 7. Only one of our native species is strictly aquatic, the anomalous Ceratopteris tha/ictroides found in Southern Florida, though Acrostichutn aureum is often found with its rhizoma rising from the water of salt marshes. Osmunda rega/is is oc- casionally found in standing water several inches deep, though this is not usual. 1 8. Among the epiphytic ferns are several species of Po/y' podium^ P. po/ypodioides, P. Scou/eri, and Ph/ebodiutn, the last always being associated with the cabbage-palmetto {Sabal pa/- mettd). Vittaria, Chei/ogramma, and Nephro/epis are also of this class, and are frequently pendent from the same plant, though occasionally found on other tree-trunks. Cheirog/ossa pa/mata, another peculiar tropical fern-ally, belongs to the same * This rare fern seems to show a decided preference for limestone rocks, and thus far has been found only above the geological formation known as the Comiferous limestone. I believe a thorough search for this fern along the outcrops of the formation in Central New York and elsewhere would show a wider distribution than is at present attributed to this tpedes. HAUNTS AND HABITS OF FERNS. 7 list. Even in the streets of Southern c\l\ts, Potypodium pofy- podioides is often seen growing with various mosses well up on the trunks of shade-trees. It is only in tropical regions, however, that epiphytes are seen in profusion. 1 G. These principles of climatic distribution are necessarily modified by the geographic range of species, which must be considered in this connection. For example, Dryopteris spinu- losa or its varieties form the leading foliage ferns of Northern New England and New York, and Dennstadtia^ less common in those localities, largely replaces them from Connecticut souths ward. This subject will be more fully discussed in a later chapter. LITERATURE. Most of the American literature bearing on this subject is in the form of short notes which have appeared from time to time in our two botanical monthlies ;* a classified summary appears below: mABiTS.— Botanical Gazette, 1,2 \ II, loo; 111,82; 17,140,177, 232 ; V, 27, 30, 43, 48 ; VI, 161, 295 ; Vli, 86. Dimorphism.— Torrey Bulletin, viii, loi, 109 ; ix, 6 ; xiii, 62. Forking Froj^bs.— Botanical Gazette, I, 50; 11,80; 111,39; VI, 220; y 111, 242.— Torrey Bulletin, Vll, 26, 85; IX, 116, 129; X,4. Relative Abundance :~ Davenport (George E.). A Bit of Fern Histoiy. In Botanical Gazette, vil, 60-64 (May, 1882). Cultivation :— Jackson (Robert T.). Cultivation of Native Ferns. In Garden and Forest, I, 317, 318; 330, 331; 340-342; 352-354 (Aug.-Sept. 1888). Robinson (John). Ferns in their Homes and Ours. X2mo, illustrated. Salem, 1878. A valuable outline of fern cultiva- tion, indispensable to those desiring to undertake the cultivation of ferns either in conservatories or Wardian cases. Smith (John). Ferns, British and Foreign, 8vo. Lon^ don, 1879. * Bulletin TVrriry ^^/antVra/ C/»4 (Columbia University, New York City) and the Botanical Ganett* (University of Chicago). Many notes in recent yean have appeared in The Fern Bulletin (Binghamton, N. Y.). V s OUR NATIVE FERNS AND THEIR ALUES, CHAPTER II. THE ORGANS OF THE GROWING FERN. Pour Uen savoir une chow, il faut en savoir les details. —La Rochefoucauld. 20. Every one familiar with the forest and its products must have seen the young ferns unrolling from the bud in spring and early summer. It will be noticed that the fronds are coiled from the apex to the base, and form crosiers, so called from their resemblance to the head of a bishop's staff. This method of vernation is called circinate^ and is rarely found except among ferns. In the grape-ferns and adder-tongues the vernation is straight or merely inclined, thus approximating that of ordinary flowering plants. 21. Rootstock. — Ferns usually spring from an under- ground stem called the rootstock. This may be simple or branched) smooth or scaly, horizontal, oblique, or even vertical. In some ferns it is fine and hairlike, while in others it is very large and stout. In some cases the rootstock creeps at the sur- face of the ground and even rises above it, as in the variety of Dry opt en's contermina which grows in Florida. In the tree ferns ot warmer climates it often forms a trunk fifty feet high, bearing the fronds at the summit, when it takes the name of caudex. 2.2,, Frond. — ^The aerial portion consists essentially of a leaf-stalk and blade ; the former is technically cdled the sttpe^ and the latter the frond. Though these are usually distinct from each other in appearance, the stipe is sometimes wanting, and in others no distinction can be made between, them. Both stipe and frond, or either one, may be glabrous (smooth), pubes- cent (softly hairy), hairy, woolly, or scaly ; when the scales are small and somewhat appressed, the surface is said to be squa- mous. The careful discrimination of these haiiy or scaly appendages becomes a matter of importance in distinguishing many of the species of Oheilanthes. In a few jf our native ferns AULD. products in spring are coiled rom their lethod of pt among mation is f ordinary aui under- simple oc n vertical. 3 it is very at the sur- variety of 11 the tree feet high, le name of tially of a d the stipe, ly distinct >s wanting, em. Both ►th), pubes- \ scales are o be squa- or scaly tinguishing native ferns THE ORGANS OF THE GROWING FERN, 9 the under surface is covered with a white or yellow powder bearing some resemblance to flour or corn-starch. For this reason a surface of this character.is cs\\t^ farinaceous. Such is the California gold-fern or " golden back " {Gytnnopteris trian- gularis), and several of the cloak-ferns {Nothokena), and such are the various gold and silver ferns of conservatories, including some of the richest and most beautiful in the world. 23. The frond may be simple, when it consists of a single undivided leaf, as in Phyllitis or Camptosorus ; ox compound, when it is divided into segments. The exquisite delicacy and the extent to which this dividing is carried in some ferns deter- mines laigely their aesthetic value. The continuation of the stipe through a simple frond is called the midvein ; through a compound frond is called the rachis, and is further distinguished as primary when the frond is much compounded. A frond is entire when the margin forms an unbroken line; when so cut as to form lobes extending half way or more to the midvein it is called pinnatifid; when these incisions extend fully to the midvein the frond is said to be simply pinnate, and the divisions are called pinnce. When the pinnae are cut into lobes the frond is bipinnatifid and the lobes are called segments, and when these extend to the secondary midveins it is bipinnate and the divisions are called pinnules. The secondary midvein then becomes a secondary rachis. In like manner we may have ferns that are tripinnatifid and tripin- nate, quadripinnatifid and quadripinnate. The last lobes are designated ultimate segments, and the last complete divisions ultimate pinnules. All these various forms from entire to quad- ripinnate are abundantly represented among our native ferns. 24. In some pinnate fronds, as in the oak-fern {Phegopteris dryopteris), the lower pair of pinnae is gfreatly enlarged and more compound than those above, so that the stipe appears to form three branches bearing similar and nearly equal portions. Fronds of this character are usually triangular or pentagonal in outline, and this method of branching is called ternate. It will be readily seen that this is merely a modified form of the ordi- nary pinnate frond. Throughout the domain of nature there is infinite variety of form and structure, and at the same time unity in plan and conformity to a few generalized types. 10 OUR NATIVE FERNS AND THEIR ALLIES. 26. Venation. — The method of veining admits of great variation, often serving to distinguish species, and more especi- ally the sections of the various genera. In some ferns, like most shield-ferns (Z?ry<7//^r/j),thc veins are free — that is, arising from either side of the midvein they do not unite with any other vein. In some of these the vein is simple (not branched), in others variously forked. In many the veins repeatedly anastomose or unite together, forming a series of network or areoloe. This may be somewhat irregular, as in Onoclea ; or forming a single row of areolae next to the midvein and thence free to the margin, as in Woodwardia Virginica; or forming many uniform areolae by the parallel transverse veinlets connecting the distinct and parallel primary veins, as in Campyloneuron phyllitidis. In case the venation does not appear when examined by reflected light, it may be brought out clearly by holding the frond between the observer and the light, and then using a lens if necessary. A few fleshy species require dissection to show the veins. CHAPTER III. FRUCTIFICATION IN FERNS. " But on St. John's mysterious night, Sacred to many a wizard spell, The hour when first to human sight Confest, the mystic fern-seed fell." 26. Spores and Sporangia. — In the flowering plants (Spermaphytes) seeds are pro- duced by a complex process in- volved in pollination.the growth of the pollen tube, and the sexual process which results in the em- FiG. I.— Bniarffed section through abryo of the new plant. The sorus of Polypodium falcaium Kellosrg, -, ^i. ^ j showing the stalked sporangia. r erns, on the contrary, produce no flowers. Instead of seeds developed from fertilized ovules, minute spores are produced asexually, from which new ferns are developed by a peculiar process of germination very unlike that FRUCTIFICATION IN FERNS. II of flowering plants. These spores are collected in little sacs known as sporangia or spore cases. The sporangia in the true ferns (POLYPODiACEiE) are collected in little clusters on the back of the frond, or are variously arranged in lines along the veins or around the margins (Fig. i). These clusters of spo- rangia are called sort, and may be naked, as in Potypodium, or provided with a special covering known as the indusiuniy as in Dryopttris (Fxf^.Z). The various forms of the sori and indusia serve as the basis for classifica- tion into genera and tribes, while each sub-order has its peculiar form of sporangia. 27. In the PoLYPODiACEiE the sporangia are more or less completely surrounded with a jointed vertical ring or annulus, and at maturity burst open transversely by the straightening of the annulus and discharge their copious spores (Fig. 2). The clusters of sporangia are said to be marginal, intramarginal, or dorsal, according as they have their position at the margin or more or less remote from it. They may be . ^ roundish, oblong, or linear in shape, or arranged dium vuiga't^i^^ in variously forking lines, or may even be spread J^res'^^wfiLch en' in a stratum over the entire under surface of the '"k*^- frond. They are called indusiate or non-indusiate according as they are covered or naked ; and the indusia may be inferior (at- tached below the sorus), as in Woodsia (Fig. 9), or superior, as in Drjfopierts iF'ig.S)t or of various intermediate methods of at- tachment. 28< In the other families of Filicales the sporangia are variously arranged. In the Hymenophyllace^ or filmy ferns the flattened spo- rangia are sessile along a filiform receptacle, and are surrounded with a complete transverse annulus. At ma- turity they open vertically Fica.— Sporen- Fig. 3. Fig. 4. Fig. 3.— Enlarged sessile sporangium of Trichomanes radicans Swz. Fig. 4.— Sporangium of SchtMaa/w silia Pursh, showing the apical ring. Much enlarged. (Fig. 3). In the Sq^izmacem the sporangia are ovate, sur- V 13 OUR NATIVE FEKNS AND THEIR ALLIES, rounded at the apex by a complete annulus. and open by a longitudinal slit (Fig. 4). In the OsMUNDACEifi or flowering ferns the sporangia are larger, globose, and naked, with the mere trace of a transverse annulus, and open longitudinally. The various methods of fructiflcation can be best understood by describing the peculiarities of the various genera in regular succession and noting the variations occurring in the sections or sub-genera. By this means we will arrive at a better under- standing of the principles of fern classification as discussed in a future chapter. As the subject of venation is closely connected with that of fructification, it will be treated in the same connec- tion. 29. Aorostlohum. — In this genus the sporangia are spread in a stratum over the under surface of the upper pinnae in our solitary species, but in some exotics they cover portions of the upper surface as well. There is no indusium. 30. Polypodium (Fig. i). — Formerly all ferns agreeing in the possession of roundish naked sori were placed in this genus notwithstanding the fact that the venation was widely different ; it seems more logical to regard some of these sections as genera. In § EuPOLVPODiUM the veins are free, yet are occasionally known to unite,'*' thus indicating a tendency to vary toward the next section. The sori are generally found at the end of a free veinlet. In § GONIOPHLEBIUM the veins unite near the margin, form- ing large areolae, each containing a single free veinlet which bears the sorus at its end. A tendency to variation is seen in P. polypodioides, whose veins are free, as well as in P. Calif or- nicum in which they are often partly free. 30a. Phlebodlum. — In this genus ample areolae are next the midvein, and frequently in one or more secondary rows, each bearing a single sorus at the junction of two or more vein- lets. A large number, however, bear the sori at the end of a single veinlet. From the fertile areolae to the margin the veins anastomose more copiously. * Catalogue of the Davenport Herbarium, p. 8. FRUCTIFICATION IN FERNS. 13 3Q|b. Campylon«uron has areolae, each usually bearing two sori ; they are found between the parallel primary veins which extend from the midrib to the margin. 3 1 . Qymnopterls. — In this genus the sori follow the course of the veins, and consequently vary with the venation, being simple, forked, pinnated, or anastomose with each other. The sori arc non-indusiate. 32. Nctholaena. — In the cloak-ferns the sori are marginal, and provided with no indusia. This genus is linked very closely to Gymiiopteris on one hand and to some species of Cheilanthes on the other. From the latter it is separable only by the ab- sence of the marginal indusium ; the two are likely to be con- founded by beginners. 33. Cheilofframma has simple fronds, the fructification in a continuous sub-marginal line near the apex of the frond. 34. Vittaria. — This peculiar genusoccupies a somewhat in- termediate position between the indusiate and non-indusiate genera, and while usually associated with the latter has consid- erable claim to be ranked with the former. The fronds are nar- row and grass like, bearing the sporangia in an intramarginal groove, often more or less covered by the inrolled edge of the frond. The venation Is very obscure. 35. Adiantum (Fig. 5). — The maidenhairs have a peculiarly smooth foliage, and usually possess no midvein. The veins are usually flabellate, and after forking one or more times bear the sori at their extremities. The margin of the frond is reflexed, thus forming an indusium which bears the sporangia on its under surface. p«g. s-— a segment of ,_. ,. _ , . Adiantum, showine the 36. Pterls (Fig. 6). — In this genus, sori covered by indusia formed by reflexions of the now excluding the common brake, the mI^lno?^thefrond--From otherwise free veins are united by a fili- Le Maout and Decaisne form receptacle which bears the sporangia. This continuous marginal line of fructification is covered by a membranous in- dusium formed of the margin of the frond. 37. Cheilanthes. — The lip-ferns found within our limits are unequally divided among four sections, all agreeing in bear* 14 OUR NATIVE FERNS AND THEIR ALLIES. ing the aori at or near the ends of the veins, covered by fan in- dusium formed of the margin of the frond. In § Adiantopsis the indusia are distinct, and confined to a single veinlet. One of our species varies from the typical species of this section, and has even been assigned to a separate genus. In § EUCHEILANTHES the indu- Fio. i.^Fttrit hngi/»iia L. Bn- sia are more or less confluent but iTeWlk^ceptwWnder th* not continuous, usually extending nal iadutium. ^^^^ j|,g apjces of several veinlets. In § Physapteris the ultimate segments are bead- like, &nd the indusium is continuous all round the margin. S Aleuritopteris has the fronds farinose below, and m< eludes a single species somewhat doubtfully assigned to cur limits. 38. Cryptofframma has dimorphous fronds, the margins of the fertile being closely rolled toward the midvein, thus cov- ering the confluent sori. At maturity these open flat in order to discharge the spores. 39. Pellsea has representatives of three sections within our limits, all agreeing in possessing intramarginal sori, which finally became confluent and form a marginal line covered by an indusium formed of the margin of the frond. § Cheiloplecton includes herbaceous species with visible veins and broad indusia. § Allosorus includes coriaceous species having wide indusia. while § Platvloma includes species similar in texture, but with extremely narrow indusia and broad segments. 40. Ceratopteris is an anomalous genus from southern Florida, having a few sori arranged on two or three veins par- allel to the midvein, and covered by the broadly reflexed margin of the frond. It properly forms the type of a family. 41. Struthlopteris (Fig. 7) is intermediate between those genera in whirh there is an indusium formed of the revolute margin of the frond and those in which the indusium is remote from the margin. Our single species has dimorphous fronds, free veins, and the fructification in a broad band next the mid- I \ FRUCTIFICATION tN FERNS. IS by ble vein, covered by acuntinuous and distinctly intramarginal indu sium. This genu and each sporangium containing 50 spores, making a total of 18,000,000 spores. The copious green spores of Osmunda cinnamomea, or the pale-yellow, powdery spores of a well-developed specimen of Botrychittm Virgtnianum, must far exceed this computation. By drying either of these species under pressure between sheets of paper great quantities of the spores may be obtained for examination. Specimens for this purpose should be selected just before the sporangia reach their maturity. I CHAPTER IV. GERMINATION OF FERN SPORES. Alle Glieder bilden sich aus nach ew'gen Gesetzen, Und die seltenste Form bewahrt im Geheimniss das Urbild. -Goethe. W 61 . The germination of the fern spores usually takes place a considerable time after they are discharged from the sporangia, but in Osmunda, which develops its fruit early in the season, they commence their growth only a few days after dissemination. 62. Thalloid Phase. — In germination the exospore splits along the side, and the protruding endospore, sometimes with its divisions already formed by septa or partitions, forms, not a fern, but a thalloid structure resembling one of the lower v^ 20 OUR NATIVE FERNS AND THEIR ALLIES. Figs, is, 13.— Prothalliumof/Vmx Linn. liverworts called the prothallium. Different ferns vary in the method of forming this prothallium, some producing it im- mediately at the spore and others after the formation of a thread- like growth known as the pro- embryo. The prothallium is en- tirely composed of cellular tissue, and in the true ferns (POLYPO- DiACEiE) is broadly cordate or reni- form in shape, and bears large numbers of root-hairs from the under part of its posterior portion (Figs. 12, 13). The prothallium varies in size from less than one tenth of an serruiaia Linn. f.. showing two mch up to one third Of an mch stages of growth. (After Moore.) j^ {jg ^j^est part. On the under surface of the prothallium two sorts of organs are produced which represent the male and female structures, respectively known as antheridia and archegonia. The position of these organs on the prothallium varies in different sub-orders. In some species, notably the ostrich-fern, the two kinds of sexual organs are produced on separate prothallia, so that the plant becomes dioecious instead of monoecious. In nurseries where ferns are grown for sale immense quantities of prothallia are regularly developed from spores. 63. Antheridia. — These are small masses of tissue developed in the same manner as the root-hairs, consisting of a single layer of cells forming the wall, and containing a number of spirally coiled threads, usually with a number of cilia on their anterior coils. At maturity the antheridium swells by the ab- sorption of water and finally bursts its wall, discharging these coiled filaments, which possess the power of locomotion, and for this reason are called antherozoids. These antherozoids often drag with them a little vesicle which seems to play no part in the process of reproduction (Fig. 14). 64. Archegronia. — The archegonium (falsely called pistil- lidium) is also a rounded mass of tissue usually less prom- GERM IN A TION OF FERN SPORES. 21 inent than the antheridia, consisting of an external layer of cells and a large central cell, which soon divides into two. The lower portion, at first the larger, develops into a roundish cell, which is analogous to the ovum among animals, and is crilled the oosphere. The upper portion of the central celi develops between those composing the neck of the archegonium into a canal filled with a sort of mucilage ; this finally swells up, forces the cells of the neck apart, and is expelled to aid in attracting Fig. 14.— Antheridium of Adiantum Fig. 15.— Young archegonium of Ptiria capillus-veneris L., showing' the es- strrulata Linn, f., showing oSsphere, caping antberozoids. (After^achs.) neck, and canal-cell. (After Sachs.) iind retaining the antherozoids at the neck of the archegonium. The oOsphere is thus left exposed (Fig. 1 5). 65. Fertilization. — The antherozoids, analogous to the sperm-cells, when discharged from the antheridium swim m the moisture always present on the under surface of the prothallium, swarm in large numbers around the neck of the archegonium, and are retained by the mucilage. Some finally force their way into the canal of the neck, a few reaching the oSsphere and disappearing within its substance. There is thus a true sexual generation among ferns, and the formerly appro- priate term Cryptogamia (hidden marriage) loses its application under the untiring scrutiny of the microscopist. After fertili- zation the neck of the archegonium closes, and the fertilized aa OUR NATIVE FERNS AND THEIR ALLIES. Jiss oOsphere, now called the odsfiore, increases in size, and finally develops into a true fern. 66. Pterldold Phase. — After the o5- sphere has been fertilized it commences its growth by the ordinary processes of cell multiplication, and for a time remains with, in the walls of the archegonium, which continue to grow, until finally the interior growth breaks through the walls, differen- tiated into its first root and leaf. The young fern draws its nourishment from the pro- thallium for a time, but soon develops root-hairs, which, extending into the soil, maintain thereby an existence independent of the prothallium. The latter growth hav- FiG 16 —Adianium ing accomplished its work, withers away i^xX'^yovkk fem (F^g- ^6). The first parts of the root, stem, seen from below; //, and frond are very small and comparatively prothalltum; b, first leaf; ... "^ , , , . , A, root-hairs of prothai- simple in Structure, but those formed later 8«:ond"'*rTOt8.*"(AfSr are successively larger, and not only bear 5***"-> a closer resemblance to the mature form of the species, but also develop increased complexity of struc- ture. " The fern continues to gain strength, not by subsequent increase of size of the embryonic structures, but by each succeS' sive part attaining a more considerable size and development than the preceding ones, until at length a kind of stationary condition is arrived at, in which the newly formed organs are nearly similar to the preceding ones." 67. The complete life-history of a fern illustrates a principle common among the lower forms of animal life known as " alter- nation of generations." Instead of the direct production of a mature sexual plant, as among the higher forms of vegetation, there is the production of a sexual growth resembling a lower form of vegetation, which in turn is followed by the growth of a mature plant producing its fruit without theassistn.i;cc of sex- ual organs. 68. Recapitulation. — To review the life-history of a fern we find the following processes : \\ GERMINA TION OF FERN SPORES. 23 ' a fern A. Production of the spores asexually by the mature plant. (Fructification.) B. Growth of the prothallium from the spore with or with- out the development of a pro-embryo. (Thalloid Phase.)* C. Production of sexual organs, archegonia (female) and ftntheridia (male), on the under surface of the prothallium, 01 on separate prothallia. D. Fecundation of the oOsphere developed in the arche* gonium by the antherozoids developed in the antheridium. (Fertilization.) E. Growth of the mature fern in successive stages from the oospore. (Pteridoid Phase.)* LITERATURE. Bessey (Charles E.). Botany, pp. 361-388. New York, 1881. (Henry Holt & Co.) GoEBEL (K.). Outlines of Classification and Special Mor- phology of Plants, pp. 189-298. (English Translation.) Ox- ford, 1887. (Macmillan & Co.) Campbell (Douglas H.). Fern Notes. In Torrey Bulletin, X, 118, 119. (Nov. 1883.) The Development of the Prothallia in Ferns. In Botan- ical Gazette, x, 355-360, with Plate ix. (Oct. 1885.) The Structure and Development of the Mosses and Ferns. 8vo. London and New York, 1895. (Macmillan & Co.) This also contains an extensive Bibliography of the entire subject. * The terms " Thalloid Phase" and ** Pteridoid Phase" in place of the older terms " sexual generation" and " asexual generation" were first sug- gested in the first edition of this work (1881). The older terms, although in common use by botanical writers, are decidedly unfortunate and misleading. A generation is properly the production of offspring resembling the parent, or the offspring thus produced, which the prothallium is not and the mature fern is not. The generation proper must then be considered as including the entire life-history of a fern, of which the prothallium and mature fern are suc- cessive phases. The terms "sexual " and "asexual " as used in this connec- tion are likewise misleading, as they might apply as well to the origin as to the producing power of the so-called " generation." The prothallium is asex- ual in origin, but develops sexual organs ; the mature fern, on the other hand, produces no sexual organs, but is itself the product of bisexuality. It is now more common to speak of these two phases of growth as the gametophytt and sporophyte respectively. v 24 OUR NATIVE FERNS AND THEIR ALLIES. Campbell (Douglas H.). On the Development of the Antheridium in Ferns. In Torrey Bulletin, xiii, 49-52, with Plate Liv. (Apr. 1886.) The Development of the Ostrich- fern. In Memoirs Boston Society of Natural History, IV, 17-52, with Plates iv- VII. (Apr. 1887.) On the Prothallium and Embryo of Osmunda Clay- ton iana L. and O. cinnamomea L. In Annals of Botany, vi, 49-94, pi. iii-vi (1892). CHAPTER V. FERN STRUCTURE. Be it oiirs to meditate, And to the beautiful order of thy works Learn to conform the order of our lives. —Bryant. 69. Tissues. — The life-history of every plant commences in a single cell, and all the complications of vegetable growth depend on two simple processes, viz., the enlargement of indi- vidual cells to their full size, and their multiplication by divi- sion. The lowest forms of vegetable life consist of a single cell, either globular or elongate. Those of a somewhat higher grade consist of a single row of cells, or at most a single layer ; while still higher forms of growth consist of masses of cells variously grouped together and specialized by differentiation from the typical form and character. 70. Cells become specialized or set apart to fulfil a certain function in the economy of plant growth in many ways. Some are lengthened for giving strength to stems or leaves; some have their walls thickened to give rigidity or hardness where protection is needed from injury to more delicate structures within ; and some are variously adapted for containing and dis- tributing the secretions or other fluids connected with the cir- culatory system of plant life. Seven distinct varieties of tissues are recognized by structural botanists, yet some of these are connected with each other by various gradations. FERN STRUCTURE, 25 7 1 . Tissue Systems. — The earliest tendency to differen- tiation of cells is seen in the arrangement of the outer row of cells to form a boundary wall. In higher forms of growth the interior cells tend to form one or more series of string-like rows surrounded by the normal cellular tissue. We thus reach the basis of the classification of vegetable tissues into three groups : (a) Epidermal Syste.n. (b) Fibro-vascular System, (c) Fun- damental System (Fig. 17). The first and third are common to both ferns and mosses. The second is first seen in the ferns and their allies, where it is a character so constant that it serves as the basis for separating the so-called " vascular" cryptogams from other flowerless plants. These three forms of tissue may be seen by examining a thin cross-section of the stipe of a living fern with the microscope. Longitudinal sections will show still further the character of the tissues composing the fibro-vascular bundle. 72. Roots. — Roots are constantly produced as the root- stock advances, and consist for the most part of little fibrils which are naked for a short distance from the apex in order that they may freely absorb moisture from the earth. The epi- dermis is also thin, and usually consists of a single layer of small cells. It differs from that of the rest of the plant in having no stomata i^i). As the apex continues to grow, the epidermis of the part behind becomes harder, and frequently develops hairs, or more frequently irregular scales. 73. Stipe. — The stipe is made up of the three forms of tissue (Fig. 17), and usually con- tains several bundles of vascular tissue. In the dried stipe these can be easily seen, by scraping off the external covering of the stem. These bundles of fibres give sta- bility to the fern, and are con- tinued through the rachises and veins, thus forming the frame- p,^ ^...cross-section of stipe of work for the softer portions of i^«7i:r/r«^tVw (L.) Underw., showing ^, , J rrii ^. two bundles of fibro-vascular tissue. the frond. The stipes are some- times smooth and polished, sometimes hairy or beset with stalked glands, and sometimes densely clothed, especially near the base, with chaffy scales. 26 OUR NATIVE FERNS AND THEIR ALLIES. 74. Frond.— In the HymenophyllacEvE the frond con- sists of a single layer of cells. This condition is also found in the leaves developed along the axis of growth among the mosses to which this sub-order is related in some of its forms. In all other ferns there are several layers of cells variously compacted together, and forming all the varieties of texture — membranous, herbaceous, coriaceous and fleshy. The epi- dermis is usually easily separable from the underlying tissue, when its peculiar markings can be studied. 76. From the epidermis a great variety of appendages are developed which are all modifications of hairs, and are all in- cluded under the tern* trichomes, however different in appear- ance or distinct in function. These are not confined to the frond, but develop here their greatest variation. They a j fre- quently found on the roots, the rootstock, and the stipe, under the form of root-hairs or scales of various forms. Scales are especially abundant in certain forms of Dryopteris, as well as in Polystichum, Cheilanthes, and other genera. 76. Trichomes. — On the fronds the trichomes may be de- veloped as simple unarticulated or articulated hairs, consisting of one or two cells at most. They may i\ppear as stalked glands like those that arise from the stipe of Cheilanthes Coopera or the margin of the indusium of Dryopteris spinulosa^ var. intermedia ; or they may be developed into scales of in- tricate cellular structure like those on the under surface of cer- tain forms of Cheilanthes, particularly C. Fendleri and C. Clere- landii. In the true ferns the sporangia are specialized, tri- chomes developed in clusters {sori) along the veins, or spread over the entire surface of the frond, or even arranged in spikes or panicles. The epidermis also develops an excrescence known as the indusium, which consists of a single layer of cells, and is variously arranged as indicated in Chapter III. In some cases a false indusium is provided, which is not a growth from the epidermis, and may consist of several layers of cells. 77. Stomata. — If the epidermis covering the under surface of a fern be examined under a high magnifying power, peculiar structures will be seen in the form of semi-elliptical or crescent- shaped cells connected at their apices and separated between. These are the guard-cells of stomata which control the open- FERN STRUCTURE. «7 ings to the air-chambers of the plant. The two elliptical ccU^ form the mouth of the passage and expand when moist, allow- ing the atmospheric gases and watery vapor to escape or enter but close the entrance by contraction in time of drought. The stomata are not confined to the fronds, but are found to a greater or less extent on all aerial portions of ferns and higher plants, as well as on subterranean stems. 78. Asexual Propasratlon* — Besides the ordinary meth- ods of sexual reproduction discussed in Chapter IV., most ferns are propagated by growth of therootstock under ground, giving rise to a succession of fronds each season. In addition to this, which is common to all perennial plants, there are some meth- ods of reproduction that deserve attention. The first is by 79. Buds and Bulblets. — In a few species of conservatory ferns adventitious buds are produced on the surfaces of the fronds. These soon develop into young ferns, and it is not un- common to see a large number in vari- ous stages of growth rising from a sin- gle frond. This peculiarity is common among several species of Asplenium, especially A. furcatum Thunb., and will be sometimes found to occur among some of our native species. Bulblets are found in the axils of the upper pinnae of our Filix bulbtfera, which often fall to the ground and develop into new plants after a manner analogous to the devel- opment of the axillary buds of the tiger- lily. 80. Another method is seen in the walking-leaf {Camptosorus rhizophyllus), in which the long, attenuated, simple fronds bend over and take root in the adjoining soil in a manner quite analo- Fig. iS.—Cam^tosorus rAi- •' , .... zopkyllus Link., reduced, gous to the propagation of strawberries showing peculiar method of byrunners(Fig. i8). The same method P'°P''&^»i°"- of rooting at the apex has also been noticed in Asplenium pinnatifidum, A. piatyneuron, and Phegopteris reptans. 1 38 OUR NATIVE FERNS AND THEIR ALLIES. LITERATURE.* Bessey. Botany. (See p. 23.) GOEBEL. Outlines of Classification. (See p. 23.) De Bary (A.). Comparative Anatomy of Phanerogams and Ferns. 8vo. Oxford, 1884. (Macmillan & GOi) Campbell (Douglas H.). A Third Coat in the Spores of the Genus Onoclea. In Torrey Bulletin, xii. 8, 9 (Jan. 1885). Schrenk (Joseph). The Dehiscence of Fern Sporangia. In Torrey Bulletin, XI 11, 68, 69 (1886). Lyon (Florence May). Dehiscence of the Sporangium of Adiantum pedatum. In Torrey Bulletin, XI v, 180-183 (Sept. 1887). Atkinson (George F.). The Study of the Biology of Ferns by the Collodion Method. 8vo. New York, 1894. (Macmillan & Co.) CHAPTER VI. THE FERN ALLIES. Beneath my feet The ground-pine curled its pretty wreath. — Emerson. A. THE GRAPE-FERNS AND ADDER-TONGUES. 8 1 . General Characters. — These peculiar plants, former- ly united with the true ferns, are now regarded as constitut- ing a distinct botanical family. They include mostly small, fleshy, terrestrial plants, and, like ferns, may usually be found in swamps or rich, moist woods. As already noticed (5), there is a marked tendency to variation in the same species, atid numerous varieties have been established from the various forms. 82. The sterile and fertile portions of the plant are borne on a common stalk, and either portion may be sessile, long * Sett other references at close of Chapter X. ! \ a V,, THE FERN ALLIES, 29 In Ophioglossum or short stalked, in the various species, the sterile portioti is simple, and in all our species appears like a leaf rising *rom the common stalk. C/tet'roji^l>>ssa has several spikes. In Botrychium (Fig. 19) the sterile segment (except in some forms of B. simplex) is somewhat pinnatcly or ternately divided, and in the larger forms of B. Virginianum is broad- ly ternate, with the divisions even tri — quadripinnatifid. The veins are free in the latter genus, but anas- tomose in the former. This charac- ter, however, is frequently obscured by the fleshy texture of the plant. 83. Vernation. — As has been before stated, ferns are rolled in the bud from the apex downward («>- a'nate), distinguishing them from the higher forms of vegetation. Among the OPHiOGLOSSACEiE, how- ever, the vernation is either straight, inclined at the apex of one or both segments, or else the fertile seg- ments are folded on the main stalk, making the vernation wholly in- clined. Until recently there has been much difficulty in distinguishing the smaller species of Botrychium, and some forms seem to connect the smaller ones with the reduced forms of B. obliquum and B, Virgini- anum. Mr. Davenport has investigated the bud characters of these intimately related species, and has made their identification a matter of comparatively easy investigation. The buds may be found enclosed in the base of the common stalk (except in B. Virginianum^ where they are placed in an upright cavity at one "-*i Fig. 19. — Plant of Botrychium lunaria^ natural size. 30 OUR NATIVE FERNS AND THEIR ALLIES. ^- i side), and may be examined with a strong lens. The three di- visions are summed up as follows : "I. Vernation wholly straight. B. simplex Hitch. (Fig. 31). II. Vernation partly inclined in one or both portions. B. lunaria Swz. (Fig. 32), B. boreale Milde, B. neglecttim Wood (Fig. 33), and B. obliquum Michl (Fig. 34) and its allies. III. Vernation wholly inclined, in the fertile frond recurved. B. lanceolatum Angs. (Fig. 35), arnd B. Virginianum Swz." The special characters of each species will be found under the descriptions of the Botrychia later in this work. The cuts will be valuable for reference, and will enable even beginners to identify the species of this complicated genus with compara- tively little difficulty. 84. Fructification. — In this order of plants the fructification consists of sporangia, which^ unlike those of the true ferns, are not reticulated, possess no trace of a ring, open by a transverse slit, and are variously spiked and panicled (Fig. 20). In the adder-tongues {Ophioglossum) the sporangia are large, and cohere in two ranks along the margins of a single spike, opening transversely to discharge their copious sulphur- yellow spores. In the grape-ferns {Botrychium) the sporangia are globular and arranged in double rows along the narrow segments, more or less in panicles. In both genera the sporangia are not developed from the epidermal cells, but arise from a transformation of the interior tissue of the leaf. This, with other characters as clearly defined, serves to separate these anomalous plants from the families of true ferns. 85. Germination. — Among the OPHiOGLOSSACEiE, so far as known, the prothallia are destitute of chlorophyll, develop under ground, and are monoecious. In Botrychium lunaria the prothallium is an ovoid mass of cellular tissue, light brown without and yellowish white within. It produces a number of antheridia and archegonia on the upper surface as well as the lower, differing in a few minor points from the true ferns in the method of their development. Fig. 20. — En- larged sporangia of Botrychium Virtiinianuni Swz. THE FERN ALLIES. 31 LITERATURE. Hooker (W. J.) and Baker (J. G.). Synopsis Filicum, pp. 444-448. MiLDE (J.). Botrychiorum Monographia. In Verhandl. der k.k. zool. bot. Geseilschaft, '^ivni, 507-516 (1868); xix, 55-190; Tafel VII, VIII (1869) ; xx, 999-1002 (1870). Davenport (George E.). Notes on Botrychium simplex. 4to, paper, with plates (1877). Vernation in Botrychia. In Torrey Bulletin, vi, 193- 199, plate (1878); VII, 115,116 (1880); viii, 100, loi (1881). Cf. also XII, 22, 23. Campbell (Douglas H.). The Development of the Root in Botrychium ternatum. In Botanical Gazette, XI, 49-53, with plate (March, 1886). A Method of Spore Germination. In Botanical Ga- zette, X, 428 (1885). Jeffrey (E. C.)- The Gametophyte of Botrychium Virgi- nianum. Ann. Bot., xi, 481-486 (1897). Prantl (K.). Beitrage zur Systematik der Ophioglosseen. \njahrb. des Kon. Bot. Garten {Berlin), III, 297-350 (1884). B. THE HORSE-TAILS. 86. General Characters. — The horse-tails or scouring- rushes belonging to the genus Equisetum are perennial, rush- like plants, that may be found in damp, gravelly, or loamy soil, some species even growing in shallow water. Our native species vary in height from a few inches up to eleven feet, as seen in some of the larger forms of E. robustum. In some species only the root is perennial, the stems which are sent up for producing fruit dying down to the ground every year. In others the stems are evergreen, continuing through the winter. Some species, like the common horse-tail {E. arvense), are dimorph- ous, the fertile stems being simple and destitute of green color- ing matter (chlorophyll), while the sterile stems are green and copiously branched, The fertile stems of some other spe- cies, as E. silvaticum, which are simple at first, after maturing their fruit produce branches and resemble the ordinary sterile stems (Figs. 21, 22). •'f m I ii 32 OUR NATIVE FERNS AND THEIR ALLIES. 87. The furrowed stems are hollow, except in E. scirpoides, and in addition to the large central cavity there is a series of smaller air-cavities opposite the furrows known as the valleciilar canals, the furrows themselves being called vallecula and the ridges carince. Opposite the carinas there are still smaller cavi- ties known as carinal canals. The carinse vary in number from I \ Figs. 31, •M.—Equisttum silvaiieum L., showing sterile and fertile stems. (Prom Thome.) five to fifty in different species. The stems are also jointed, and at each node some species produce a whorl of branches which may bf; simple or compound. Some species, however, like the common scouring-rush {E. hiemale), produce simple stems. 88. The leaves are produced also at the nodes, and by the union of their margins form a short sheath which ends in a row THE FERN ALLIES. 33 ides, s of ular the avi- rom -\ of teeth. These teeth may be deciduous or persistent, and their number, varying from three upwards, indicates the number of leaves forming the united whorl. 89. Stomata {^^^ are distributed along the valleculae either irregularly or disposed in ranges on either side of the valleculae. The epidermis frequently contains much silica, and the rough- ened tubercles of some species give the surface a harsh feeling. 90. Fructification. — The fructification in Equisetum is arranged in cone-like spikes borne at the apex of the fertile stems. These spikes are composed of successive closely-placed whorls of shield-shaped, stalked scales or modified leaves, each of which bears from five to ten one-celled sporangia on its under side. The sporangia open along the inner side to discharge their numerous spores, whose outer coat is spirally split into two bands, forming the so-called elaters. The elaters when dry are spread out at right angles to each other in the form of a cross, and probably assist in scattering the spores ; when moist they rapidly absorb water, and become closely coiled around the spore.* 9 1 . Germination. — The spores of Equisetum retaining their powers of germination only a few days, soon develop branched and irregularly lobed prothallia, which are provided with chlorophyll. These are usually dioecious, the male being smaller, and producing antheridia at the end or margin of the larger lobes. The antherozoids are large, and provided with a peculiar appendage known as the " float." The female prothal- lium may reach one half inch in length, and develops archegonia on the anterior margin of the fleshy lobes. The process of fertilization is similar to that of ferns. nd ch he •: * An interesting illustration of this can be seen by placing a mass of fresh spores on a slide uncovered, and examining it with a low power. By breath- ing on the slide the elaters coil closely about the spore ; as soon as the moist- ure evaporates they uncoil, and in their activity jostle each other in great confusion. 3 uf he )W 34 OUR NATIVE FERNS AND THEIR ALLIES. i LITERATURE. Baker (J. G.). Fern Allies, pp. i-6 (1887). Braun (Alexander). A Monography of the North American Species of the Genus Equtsetum. With additions by George Engelmann, M.D. In Stlliman's Journal, XLVi, 81-91 (1843). Describes the then known North American species of Equtsetum. Campbell (Douglas H.). The Development of the Male Prothallium of the Field Horse-tail. In American Naturalist, XVI, i-io (Jan. 1883). MiLDE (J.). Monographia Equisetorum. 4to, pp. 607, with 35 plates. Dresden (1865). Newcombe (F. C). Spore dissemination in Equtsetum. In Botanical Gazette, xiii, 173-178 (1888). C. THE CLUB-MOSSES. 92. General Characters. — The club-mosses are chiefly small perennial plants usually growing in dry or moist woods, or even on exposed rocks with little soil for nourish- ment. Most of the species are somewhat moss-like in habit, as might be suspected from the popular names given to these plants, the genus Lycopodium taking the name of club-moss and Selaginella that of rock- moss. Various species of Lycopodium are also known as ground-pine, ground-fir, ground-cedar, running-pine, etc., from more or less marked resemblances (Fig. 23). In the curious Sela-^ ginella hpidophylla from Arizona the branches of the closely coiled central stem roll up when dry into a nest-like ball, and when moistened expand so as to appear flat or saucer- FiG. 23. — Portion oi T ycopodium clavatuni L, ^ natural size. (After Prantl.) THE FERN ALLIES. 35 shaped. As the plant retains this power indefinitely, it has sometimes been called " the Resurrection-plant." 93. The stems are usually creeping, yet in some species show a tendency to become erect, and most species send up erect branches which bear the fruit. Most species bear roots at irregular intervals along the under side of the creeping stems, but our solitary species of Psilotum is rootless, bearing only underground shoots which perform the functions of roots. The leaves are small and unbranched, in some instances resembling appressed scales, in others resembling the acicular leaves of Conifers, and are arranged in four, eight, or many ranks. In some species the leaves are of one kind, while in others two or even more forms may occur on the same plant. In Psilotum the leaves are all rudimentary. 94. Fructification. — The fructification of the club-mosses is chiefly borne on upright branches in solitary or clustered (2-5) spikes, which are formed of numerous scales or scale-like leaves, each bearing a single large sporangium in its axil. The sporangia open transversely, and are one-celled, except in Psilotum^ where they are three-celled. In a few species of Lycopodium the sporangia are borne near the summit of the fertile stems in the axils of ordinary leaves. The usual shape of the fruit-bearing scales is represented in Figs. 24-26. 95. The spores of Lycopodium and Psilotum are of one kind (Fig. 24), but in Selaginella two kinds of spo- rangia are developed — the microspo- rangia, producing numerous micrO' spores (Fig. 25) not unlike the spores of Lycopodium; and the macrospo- rangia, producing usually four macro- Ljc?^di^lfc^rJiiaS^ {:\ spores (Fig. 26), so called from their ''^MS^^^.'lSsfriS"?: larger size. This character of Sela- tile spike of Stiagineiia rupet- • // V.' \^ '„ t, -..u ..u n ''•«> Spring,disclo8ing two sorts gtnella, which it shares with the quill- of spores. (After Sprague.) worts and pepperworts soon to be described, serves as the basis for the division of the fern allies into two groups : the I? p ,si $6 OUR NATIVE FERNS AND THEIR ALLIES. isosporous, producing spores of one kind ; and the heterosporous, producing spores of more than one kind** 96. Germination. — The germination of Lycopodium is only partially known, as the prothallia have been seen in only three species, and in these they have not been carried through all the stages of development. That of L. annotinum is a yel- lowish-white mass of tissue with a few small root-hairs.t The antheridia and archegonia are developed from the upper side of the prothallium. In L. cernuutn, TreubJ found the pro- thallia much smaller (one twelfth of an inch long), vertical in growth, yellowish below and bright green above. The anthe- ridia and archegonia are found round the summit of the cylin- dric prothallium. 97. The germination of Selaginella is better known. The contents of the ripened microspores are transformed into a mass of tissue consisting of a few cells, one of which remains sterile and is considered a rudimentary prothallium, while the others give rise to antherozoids, and are consequently considered as a rudimentary antheridium. The macrospores, on the other hand, produce a many-celled prothallium, which develop a few root-hairs and numerous archegonia, which after fertilization give rise to a new plant. Two plants are sometimes produced on the same prothallium. 98. The microspores are thus seen to be male and the macrospores female, showing a clearer differentiation of sex in the products of the mature plant than appears in any other group of the fern allies already studied. This may be consid- ered a foreshadowing of the vastly more complicated repro- ductive processes of the flowering plants. In the method of formation of the embryo the Selaginella also differs from all other plants of this group, and approaches the flowering plants. \' * This division, though used by some of the best botanists, is at best an artificial classification, as it separates genera otherwise closely allied to each other. ^ Cf. "i- Fankhauser, Botanische Zeitung, 1873, pp. 1-6; Bruchmann, Botanisches Centralblatt, xxi (1885). X C/. Trevh^ Ann. d. Jard. Bot. d. Buitenzorgf w {iS&^i. THE FERN ALLIES. 37 LITERATURE. Baker (John G.). Fern Allies, pp. 7-123. London, 1887. (George Bell & Sons.) Spring (A.). Monographic de la Famille des Lycopod la- ches. In Mimoires de VAcadimie Roy ale de Belgique, XV, i-i 10 (1842); XXIV, 1-358 (1849). ^ir "in 5, :i \ D. THE QUILLWORTS. 99. General Characters. — The quillworts, so named from the appearance of the leaves, are principally incon- spicuous aquatic plants of a grass-like or rush-like aspect (Fig. 27). Some species are always submerged — often in several feet of water; others grow in marshy soil or in the shallow margins of ponds or streams, where they become ap- parently terrestrial in time of low water; while others still are found between high and low water marks, where they will be covered by water at high tide. The leaves are awl-shaped or linear, and are attached to a short fleshy trunk. They vary in number from ten to one hun- dred in each plant, and in length from two to twenty inches in various species. On account of their resemblance to the im- mature forms of rushes and other aquatic vegetation of a higher order, they have been very sparingly collected. Many questions of distribution, habits, and life-history may be studied by even amateur botanists in various sections of the country. Fig. vj.—Iso'ites lacustris L., natural size. (Redrawn from Sprague.) i: 38 OUR NATIVE FERNS AND THEIR ALLIES. In this way valuable additions to science may be contributed by those whose labor misdirected might be wasted. 100. Fructification. — The sporangia of the quillworts, like those of the club-mosses, are sessile in the base of the leaves. The leaf base, sometimes called the sheath, is some- what triangular from the broad insertion, convex behind and concave in front, where there is a large depression known as \)c\& fovea, which contains the sporangium. The margin of the fovea rises in the form of a delicate membrane called the velum, which in many species lies above the sporangium and en- closes it. The sporangia of the outer Figs. aS. aa — ^Two kinds of , ^ . , i • i sporangirin 7. lacustris L., en- leaves contam large spherical ma- larged. (After Sprague.) crospores ; those of the inner con- tain numerous oblong, triangular microspores. The size and marking of the spores form important characters in distin- guishing species. 1 0 1 . Germination. — The microspore after remaining dor- mant through the winter forms a few-celled structure which produces the antherozoids, which are long and slender, and provided with a tuft of cilia at each end. The macrospore produces a prothallium much as in Selaginella (97) ; from this the germ of the mature plant arises after fertilization by the antherozoids. LITERATURE. Baker (J. G.). Fern Allies, pp. 123-134 (1887). Braun (Alexander). On the North American Species of Isoetes and Marsilea, Communicated by Dr. G. Engelmann. In Silliman's Journal, Second Series, in, 52-56 (1847). Campbell (D. H.). Contributions to the life-history of Isoetes. In Annals of Botany, V, 231-258, pi. xv-xvil (1891). Engelmann (George). Isoetes of Northern United States. In Grafs Manual, Fifth Edition (1868). The Species of Isoetes of the Indian Territory. In Bo- tanical Gazette, III, i, 2 (Jan. 1878). The genus Isoetes in North America. In Trans, St, THE FERN ALLIES. 39 Louis Acad. Set., iv, 358-390 (1882). A valuable monograph of this most difficult genus of the fern allies. Underwood (L. M.). The distribution of Isoetes. In Botanical Gazette, Xlll, 89-94 (1888). See also notes in Botanical Gazette, vi, 228. E. THE WATER FERNS. 102. General Characters. — This group includes plants of very diverse characters. Some, like Marsilea, root in mud and produce quadrifoliate leaves. Others, like Pilularia, re- semble the sterile forms of Eleocharis, or other sedges. Others, like AzoUa or Salvinia, float on the surface of water, sending numerous roots into the watet'. Marsilea and Pilularia have a circinate vernation like the ferns. 1 03. Fructification. — The fruit of Marsilea consists of a hollow-stalked receptacle known as the sporocarp, which is oblong or rarely globose, and bears the spo- rangia in sori on the inner walls of its two valves. The spores are of two kinds, as in all rhizocarps. The numerous microspores are con- tained in microsporangia, while the macrospores are solitary in the few macrosporangia. 104. The sporocarp of Pilu- Iraia is globose, containing from two to four cells, which produce microsporangia in the upper portion and macrosporangia below ; the microspores are numerous, while a single macrospore is found in each sporangium. 1 05. In Azolla the sporocarps are of two kinds, borne in the axils of the leaves ; the larger are glo- bose, and contain numerous microspores, which are aggregated in masses ; the smaller are ovoid, and contain a single macro- spore. / - Fig. 30. — Salvinia natan^ Hoflm., natural size. (Re. drawn from Thom^.) t Bit p 'lill;! if. t^ !•■ n 40 OC/^ NATIVE FERNS AND THEIR ALLIES. lOe. Salvinia (Fig. 30), more often seen in cultivation, has the sporocarps borne in clusters on short branches of the floating stem, one or two of each cluster bearing ten or more macrosporangia, each of which contains a single macrospore, the remainder bearing numerous globose microsporangia with numerous microspores. 1 07. Qerminatton. — In Marsilea the antherozoids are produced in a rudimentary prothallium which develops from the microspore and are corkscrew-shaped, consisting of several coils. The prothallium, developed from the apex of the ma- crospore is a hemispherical mass of tissue, and contains a sin- gle archegonium. Much is yet to be learned of the habits and life-history of our native species. LITERATURE. Andrews (W. M.). Apical growth in roots of Marsilia qtiadrtfolia and Equisetum arvense. In Botanical Gazette, xv, 174-177 (1890)- Baker (J. G.). Fern Allies, pp. 134-149 (1887). Braun (Alexander). On the North American species of Isoetes and Marsilia. In Silliman's Journal, Second Series, III. 52-56(1847). Uther Marsilia wndPilularia, InMonatsb.der Konigl. Akad. der Wissenschaft, 1863, 413-436; 1870, 653-753; 1872, 635-679. Campbell (D. H.). The systematic position of the Rhizo- carpeae. In Torrey Bulletin, XV, i^Z-2(iZ (1888). The development of Pilularia globulifera L, In An- nals of Botany, IH. 233-264, pi. XIII-XV (1888). On the Prothallium and Embryo of Marsilia vestita. In Proc, Cal. Acad. Science, III, 183-205, pi. Ill, IV (1892). Some notes on Azolla. In Zoe, ill, 340-343 (1893). The development of the Sporocarp of Pilularia Ameri- cana A. Br. In Torrey Bulletin, XX, 141 -148, pi. CXLVI (1893). Engelmann (George). New Species of Marsilia. In Silli' man's Journal, Second Series, vi (1848). Strasburger (L.). IJeber Azolla. 8vo, 7 plates. Jena (1873). Underwood (L. M.) and Cook (O. F.). Notes on the American Species of Marsilia, In Torrey Bulletin^ xiv, 89-94 (May, 1887). CLASSIFICATION AND NOMENCLATURE, 41 \i 'i CHAPTER VII. CLASSIFICATION AND NOMENCLATURE. The education of a naturalist now consists chiefly in learning how to compare. — AOASSIZ. |873). the 9-94 108. Nomenclature. — The attempts in later years to bring the system of plant nomenclature to a stable basis has resulted in a number of annoying changes in the names of species, and as the present edition contributes something to the matter of change, it may justly be expected to give some reasons for these changes. It is well known that before the time of Lin- naeus, the method of naming plants and animals was a subject of much embarrassment to science, and the lack of a definite system gave rise to much inconvenience and endless confusion. Linnaeus adopted a simple method of naming living organisms, and to him belongs the merit of first extensively and systemat- ically introducing the binomial system of nomenclature which still remains universally in use. Many suppose that this was his own invention, but binomial Latin names for plants were used a hundred years before Linnaeus was born. Comut, for example, in a rare book published in 1635 * illustrates two of our common ferns under the names " Fz/tx baccifera " and " Adiantum Americanum " — probably the first illustrations ever published of American species. Genera existed prior to Linnaeus, and he was not always either wise or just in his selec- tion or use of names for those he recognized. For example, he changed the application of some of the names of that acute botanist, Tournefort (1656-1708), who in 1700 published one of the first accounts of genera,t a much more scientific treatise than anything Linnaeus ever produced. Linnaeus also arbitrarily changed names which his predecessors had used. Mitchell, in * Canadensium plantarum historia. ' t Institutiones rei herbariae. M ; 1.:' t' ' I' •f 1: m $ t 3!' I ■ i 42 OUR NATIVE FERNS AND THEIR ALLIES. 1751, had used the name Attgiopteris for one of our American ferns, but Linnaeus arbitrarily substituted one of his own {Ono- clea) in its place. 1 09. The Linnaean system involved two names for every organism, a generic or group name and a specific or individual name. Generic names are often from the Greek, derived from some characteristic of growth or structure {jCryptogramma, Cheilanthes), or are Latinized in honor of some botanist or patron of botany ( Woodsia), or occasionally from some symbol- ical character {Osmunda). 1 1 0. Specific names are usually Latin or Latinized, and must agree in gender with the generic name, according to the rules of Latin syntax. Specific names frequently indicate some- thing regarding habit or mode of growth {bulbi/era, tomentosa, atropurpurea), or may indicate the locality in which the organism was first discovered {Calif ornica, Ilvensis). A few take their name from their discoverer, in which case the name is Latinized and takes a genitive ending {Boottii, Cooperce), or else an ad- jective form {Goldieana, Wrightiana). 111. The advantage of this binary nomenclature is at once evident when we consider the immense number of ferns alone, to say nothing of the remainder of the vegetable world and the hosts of the animal creation. By this means organisms of complex structure can be definitely characterized with com- paratively few words, and the scientific name once established, is recognized among scientists of all nations and languages. 1 1 2. Among some there is a tendency to regard scientific names with disfavor, on the ground that they are long and dif- ficult. But what shall we say of Geranium, or Gladiolus, or Fuchsia, or Phlox Drummondii, or a hundred others familiar to every lover of flowers ? Are these less difficult than Adiantum, Notholana, Woodsia, or Pellcea Breweri} A little reflection will convince a person of sense that such a criticism is unjust. 1 1 3. A worse tendency is perhaps that which prompts the introduction of " popular names" for ferns : occasionally a name of this kind is highly appropriate, and deserves wide-spread adoption, as in the case of " Christmas-fern " for Polystichum acrostichoides, suggested by Mr. Robinson; the greater part, however, have no merit, and when such monstrosities appear CLASSIFICA TJON AND NOMENCLA TURE, 43 as " Leather-leaf Polypody " for Polypodium ScouUri, " Mr. Goldie's Shield-fern " for Dryopteris Goiiiteana, nomenclature is made cumbrous instead of simple. I 1 4. The Linnsean system, however, did not prove entirely stable. )n the early days when communication among botanists was not easy, the same plant would be described independently by two botanists under different names. Or, in other cases, two botanists would independently establish a certain generic group under different names. For example, Swartz separated a group of plants under the name Botrychium which Linnaeus had included in Osmunda, leaving the latter name for the species we now know under that name. In the same year, and in fact in an article immediately following that of Swartz, Bernhardi separated the same two genera, but left the Botrychium species under the name Osmunda, and took the true Osmunda species out under the name Struthopteris. But errors of this kind were not the worst that existed. Botanists frequently cancelled good names that already existed, and deliberately substituted some of their own. Lamarck in 1797 called one of our Southern fern allies Osmunda biternata ; in 1803 Richard called \l Botrypus lunarioides, recognizing it as belonging to a genus distinct from Osmunda and unaware of the establishment of the genus Botrychium by Swartz. When Swartz in 1806 published the first manual of all known ferns * he properly transferred this species to his own earlier named genus Botrychium, but instead of adopting the oldest specific name he adopted the later one and called this fern Botrychium lunarioides. Willdenow enu- merated the ferns known to him in i8iot and quoted all three of these names, but rebaptized the plant as Botrychium fumarioides. Sprengel, seventeen years later, quoted all these names, includ- ing that of Willdenow, and gave the plant still another name, Botrychium fumarice. It will thus be seen that the period of eruption in nomenclature was in the early part of the century instead of the later, and largely on account of these early irregularities of procedure we have recently been undergoing something of an upheaval of nomenclature. m- ♦ Synopsis Filicum, 1806. t Species Plantarum, vol. 5. Jlr ; J- 44 OUR NATURAL FERNS AND THEIR ALLIES. 1 1 6. Synonymy. — It may also be remarked in this con- nection that different authors have described the same fern under widely different generic and specific names, owing (i) to the different conceptions that have prevailed at different times as to what constituted generic characters, and (2) to ignorance of what others had already written on species, redescribed as new. For example, the delicate Woodsia Ilvensis of Robert Brown was described as Acrostichum Ilvense by Linnaeus, Polypodiutn II- vense by Swartz, Nephrodium rufidulum by Michaux, Aspidium rufiulum by Willdenow, and Woodsia rufidula by Beck. Mai>y other species have been as variously classified. The oppor- tunities for errors of this character are much less now than for- merly, yet redescription is not unknown in our day. 1 1 6. Species. — Goethe tells us that nature knows only in- dividuals, and that species exist only in the school-books. From this extreme there has been every grade of opinion respecting species to the one which regards species as invariable, actual existences, types originally ordained and summoned to existence by the Creator. Linnaeus, for example, defined species in these words : " Species tot sunt diver see, quot diver sas formas ah initio creavit infinitum ens." * Various definitions have been given to species, but none accord with the actual practice of systematists, who seem inclined to make a species what they choose ; and indeed the existence of various connecting forms between many species distinct under normal conditions makes the prac- tical definition of the term almost an impossibility. We may, however, for practical purposes, regard as a species an assem- blage of individuals not differing essentially from each other, and capable of producing like individuals by the ordinary pro- cesses of reproduction. A recent writer defines species as " the present aspect of a line of organic development, destined to become Something else in tha future, as it was something else in the past," — a definition in accord with the now universally accepted biological doctrine respecting the origin of species. Species among ferns are founded chiefly on differences in the cutting of the fronds and their method of venation. * There are as many different species as the Infinite Being created in the beginning. i I :ies. the the CLASSIFICATION AND NOMENCLATURE. 4S 1 1 7. Varieties. — Many forms differing only slightly from the ordinary specific types, and yet capable of transmitting their variations from generation to generation, are regarded as vari- eties. It was the opinion of a prominent botanist, that all so-called varieties among the lower plants " were purely the result of the accident of environment, and never of cross-fertili- zation." Since a species which varies in some minor particular is likely to revert to the ordinary form as soon as the normal conditions of soil, moisture, or environment are restored, there is no scientific foundation for the multiplication of varieties to serve as rubbish in works on systematic botany. A true variety is an incipient species in process of formation ; when it becomes sufficiently distinct to be regarded as a distinct thing with a certain constancy of characters it is more logical to regard it as a distinct species. In cases where species have been more recently separated from each other in their evolutionary progress, some intermediate forms may still persist. If the typical form is clearly marked, these intermediate forms need not invalidate its specific rank. 1 1 8. Genera, — The limits of genera among ferns have given rise to much difference of opinion. The few comprehensive and heterogeneous genera recognized by Linnaeus were soon divided by various authors, and other new genera were based on new dis- coveries resulting from the exploration of newer portions of the world. Adanson, Smith, Roth, Swartz, Bernhardi, Robert Brown, and others added genus after genus, often passing over the work of other post-Linneean authors and often unwittingly or even purposely renamed genera which had already well- established names. Genera were largely based on the varying arrangement of the sporangia on the veins, as well as the char- acter, shape, and position of the indusia. The English of the Hookerian school who have written on ferns have largely depended on these characters and have tended to recognize fewer genera than others of their countrymen or than are usu- ally recognized by Continental botanists. 1 19. Presl (1836) was one of the first to establish genera based on the vascular systems of tlie plants, particularly their methods of venation, and laid the foundation of a more logical classification of ferns. John Smith, whose life had largely been m ■i ri v 46 OUR NATIVE FERNS AND THEIR ALLIES, spent in one of the largest collections of growing ferns, added to Presl's system, characters based on methods of growth. Fee, who gave much attention to the ferns of the West Indies, Mexico, and Brazil, also established numerous genera. To these three writers and to Moore, who followed them, we are indebted for a more liberal and more consistent conception of fern genera. The modern tendency is toward this recognition of a larger number of fern genera, depending on characters drawn from venation and from habit of growth. Such unnatu- ral aggregations of species as have hitherto been grouped to- gether under the name Gymnograme^ because of the fact that the species all had elongated naked sori on the back of the leaf, cannot be tolerated in a system that professes to be founded on natural relationships. Natural genera must contain only species that are more closely related to each other than they are to any other species. 1 20. Families. — Genera are grouped in families according to the characters of the sporangium itself, its method of dehis- cence, and especially its origin from the tissues of the leaf. Eight families of ferns, if we include the eusporangiate Ophto- glossacea and the heterosporous Salviniacece and Marsileacea, are found in our flora. Besides these there are the Marat- tiacece, Gleicheniacece, and Matoniacecs among the ferns of trop- ical regions. Families of plants now have the uniform termina- tion -acea. 121. Orders. — Families are grouped into orders based on still wider characters. The plants with fern-like habit make up the order Fi Lie ales, though it is an open question if the eusporangiate types and heterosporous types ought not to be separated as distinct orders. The rush-like species forming the single family Equisetacecs constitute the order Equisetales, and the club-moss types, isosporous and heterosporous, form the order Lycopodiales. 1 22. Principle of Classification. — The true idea of clas- sification is the grouping together of objects according to essen-' tial and fundamental resemblances. Every system is more or less artificial, yet there is a continual approach toward the true * C/. Hoolcer and Baker, Synopsis Filicium, pp. 376-390. CLASSIFICATION AND NOMENCLATURE. 47 natural system, which is the tiltimatum of scientific classifica- tion. The study of life-histories will continually clear up points of relationship before unknown, and it will not be long before the classification will become fixed and constant. Every real study contributes to this end. 1 23. Changes of Nomenclature. — Exactness of citation is of prime importance, and in later years stability in nomen- clature has been an end constantly sought. Nineteen years ago, when the first edition of this work appeared, the serious study of the higher flora of America was largely confined to a single botanical centre, and that centre followed the practice of Kew, the great royal herbarium of England, in adopting names without particular reference to principles of priority. So long as one centre existed, this system was little questioned. But soon new centres of study of our flora were formed, new workers appeared fresh from fields where the study of plants in life had been added to the study of plants in the herbarium. These workers recognized the fact that in neglecting priority and in following no fixed principles of nomenclature, grave difficulties were con- stantly arising, and confusion followed ; they could not follow a system based on so uncertain and variable a standard as the personal system of nomenclature. European botanists, even Englishmen outside of Kew, recognized the same difficulties. There must be a common starting-point accepted ; there must be some common principles adopted and followed for taking up generic and specific names. During the past ten years the bo- tanical world has quite generally settled down to 1753 as the starting-point of nomenclature,* and most adopt the principle of priority as fundamental ; i.e., each generic group is given its oldest tenable name, and each species bears the original specific name assigned it whether it remains in its original genus or is transferred to some other. Some examples will make this clear if we follow the history of individual species. 1 24. — A simple case that has been involved in a recent change of name is seen in our Eastern lip-fern {Cheilanthes) ; the :' 'iy ir i • * This is purely arbitrary and has been selected for convenience merely. Genera and species were clearly defined before this time. V 48 OUR NATIVE FERNS AND THEIR ALLIES. history of this fern is summed up in the following chronological synonymy : Nepkrodium lanosum Michx., 1803. Chiilanthes vestita Swz., 1806. Cheilanthes lanosa Watt, 1874. Watt rightly discarded the specific name given this plant by Swartz, and adopted the earlier specific one given by Michaux. The full name is then written Cheilanthes lanosa (Michx.) Watt, the parenthesis noting the fact that the specific name was given by Michaux with a generic combination different from the one in which it now stands. The practice of a few botanists has been to give a species the first name it bore under a genus. In this case the plant in ques- tion would bear the specific name vestita so long as it remained in Cheilanthes, but if transferred to the genus Nephrodiutn it would have to bear the specific name lanosum,sind if transferred to some other genus might bear still a third. The absurdity of such a practice is clearly apparent.* 1 25. — A case slightly more complicated is seen in the hart's- tongue — the lingua cervina of the pre-Linnaean botanists. Its chronological synonymy is as follows : Asplenium Scolopendrium L., 1753. Scolopendrium vulg-arej. E. Smith, 1793. Phyllitis Scolopendrium Newman, 1854. Scolopendrium Scolopendrium Karsten, 1883. The last name is perfectly legitimate although a duplication, and so long as the plant remained in the genus Scolopendrium that was its appropriate name. But the genus Scolopendrium was founded by Adanson in 1763,! and the generic name Phyl- litis founded in pre-Linnaean times on the same plant was used since Linnaeus at least as early as 1757. This being true, New- man's combination is the correct one to follow, and the full cita- tion would be Phyllitis Scolopendrium (L.) Newm. I' \ * For each plant or group there can be only one valid name, and that is always the most ancient if it is tenable. — A. Gray. t Not by J. E. Smith, 1793, as usually supposed. CLASSIFICATION AND NOMENCLATURE. 49 I 26.— Still more complicated is the ostrich-fern. Its syno- nymy is as follows : Osmunda struthiopteris L., 1753. Onoclea struthiopteris Hoflfm., 1795. Osmunda nodulosa Michx., 1803. Struthiopteris Germanica Willd.,* 1809. , Struthiopteris Pennsylvanica Willd., 1810. Matieuccia struthiopteris Todaro, 1866. Struthiopteris Germanica var. Pennsylvanica Lawson, 1889. Now this case involves several independent problems that are not mere " battles with synonyms " : (i) Is the American spe- cies the same as the European ? (2) Are we to take a superficial resemblance like the rolling of the sporophyll into a necklace- shaped structure as a ba^is for comparison, and unite a species with leaves growing in crowns from an upright rootstock and having free veins, in the same genus {Onoclea^ with a plant that has horizontal creeping stems, scattered leaves, and copiously anastomosing veins? These are problems on which human judgment will disagree as it has disagreed in the past. In regard to the latter question the practice of the Kew botanists followed too implicitly by us Americans was adopted in previous editions not without many misgivings. We believe that the two ferns form two as valid generic groups as exist ; that there is nothing in common between them to indicate community of origin or even an3rthing but the most distant relationship. They are therefore treated in this edition as two genera. In regard to the question of the identity of the European and American plants, we will say that, having been familiar with our American species from childhood, and having studied the European form in its native soil, we are forced to the conviction that there is but one species on the two continents. If this be the case, whatever the generic name may be, the specific name of our species must be Struthiopteris, the Linnsean specific name for the plant. If we adopt the view that the American plant is distinct from the European, our plant would then bear the specific name * Applied to the American plant by various authors, but limited bjr Will- denow to the European plant. i >» II I If % t 't V 50 OUX NATIVE FERNS AND THEIR ALLIES. 't If- (■;: w m i nodulosa, the earliest name; Willdenow in establishing the genus Struthiopterts in 1809 incorrectly stating as a fact that the American plant was " eine noch nicht beschriebene aus Penn- sylvanien," and not assigning it a name until 18 10. 1 27. — Having thus fixed the specific name, what of the generic ? The name Struthiopterts cannot be used for this plant, for wlien Willdenow assigned it to this use it had been used already twice before. In 1760 Scopoli used it for a genus of which Osmunda spicant was the type ; Bemhardi used it again in 1799* to include the species of the genus we now know as Osmunda to separate them from the ill-assorted aggregate which Linnaeus had brought together under this name. Struthiopterts must then stand for a genus which hitherto has commonly been called Lomaria, and our ostrich-fern must look farther for a name. Matteuccia, proposed by a Sicilian botanist in 1 866, appears to be the first tenable generic name, and is here used in that sense. It will thus be seen that the question of the proper use of botanical names is by no means a simple one. The botanical literature of the world must be ransacked before stability can be reached. An obscure local publication in the Italian language on the plants of Sicily in this case furnishes the generic name for a plant which grows in the northeastern United States ! 1 28. After specific stability is settled comes the equally interesting problem of generic stability which is still more diffi- cult. This, however, involves principles that have never been thoroughly discussed, and this subject will not be considered here,t except to give a single illustration. In 1799 Bernhardi established a genus of plants under the name Gymnopteris based on a single West Indian species which Linnaeus first described as Pteris ruffa, but afterwards referred * Bernhardi's orthography was Struthopteris, a fact that has led an over- ardent nomenclaturist to abolish the genus Osmunda. t Those interested in this phase of the nomenclature question will find a paper by the writer on " The Genera of Ferns established prior to 1832," in the' Memoirs of the Torrey Botanical Club, vi, 247-283, 1899. On the general question of botanical nomenclature one of the best discussions of the subject will be found in the Bulletin of the Torrey Botanical Club, xxii, 308-329, Mm. V CLASSIFICATION AND NOMENCLATURE. 51 to the genus Acrostichum. Ten years later Desvaux established the genus Gymnogramma, based on this same West-Indian species, and eleven others with a similar method of forming their sori. Now under any rational system the name Gym- nopteris must stand for that group of species which includes the Pterts ruffa of Linnaeus, and it is equally true that it could not be used legitimately for any other group of plants. To illustrate how wide of the mark certain modem usage is, it is only necessary to cite a recent revision of fern names * in which Gymnopteris is used for a wholly different group of ferns from that which contains the plant Linnaeus called Pterts ruffa, and for the group to which Pterts ruffa actually belongs a name is selected that was not established until 1844, namely, Lepto- gramme Link, thus passing over two earlier names which had priority ! \\ LITERATURE. The references to original writings would include all tlie botanists who have named or classified ferns since the time of Linnaeus (1707-1778). Among the more prominent of these wc may mention Swartz (1760-1818), Willdenow (1765-1812), Presl (1791-1849), Mettenius (1823-1866), Hooker (1785-1865), Fee (1789-1874), Milde (1824-1871), Al. Braun (1805-1875), and J. G. Baker ( - )• The following work gives a good review of the various systems : Smith (John). Historia Filicum. London, 1875. (Mac- millan & Co.) The American literature bearing on the subject is as follows : Beck (Lewis C). Synoptical tables of the Ferns .ind Mosses of the United States. In Silliman s Journal, iv (1829). Davenport (George E.). Aspidium spinulosum (Swz.) and Its varieties. In American Naturalist, xii, 707-717 (1878). New species of Ferns. In Bulletin of the Torrey Bot. Club, VI, 190, 191 (1877) ; VII, 50, 51 (1880) ; viii, 61, 62 (1881); X, 61, 62 (1883). * Die naturlichen Pflanzenfamilien. (Engler-Prantl.) ^: % I- i'>; 1 Mm M h. 52 OUR NATIVE FERNS AND THEIR ALLIES, Fern notes. In Bulletin of the Torrey Bot. Club, VII, 85, 86 (1880) ; VIII. 88. 89 (1881) ; IX. 20-23. 68. 69. 99-101 (1882) ; X. 4-7 (1883); XII. 21-24 (1885); XIII. 81. 82, 129-135 (1886); XV, 225-229 (1888). Eaton (Daniel C.)< Ferns of the Mexican Boundary. In Mexican Boundary Survey (1857). Ferns of the Southern States. In Chapman : Flora of the Southern States (i860). Ferns of the Northern United States. In Gray : Man- ual of Botany, 6th edition {\Z^). Notes on some of the plants in the herbaria of Linne and Michaux. In Canadian Naturalist (1870). New and little known Ferns of the United States. In Bulletin of the Torrey Bot. Club, IV, 11. 12, 18, 19 (1873) ; vi. 33 (1875), 71.72 (1876), 263-265 (1878), 306. 307,360,361 (1879); VII, 62-64 (1880) ; VIII, 4, 5, 99, 100 (1881) ; IX, 49, 50 (1882) ; X, 26-29, loi, 102 (1883). Ferns of North America. Illustrated with colored plates by J. H. Emerton and C. E. Faxon. Ferns of the Southwest. In Wheeler : Report of the U. S. Geog. and Geol. Surveys west of the 100th meridian, vi (1877). Vascular Acrogens of California. In Watson : Botany of Calif ornia, 11 (1880). Gray (Asa). On the discovery of two species of Tricho- manes in the State of Alabama. In Silliman's Journal, 2d ser., XV (1853). KuNZE (G.). Notes on some Ferns of the United States. In Silliman' s Journal, 2d sen, VI, 80-89(1848). Underwood (L. M.). American Ferns, I, II. Bull. Torrey Club, XXV, 521-541 (1898) ; XXVI, 205-216 (1899). The literature relating to exotic species is very extensive. Some of the more important works are the following : Baker (J. G.). A summary of the new Ferns which have been discovered or described since 1874. (1892.) Fee (F. L. A.). Memoires sur la Famille des Fougeres. 4to. (i 844-1 873.) 289 plates. V CLASSIFICA TION AND NOMENCLA TURE. 53 Hooker (W. J.). Genera Filicum. 4to. (1842.) 120 col- ored plates. Species Filicum. 5 vols. 8vo. (i 846-1 864). 304 col- ored plates. Hooker (W. J.) and Baker (J. G.). Synopsis Filicum. 2d ed., 8vo. (1874.) Contains descriptions of all the ferns of the world recognized at Kew to the date of publication. Hooker (W. J.) and Greville (R. K.). Icones Filicum. 2 vols, folio. (1831.) 240 colored plates. Mettenius (G.). Filices Hort Botanici Lipsiensis. 4to. (1856.) Ueber einige Farngattungen. Five parts. 4to. (1857- 1859.) Luerssen (C). Die Farnpflanzen. In Rabenhort: KryP^ togameit'Flora Deutschlands. 8vo. Besides numerous scattered papers by all of the above-men- tioned writers, together with Moore, Kuhn, A. Braun, Prantl, Milde, Christ, Jenman, Fournier, and many others. \ m 1 %:\ > :, : 54 OUR NATIVE FERNS AND THEIR ALLIES. CHAPTER VIII. THE FERN'S PLACE IN NATURE. 129. The popular conception as to what constitutes a plant needs to be considerably enlarged and otherwise modified, for as soon as we commence to look about us after our eyes have been really opened, we find a vast array of forms varying in size and complexity of structure from the simple cells of the yeast- plant that we use in bread-making to the highly organized tree of the forest, and including such diverse forms of growth as the green scums that accumulate on ponds in summer, the gray lichens covering rocks and trees, the puff-balls and mushrooms that seemingly develop in a single night, the mosses, ferns and flowers in all their variety and beauty. Where in all this array of plants do our ferns stand, and what relations do they susb^'n to other plants ? In answering this question we will have to give some account of the various groups of plants, pointing out their structural peculiarities and noting here and there in their appropriate place in the system such forms as are likely to be popularly recognized. 1 30. Aside from the plants producing flowers, the ferns and the mosses,'" all of which are widely known and generally * It should be noted that even this name is often misapplied. The lichens, which are :n no way related to the true mosses, are sometimes popularly called "gray mosses." In "Evangeline" where Longfellow speaks of the trees " bearded with moss" he evidently alludes to the lichen, Usnea barbata ; the " hanging moss" of the Pacific coast is also a lichen, Ramalina reticulata, which has a much more appropriate name in "lace-lichen." The "hang- ing moss " of the Gulf States, on the contrary, is a flowering plant whose near- est allies are in the pineapple family. Another flowering plant, Euphorbia cyparissiaSy is often called " graveyard moss " in the Northern States. This loose and confusing use of language is to be deplored, and those who know better ghould assist in relegating these incorrect usages to a merited oblivion. / THE FERN'S PLACE IN NATURE. be 55 recognized, we find two types of plants of lower grade which stand out prominently to even the unpracticed eye. Of these the first are mostly green,* and though variously known and named may be called collectively a^e ISOETUtOBC MerPODIAeE>E 8PHAQNA0EX PHYCOMVCETES MasocaooQiOEX Pmvisiohal Pbdigrbb of the Lbading Groups of Plants. 60 OUR NATIVE FERNS AND THEIR ALLIES, CHAPTER IX. DISTRIBUTION IN TIME AND SPACE. 1 37. Geoerraphic Distribution. — Ferns are found in all parts of the world. The number of described species is not certainly known, and the uncertainty is largely increased for the reason that our best systematists do not agree as to what constitutes a species. Baker places the estimate at about 3000 species. Added to these are 565 fern allies as recognized by the same author. The full number is probably much greater than this very conservative estimate. From what has been said respecting the climatic conditions of fern growth we would naturally expect to find them most abundant in countries where warmth and moisture predomi- nate. These conditions seem most completely met on tropical islands or in tropical continental areas with insular climates. The little island of Mauritius, having an area of 676 square miles, or less than one third the area of Delaware, has 235 na- tive species, while Java, little larger than New York, has 460. Brazil furnishes 387, and the Isthmus of Panama 117. Com- paring these with colder climates, we find 67 in all Europe, and only 26 grow within the borders of the arctic zone. " Our Native Ferns," as described later in this volume, in- cluding those species that are classed in the order Filices, number 170 species. Adding to these the 22 species of the order OPHiOGLOSSACEiE, which have frequently been enumer- ated with the ferns, we have a total of 192 species. The remain- ing fern allies number 87, making a grand total of 279. 138. Divisions of our Flora.— It has been found con- venient to divide the surface of the earth into faunas and floras, limited by the natural distribution of the various species of DISTRIBUTION IN TIME AND SPACE. 6i animals and plants. These limits are by no means sharply defined, for wherever the limit is made some species will pass beyond it ; yet the majority found on one side are different from the majority of those on the other. North America (ex- cluding Mexico) forms the Nearctic realm or fauna {Regnum Nearcticwn), and the same boundaries may be used in the limi- tation of our fern flora, although some species from tropical regions invade our borders in Florida, Texas, and Arizona. Leaving out of question the species that are widely distributed over the greater part of our country, many of which are cos- mopolitan species, we may divide the Nearctic realm into five provinces, each of which possesses many species peculiar to itself. 139. The provinces* are as follows : I. Boreal : inhabiting (with a few exceptions) the northern portion of the United States, extending through Canada and British America, some species even reaching Labrador, Green- land, and Alaska, and nearly all represented also in the north- ern portions of the Old World. II. Medial : extending throughout the mountain and hilly region of the States east of the Mississippi, westward to the mountains, and northward into Canada, and in a few instances also inhabiting the Old World. III. Occidental : extending along the western border of the continent from British Columbia to California, in a few cases appearing also in the Rocky Mountain region. IV. SoNORAN : inhabiting the central mountain regions of Western Texas, Arizona, and Colorado, many of the species ex- tending thence into Mexico, and some even to South America. V. Austral : inhabiting the border of the Gulf of Mexico, many of the species extending into the West Indies and Tropi- cal America. 140. Geolofiric Distribution. — It is well known that the plants and animals now existing on the earth are not the same in kind as those of former ages. Geologists have carefully studied the stony heart of nature, and have drawn therefrom * This division is a slight modification of one proposed by John H. Red- field in 1875. C/. Bulletin Torrey Botanical Club^ VI, 1-7. V 62 OUR NATIVE FERNS AND THEIR ALLIES, the story of the development of land and sea, and the succes- sive populations that from time to time have held possession of our globe. Plants furnishing the natural food for animals must have preceded animal life, yet in the earliest geologic ages the remains of animals are far more numerous. The abundance of the deposits of graphite and iron-ore in the earli- est or Archaean rocks indicates the existence of extensive plant growth, but the remains are so transformed as to mak^ it im- possible to determine the character of this primeval vegetation. 141. In the succeeding Silurian age the fossil remains in- dicate the existence of algae or sea-weeds in abundance, and a single small species of ground pine attests the existence of some of the higher Cryptogamia ; no ferns, however, have been found in America older than the Devonian. Over fifty species of Devonian ferns have been described from the American rocks chiefly, by Principal J. W. Dawson of Montreal. 142. It is in the coal measures, however, that ferns and other Cryptogamia are found in the greatest abundance and profusion. Their delicate foliage is impressed on the various rock strata above the beds of coal, and so perfectly are they preserved that not only the methods of fructification but even the microscopic spores have been detected ! In the coal meas- ures of the United States and Canada (counting from the base of the Catskill), 381 species of ferns have been described, chiefly by Prof. Leo Lesquereux. The most abundant Ameri- can genera are Neuropteris 45 species, Pecopteris 50 species, Sphenopteris 31 species, Pseudopecopteris 25 species, and Rha- cophyllum 24 species. The frontispiece gives an ideal representation of the vegeta- tion of the Carboniferous age. The luxuriant tree-ferns, the Lepidodendrids, ancient representatives of the diminutive club- mosses or ground-pines, the Calamites, allies of the modern scouring-rushes, and other forms no less wonderful, are seen in their profusion. 143. In the later geologic ages, Mesozoic and Tertiary, ferns are found preserved in the rocks, with the leaves of many trees and shrubs of existing genera. The indications are that DISTRIBUTION l!f TIME AND SfACB. 63 terns formed a far smaller part of the vegetation of these later ages than in the preceding Carboniferous, and even approxi- mated to that of the present. Six Cretaceous and twenty-four Tertiary species have been catalogued,"" including species in the existing genera Lygodtum, Pteris, lVoodwardia,Dryopt£rts, Gymnogramme, etc., as well as some related to genera abundant in earlier formations. No living species is found fossil, unless Dr. Newberry's variety of Onoclea sensibilis becomes estab- lished.! In the course of geologic history, however, we can trace a gradual approximation to the modern types from the generalized forms of Devonian and Carboniferous times. 1 4.4. Fern Allies. — Ophioglossum dates back to the Ter- tiary period with one species. The order EQUiSETACEiE have existed since the coal period and the genus Equisetum since the Triassic. The order CALAMARiACEiE, which combined charac- ters of modern Equiseta and Conifers, came into existence in the Devonian, but became extinct before the close of the Permian. Illustrations of Calamites can be seen at the left-hand corner of the frontispiece, also under the tree-fern in the centre. The club-mosses proper have been in existence since the Devonian, and the genus Lycopodium since the Carboniferous. Selaginella has never been found fossil, but its near relatives belonging to the extinct orders LEPiDODENDRACEiE and SiGiLLARiACEiE were very abundant in the Palaeozoic era, particularly during the Carboniferous, where they formed the largest part of the forest vegetation, reaching in some instances a height of sev- enty to one hundred feet. The former possessed characters connecting modern club-mosses with Conifers, while the latter ■M%\ * Tenth report, Hayden Geological Survey of the Territories. Washing- ton, 1878. t Prof. Lesquereux writes me: "Though analogous by the nervation, I doubt the identity on account of the coriaceous character of that fossil fern, which I have not seen in any variety of O. sensibilis now living. " Principal Dawson, however, writes : " The Onoclea sensibilis of tht T.aramie is truly that species, and I have found with it in our Manitoba formations another modern fern, Davallia tenuifolia,^^ 64 OUR NA TIVE FERNS AND THEIR ALLIES, seem to connect the club-mosses with the Cycads. Restor^i tions of Lepidodendron may be seen on the left-hand side of the frontispiece, and of StgiUaria on the right. Isoites dates back to the Miocene (Tertiary) and MarsiUa and Pilularia to the same pe^ ;od. I ( the ack the OUR NATIVE PTERIDOPHYTES. PTERIDOPHYTA Cohn. Plants containing vascular tissue and manifesting two distinct phases in their life-history: (i) An asexual phase {sporophyte) differentiated into stem and leaves, producing spores and de- veloping vascular tissue in bundles throughout their stems and leaves ; and (2) A sexual phase {gametophyte) developed from the germination of the spore in the form of a cellular thallus {jprothallium) on which the sexual organs — antheridia and arche- gonia — are produced ; from the egg of the archegonium fertilized by the antherozoids from the antheridia arises the asexual phase from which the characters used in classification are largely drawn. Besides several groups that have become extinct the Pteridophytes are represented by three orders : I. Filicales, containing the ferns and waterworts; II. Equisetales (see p. 126) including the horsetails and scouring rushes; and III. Lycopodiales (see p. 130), containing the ground-pines and quillworts. synopsis of the order filicales. (families.) X. Spores uniform, of one aort a Spores of two sorts (minute microspores and large macrospores) ... 6 a. Sporang;ia rising from the tissues beneath the epidermis (eusporangiate), borne in spikes or panicles ; vernation straight or inclined. Family i. Ophioglossace^, p. 66 Sporangia rising from the epidermal cells (leptosporangiate), borne on the back or margin of a leaf (frond) or rarely in panicles ; vernation circi- nate 3 3. Sporangia sessile, with a complete ring borne on a thread-like receptacle from a cup-like involucre ; texture filmy. Family a. HVMENOPHYLLACEiG, p. 74 Sporangia borne on the back or margin of the leaf or in spikes or panicles. 4 4. Plants terrestrial 5 Plants aquatic ; sporangia sessile, scattered, in a specially folded leaf. Family 5. CERATOPTERIDACEiE, p. 78 5. Sporangia sessile, ovate, with an apical ring, opening longitudinally, mostly in panicles or solitary under a scale. . Family 3. Schizaace/c, p. 75 fj'J i in V ^^ OUR NATIVE FERNS AND THEIR ALLIES. I Sporangia with a rudimentary ring, opening longitudinally, in panicles. Family 4. OsMUNDACEiE, p. 77 Sporangia stalked, with a complete ring, opening transversely. Family 6. PoLYPODiACEiE, p. 78 6. Rooting in mud ; leaves filiform or quadrifoliate. Family 7. MARSlLEACEiC, p. 133 Floating ; leaves spongy Family 8. SALViNiACEiE, p. 135 Family I. OPHIOGLOSSACE>iE Lindl. Plant-body consisting of stem and leaf, usually from a fleshy sometimes bulbous root, straight or inclined in vernation. Eusporangiate, the sporangia formed of the interior tissues, variously clustered on sporophylls in the form of spikes or pani- cles, destitute of a ring, opening by a transverse slit into two valves and discharging their copious sulphur-yellow spores. Prothallium (so far as known) subterranean, not green, monoe- cious. The family contains about six genera, three of which are represented in America. Our genera may be distinguished as follows : 1. Sporangia in spikes cohering in two ranks 2 Sporangia free, in compound spikes or panicles ; leaf mostly divided III. Botrychium. 2. Spike solitary; leaf simple, entire, attached to the middle of the common stalk or below ; terrestrial. I. Ophioglossum. Spikes several, pendent from near the base of a palmately divided leaf ; epiph3rtic 11. Cheiroglossa. I. OPHIOGLOSSUM L. Addbr-tongue. Sporangia large, coriaceous, connate, coherent in two ranks on the edges of a simple spike. Leaf simple, attached at the middle of the main stalk or below, entire ; veins anastomosing. Spores copious, sulphur-yellow. Terrestrial. Name from Gr. oifft'er secondary segments from a narrowed base, ovate, o*'«nport.) acute, serrate, the upper spreading, quickly decreasing, finally elliptical, acute ; fertile segment bi — tripinnate, panicled. Apex of sterile segment bent over inside of the nearly erect fertile one in vernation ; divisions of the sterile segment arranged on an angle. Unalaska. 1 1 Vernation wholly inclined^ recurved in the sporophyll; Itaf triangular t sessile. OPHIOGLOSSA CEiE, 71 Fig. 33.— Veraation of B. simplex E. Hitchcock. Plant 2'— (Afte^DavelfJ^.) "''' B. 6. B. lanoeolatum (S. G. Gmel.) Angs. Plant 3'— 12' high, •omewhat fleshy. Leaf closely sessile near the summit of the stem, I'— 2' wide, 3-lobed or broadly trian- gular and 2-pinnatifid, the ultimate segments lanceolate, acute, oblique, entire or dentate ; midvein continuous, with forking veinlets ; sporophyll slightly overtopping the leaf, short-stalked, 2— 3-pinnate, recurved its whole length with the shorter leaf reclined upon it in vernation. Nova Scotia to Alaska, south to New Jersey, Ohio, Colorado, and Washington. Also in Europe and Asia. June-July. ♦ ♦ Ltaf rising from near the rootstock. t Vernation wholly straight ; bud smooth ; leaf entire or 2—6-lobed. 7. 5' high, slender, very variable. Leaf ovate, obovate, or oblong, entire, lobed, or pinnately parted, borne near the base of the stem ; sporophyll a simple or slightly compound spike, sometimes reduced to [only a few spo- ranges ; spores large for the genus, minutely tuberculate; apex of leaf and sporophyll Pig. 34.— Vernation of erect in vernation. Nova Scotia to Maryland, B.timpltx Hitchcock. >. .. -iir j /- i-r • /■^^ (After Davenport.) west to Wyommg and California (?). tt Vernation inclined; leaf ample, ternately compound. X spores maturing in early spring ; leaf sessile or nearly so, 8. B.'blternatum (Lam.) Underw. — Plant 3'— 6' high, bear- ing a nearly sessile, broadly triangular ternately compound leaf, 3'— 4' wide, 2' high; middle division slightly larger than the lateral ones and like them nearly bipinnate ; ultimate divisions rather lunate, usually not exceeding 2" — 3" in width, the outer margin crenulate, the lateral margins decurrent into the short branches of the rachis; sporophyll on a rather stout stalk, bipinnate, with a rather broad rachis ; bud smooth or slightly hairy, the segments nearly erect. {B. lunarioides Swz., B. fumarioides Willd., B. fumarice Spreng., Osmunda biternata Lam.) South Carolina to Louisiana, apparently not common. ^2 OUR NATIVE FERNS AND THEIR ALLIES, J, I ■III! X \ spores maturing in autumn ; leaf long~stalked. \ Ultimate leaf-s^ments laciniate, narrow, ^' or less wide. 9. B. disseotum Sprang. — Plant 6' — 15' high, with slender fleshy stems. Leaf long-stalked from near the base of the stem, with broadly deltoid basal divisions, decompound ; sec- ondary pinnae lanceolate from a broader base, pinnate with laciniate and deeply cut pinnules, the ultimate divisions diver- gent, often 2-toothed at their apices, usually less than i mm. wide ; sporophyll long-stalked, 2 — 3-pinnate ; bud pilose, en- closed in the base of the stem, both portions bent in vernation. New England (where a more compact variety is more common) to Virginia and inland to Kentucky and Indiana. I II Leaf-segments small, rounded, or obliquely ovate, i\" — 3" wide ; plant small {leaf i' — 2' wide). 10. B. matrlcarlae (Schrank) Spreng. Plant 4' — 6' high, with slender flesh/ stems; leaf moderately short-stalked, ternate, small, i' — 2 wide and high, the three divisions similar, bipin- natifid or bipinnate; ultimate segments small, i^' — 3" wide, rounded or somewhat obliquely ovate, the margins undulate or crenate; sporophylls rather long-stalked for the size of the plant, 2 — 3-pinnate with large sporanges ; bud pilose. Northern New England and New York and northward. II III Leaf -segments obliquely ovate, large, 5"- 10" long. {Eastern.) 11. B. obllquum Muhl. Plant robust, 7' — 20' high; leaf rising from near the base on a stalk 3' — 4' long or more, ternate with the three divisions nearly equal, bipinnate or somewhat tripin- natifid in larger forms, the ultimate segments obliquely ovate or oblong-lanceolate, the terminal one of each division elongate, all 5" — 10" long, 2i" — 4" wide, the margins cre- nate or serrate; sporophyll long-stalked tri- quadripinnate ; bud densely pilose, both por- tions bent in vernation. {B. ternatum in part, of former editions, not of Swz., which was Thunberg's Osmunda ternata from Japan.) New Brunswick to Florida and Mexico and westward to Minnesota. Var. intermedium (D. C. Eaton) Underw. —Vernation 'iquum Muhl. (After Davenport.) plant larger, the leaf on a shorter stalk i'— 2' rv ■, i i n I( h ti OPHIOGLOSSA CEM. 73 -4' ions pin- ents the all cre- tri- >or- in lich Ian.) land ;rw. -2' long, the leaf sometimes reaching 6' each way ; lateral divisions smaller than the terminal; ultimate segments similar to the type, but mostly shorter. Northern New York and New England. The limits of this variety are not fully understood. H II I li Ltaf-segments ovate or roundish ; plant large and stout. (Western). A. Leaf -stalk short {i' or less) ; segments crowded. 12. B. Coulter! Underw. A stout fleshy plant growing in geyser formations. Roots numerous, fleshy, stout : stem very short, i' or less long, very stout, 7" — 10" in diameter, swollen with the contained bud of the succeeding season, soon dividing to form the sporophyli and leaf ; petiole very short, \' or less long, stout, sulcata in drying ; sterile lamina about 6' wide, the x,entral portion nearly 4' long, this and the lateral ones tripin- nate, or quadripinnatifid ; segments obliquely ovate, 5" or more long, 2" or more wide, thick, fleshy, tht margin entire -^ slightly repand; veins few, scarcely perceptible; sporophyli about ^' long ; panicle quadripinnate below, the pinnae crowded, gradu- ally simpler above ; sporangia very numerous, bright yellow ; spores copious, pale yellow. In geyser basins, Yellowstone National Park ; Montana. B. Leaf "Stalk longer (4' — 6') or more ; segments more scattered. 13. B. occidentale Underw. Roots fibrous, fleshy; stem short, i' — 2' long, 2" or more in diameter; leaf-stalk 4' — 5' long, rather slender; leaf very large, 7' — 8' broad, 5' — 6' high, the lateral divisions bipinnate with about five pairs of mostly oppo- site pinnae; terminal division tripinnatifid, gradually simpler above; ultimate segments nearly oval, mostly narrow (under 3" wide), the margins finely and irregularly crenulate ; texture fleshy, the veins indistinct ; sporophyils 16' long, including the panicle, which ranges from 4' — 6', tripinnate almost throughout its entire length ; bud densely covered with white silky hairs. Washington, Oregon, and British Columbia. 14. B. sllalfolium Presl. Plant robust, 15' — 2° high; com- mon stem rather short, i' — 2' long ; leaf-stalk ; stout 3' — 6' long ; leaf very large, 8' — 10' or more wide, 5' — 8' high; formed of a larger central division and two lateral ones; divisions nearly tripinnate; ultimate segments ovate, the lowest outer series \i YM \ '^ lii 74 OUR NATIVE FERNS AND THEIR ALLIES, often trilobed ; sporophyll long-stalked, much overtopping the leaf, the panicle ample, 5' — 8' long. California to British Columbia. §2. OsMUNDOPTERis Milde. Bud pilose, enclosed in a smooth upright cavity at one side of the lower part of the stalk. 15. B. Vlrerlnlanunn (L.) Swz. (Rattlesnake-FERN.) Plant from a few inches to two feet high ; sterile segment ses- sile above the middle of the stalk, broadly triangular, thinly herbaceous, ternate; the short-stalked primary divisions once to twice pinnate, then once or twice pinnatifid ; lobes oblong, cut-toothed toward the apex ; fertile segment long-stalked, bi — tripinnate. Bud pilose, enclosed in a smooth upright cavity at one side of the lower part of the stalk ; fertile segment recurved its whole length, the longer sterile segment reclined upon it Reduced forms are B. gracile Pursh. (BotryPus Virginicus Michx., Osmunda Virginiana L.) New Brunswick to Florida, and westward to Arizona and the Pacific Coast. Family 2. HYMENOPHYLLACEiC Endl. Plant body consisting of a creeping stem bearing scattered leaves of a filmy consistency, usually translucent. Sporangia provided with a ring, sessile on a thread-like receptacle which is surrounded at base by a cup-shaped or two-valved involucre. The family contains several genera, mostly of tropical regions, only one of which is represented in our flora. I. TRICHOMANES Sm. Filmy-fkrn. Sori marginal, terminating a vein, more or less sunken in the frond. Sporangia sessile on the lower part of a cylindrical, filiform, often elongated receptacle. Indusia tubular or funnel- shaped, entire or two-lipped at the mouth. Fronds delicate, pellucid. Name from Gr. rpixofiavei, the name of some fern, from Tftix, hair, and fxaivofxai, producing frenzy, alluding to some supposed property. A tropical and temperate genus con- taining nearly 100 species. , § EUTRICHOMANES. I. T. Petersll Gray. Stipes i" — 2" long; fronds 3"— 10" long, i" — 2" broad, oblong-lanceolate or obovate, entire or vari- SCHIZ^ACE^. n :h ously pinuatifid, the younger ones with a few black hairs along the margins ; indusium solitary, terminal, funnel-shaped, the mouth expanded and slightly two-lipped, the receptacle in- cluded. Winston County, Alabama {Peters). 2. T. radlcans Swz. Rootstock wiry, tomentose ; stipes ascending, i' — 3' long, naked or nearly so, usually broadly winged ; fronds 2' — 8' long, i' — i^' wide, lanceolate or ovate- lanceolate, bipinnatifid ; pinnae ovate, obtuse, the upper side of the base parallel and appressed to the winged rachis, the lower side cuneate ; divisions toothed or divided into linear lobes ; indusia terminal on short lobes, tubular or funnel-shaped, the mouth slightly two-lipped; receptacle exserted little or very much. (7". specwsum Willd.) Alabama, Tennessee, Kentucky. Family 3. SCHXZ/EJKCE/E Presl. Plant-body consisting of a short or creeping stem (rootstock) bearing clustered or scattered leaves. Sporangia ovat ) or pyri- form, provided with an apical ring, bursting longitudinally at maturity. The family contains about ten genera, three of which are represented in our region. Our genera may be distinguished as follov»rs : 1. Leaves twining; leaflets in pairs, palmate . . I. Lygodium. Leaves erect or merely curled 2 2. Sporophylls distinct from the grass-like leaves. in. SCHIZiEA. Sporophylls borne on the elongate lower pinnae of an ordi- nary leaf IL Ornithopteris. L LYGODIUM Swz. Climbing-fkrn. Sporangia ovoid, solitary or occasionally in pairs, in the axils of large imbricated scale-like indusia, which are fixed by their broad bases to short oblique veinlets. Fronds scandent, twining, bearing stalked and variously lobed divisions in pairs. Veins mostly free. Name from Gr. Xvyoodrji, flexible, alluding to the scandent stems. Includes 25 species. § EULYGODIUM. I. L. palmatum (Benih.) Swz. Stipes slender, twin- ing; fronds 1° — 3° long, the short alternate branches or peti- ■J i 76 Or/X NATIVE FERNS AND THEIR ALLIES. oles 2-forked, each fork bearing a round-cordate palmately 4 — 7- lobed pinnule ; fertile pinnules above, contracted, several times forked, forming a terminal panicle ; surfaces naked ; texture thinly herbaceous. (Hydroglossuvi palmatum Willd.) Mas- sachusetts and New York to Kentucky and Florida. II. ORNITHOPTERIS Bernh. Sporangia ovate, sessile, placed in two aOws on the back of the very narrow branchlets of the two long-stalked, panicled, lower branches of a pinnately divided frond, the fertile branches in a few species entirely distinct from the sterile frond. Veins free. Name from Gr. u/jviS, bird, and itrepii, fern. A genus mostly of tropical America containing about 35 species. 1. O. adiantlfolla (L.) Bernh. Rootstock creeping ; stipes li* long, firm, naked ; fronds sparingly pubescent, the two lower branches elongate, pinnately decompound, fertile ; sterile por- tion deltoid-ovate, bi — tripinnate ; ultimate segments obovate or cuneate, entire or lobed, striate above with numerous flabellate veins. {Anemia adianttfolia Swz.) Florida. 2. O. Mexicana (Kl.) Underw. Rootstock creeping, cov- ered with narrow blackish chaff; stipes slender, scattered, 6'— 12' long; the two lower branches of the frond fertile, long-stalked, glandular bipinnate with densely clustered fructification; the rest of the frond like the sterile ones, deltoid-ovate, simply pin- nate ; pini.ae about six pairs and a rather large terminal one, short-stalked, ovate-lanceolate, subcoriaceous, smooth and some- what glossy ; midrib distinct, veins free, oblique, parallel, ciosely placed. {Anemia Mexicana Klotzsch.) Western Texas. III. SCHIZiCA Sm. Curly-grass. Sporangia large, ovoid, striate rayed at the apex, naked, ver- tically sessile in a double row along the single vein of the nar- row divisions of the fertile appendages to the slender and sim- ply linear, fan-shaped, or dichotomously many-cleft fronds. Name from Gr. a-x'^eiv, to split, alluding to the forked sterile fronds of foreign species. Includes 16 species. § EUSCHIZyEA. I. S. pusllla Pursh. Sterile fronds linear, very slender, flattened and tortuous ; fertile ones equally slender, 3'— -4' high, OSMUNDACEjE. 77 ly r- r- Is. le K, and bearing at top the fertile appendage consisting of about five pairs of crowded pinnae, forming a distichous spike. New Jer- sey; Grand Lake, Nova Scotia (£". G. Knighi) \ Newfoundland {De la Pytaie, Wagkorne). Family 4. OSMUNDACE^E R. Br. Plant body a stout suberect stem (rootstock) with clustered leaves. Sporangia with a rudimentary ring, opening longitudi- ally, borne in panicles on altered portion of the leaf. The family contains three genera, only one of which is represented with us. I. OSMUND A L. Flowering-tern. Fertile fronds or fertile portions very much contracted, bearing short-pedicelled, naked sporangia on the margin of the rachis-like divisions. Sporangia large, globular, openiv*g by a longitudinal cleft into two halves, bearing near the apex a few parallel striae, the rudiment of a transverse ring. Spores green. Named for Osmunder, a Saxon n .me for the divinity Thor. A genus containing six species mostly north temperate. * Fronds bipinnate,Jcrtile at the apex. 1. O. regalls L. Stipes tufted, i" — 1|^° long, erect, naked ; fronds 2° — 4° long, i^or more broad ; sterile pinnae 6' — 12' long, 2 — 4' broad ; pinnules oblong-ovate to lance-oblong, sessile or slightly stalked ; the fertile pinnules cylindrical, panicled ; tex- ture subcoriaceous ; rachis and both sides naked. {O. specta- bilis Willd., O. glaucescens Link.) Canada to Florida and Mis- sissippi. ** Sterile fronds biptnnaiifid, 2. O. Claytonlana L. Stipes tufted, 1° or more long, clothed with loose woolly tomentum when young, naked when mature; fronds 1° — 2° long, 8' — 12' broad; pinnae oblong-lan- ceolate with oblong, obtuse divisions; 2 — 5 pairs of central pinnae fertile fertile pinnules dense, cylindrical ; texture her- baceous. {0. interrupta Michx.) Canada to Kentucky, and northward. 3. O. cinnamomea L. (Cinnamon-fern.) Stipes dense- ly tufted, r or more long, the sterile and fertile fronds dis- '! 1 'm ill'- 'A ■A ni i V I ji 78 OUR NATIVE FERNS AND THEIR ALLIES. tinct, clothed when young with ferruginous tomentum ; sterile fronds smooth when mature, the pinnae bearing a tuft of tomen- tum at the base beneath, lanceolate, cut into broadly oblong, obtuse divisions ; fertile fronds contracted, bipinnate, with cin- namon-colored sporangia. In var.frondosa Gray, some of the fronds are sterile below, and sparsely fertile at the summit. (0. Claytoniana Conrad.) New England and Wisconsin to Florida. Family 5. CERATOPTERIDACEyE Underw. Plant body a short succulent stem with copious aquatic roots bearing a rosette of succulent leaves of two sorts. Spo- rangia irregularly scattered, sessile, with a broad ring or often devoid of one altogether. The family is represented by a single genus and species. I. CERATOPTERIS Brong. Floattng-fern. Sori placed on two or three veins which run down the frond longitudinally, nearly parallel with both the edge and midrib. Sporangia scattered on the receptacles, sessile, subglobose, with a complete, partial, or obsolete ring. Indusia formed of the reflexed margins of the frond, those of opposite sides meet- ing at the midrib. Name from Gr. Kepa^, horn, and nTeini, a fern. Contains a single tropical species. 1. C. thallctroides (L.) Brong. Stipes tufted, inflated, filled with large air-cells; fronds succulent in texture, the sterile ones floating in quiet water, simple or slightly divided when young, bi — tripinnate v/hen mature ; fertile ones bi — tri- pinnate; ultimate segments pod-like. Southern Florida. Family 6. POLYPODIACE/E Pres^ Plant body consisting of a creeping or erect stem (root- stock) bearing scattered or clustered leaves (fronds). Spo- rangia borne on the back or margin of the leaf in lines or rounded masses (sori), stalked, provided with a vertical elastic ring, breaking open transversely at maturity. Sori either naked POLYPODIACEjE. 79 )po- or Lstic ked or covered when young with a membranous indusium which is either a special outgrowth of the leaf or is formed from tho more or less altered leaf margin. Prothallium green, monoe- cious or rarely dioecious. The family contains four-fifths of all the ferns, comprising a hundred or more genera, of which twenty-nine are represented within our limits. They may be distinguished by the following ARTIFICIAL SYNOPSIS OF GENERA. X. Sporophyll closely rolled together, entire!/ unlike the sterile leaf, its seg- ments berry-like or necklace-like ao Sori covered with indusia a Sori naked 3 a. Sori marginal, covered with a reflexed portion of the leaf margin . . 10 Sori dorsal or submarginal, provided with special indusia .... 14 3. Sori spread in a stratum on the under surface of the leaf. I. ACROSTICHUM, p. 81 Sori roundish, or not more than twice as long as broad 4 Sori usually linear, always more than twice as long a& broad ... 7 4. Stipes articulated to the rootstock ; leaves (in our species) entire or simply pinnate 5 Stipes not articulated to the rootstock ; leaves (in our species) bi — tripin- natifid or ternate XXII. Phegopteris, p. 108 5. Veins free or uniting irregularly (often indistinct). II. Polypodium, p. 81 Veins copiously uniting (species sub-tropical) 6 6. Primary veins distinct to the edge, connected by parallel transverse veinlets IV. Campyloneuron, p. 83 Areolae regular, each with two or more free veinlets bearing sori on their apices III. Phlebodium, p. 83 Areola; copious, inegular with free veinlets spreading variously. V. Phymatodes, p. 84 7. Leaves simple 8 Leaves pinnate to quadripinnate 9 8. Leaves very narrow, grass-like ; veins indistinct, free. IX. VlTTARIA, p. 89 Leaves broader ; veins anastomosing . . VIII. Cheilooramma, p. 88 9. Sori marginal, more or less confluent in a marginal band. VII. NOTHOLiENA, p. 85 Sori dorsal, following the veinlets, simple, forked, or pinnate. VI. Gymnoptkris, p. 84 Hi;: -*• ii SI I t 80 OUR NATIVE FERNS AND TffBIR ALLIES. 10. Sporangia at the ends of the veins, borne on a reflexed portion of the margin of the leaf X. Adiantum, p. 89 Sporangia borne on a continuous, marginal, vein-like receptacle con- necting the apices of the veins 11 Sporangia at or near the ends of unconnected veins, borne on the under surface of the leaf 13 IX. With an inner indusium inside the receptacle . XII. Ptkridium, p. 91 With no inner indusium ; our species pinnate . . XI. Ptcris, p. 90 za. Fronds conspicuously dimorphous ; stipes light-colored. XIV. Cryptooramma, p. 97 Fronds nearly uniform ; stipes usually dark-colored .... 4. 13 13. Sori on the upper part of the veins, mostly forming a continuous mar- ginal band : i.^dusium membranous, continuous round the segment. XV. PELLiEA, p. 97 Sori minute, at the ends of the veins ; indusium interrupted, or if con- tinuous, the ultimate segments usually small and bead-like ; leaves mostly chaffy, woolly, or farinose, , . . XIII. CheilantHES, p. 91 14. Sori roundish ; indusia not more than twice as long as broad ... 15 Sori linear or oblong ; indusia more than twice as long as broad . . ai 15. Indusium superior, attached by the centre or sinus 16 Indusium convex, fixed by a broad base partly under the sorus, XXVIII. FiLix, p. 119 Indusium inferior 19 16. Sori mostly on the back of the veins 17 Sori at the end of a free vein ; indusium reniform, opening toward the margin of the leaf, which is simply pinnate, the pinnae articulated totherachis XXVII. Nephrolepis, p. 118 17. Indusia centrally peltate 18 Indusia cordate or reniform, attached by the sinus. XXIII. Dryopteris, p. no 18. Veins free, i — a-forked XXIV. Polystichum, p. 115 Veins 3 — 5-forked, often uniting in irregular areolx ; leaf pinnate. XXV. Phanerophlebia, p. 117 Veins copiously anastomosing ; leaf trifoliate. XXVI. Tectaria, p. n8 19. Indusium roundish or stellate, delicate , . . XXXI. WoODSiA, p. xao Indusium cup-shaped, somewhat two-valved. XXXII. DENNSTiEDTIA, p. laa 20. Leaves growing in crowns ; veins free . . XXX, Matteuccia, p, lao Leaves scattered ; veins copiously anastomosiof XXIX. Onoclea, p. ISO at. Sori all parallel to the midribs or rachises aa Son all oblique to the midribs •34 ■ Ii; POLYPODJACEjE. 81 Sori partly oblique and partly parallel to the midrib ; leaf timple, tapering to a point XXI. Camptosorus, p. io8 aa. Veins free 33 Veins reticulate XVIII. Woodwardia, p. loa 33. Indusium near the margin ; sporophyll much contracted. XVI. Struthiopteris, p. 101 Indusium remote from the margin; leaves nearly uniform. XVII. Blechnum, p. I02 34. Sori on the upper side of a veinlet, rarely on both sides. XIX. ASPLENIUM, p. 103 Sori confluent in pairs, with an apparently double indusium opening in the middle XX. Phvli.itis, p. 107 I. ACROSTICHUM L. is Sporangia spread over the whole surface of the frond or upper pinnae, or occasionally over both surfaces. Venation anasto- mosing (our species simply pinnate). Name from Gr. aKfjos, the summit, and anxoi, a row. A tropical genus containing about five species as here limited. 1. A. aureum L. Rootstocks erect, solitary or in masses; stipes cespitose, erect ligneous, H° — 2^° long, flattish-channelled, with two or three alternate pairs of black indurated spurs, fronds erect, stiflf, coriaceous, glossy, light green; 3° — ^4" long; 1* — ii° wide ; pinnae 12 or more pairs, rather distant; sporangia confined^ to the upper half or third of the frond ; venation fine, oblique to the margin. Southern Florida. 2. A. lomarloldes Jenman. Rootstocks erect massive ; stipes cespitose, erect, subfleshy, li* — 2^" long, longitudinally ribbed, subangular ; fronds erect-spreading, 3" — 4° high, i" — 2" wide, slightly reduced at base, suddenly reduced at the apex ; pinnae patent, close or crowded, 25 — 30 pairs, the face turned upward and transverse with the rachis ; sporangia covering all the pinnae of fertile fronds ; areolae very fine, directed toward the margin. South Florida. 'i] II. POLYPODIUM L. Sori round, naked, dorsal, in one or more rows each side of midrib, or irregularly scattered, Stipes articulated to root- 8a OUR NATIVE FERNS AND THEIR ALLIES, Stock. Name from Gr. ;roAtJ5, many, and wot??, no^oi, foot, alluding to the branching rootstock. The largest, most cosmo- politan genus of ferns, containing 1 50 or more species. § I. EUPOLYPODIUM. Vtins free ; fronds {in our species) pinnate. * Sori large, 1. P. vulffare L. Stipes 2' — 4' long, firm, erect; fronds 4' — 10' long, i' — 3' broad, cut nearly or quite to the rachis into entire or sli^^.htly toothed, usually blunt pinnae ; veins once or twice forked. Larger fronds with their pinnae sharply serrated and long-pointed form the var. occidentale Hook. New England westward to Oregon and southward to Alabama. 2. P. falcatum Kellogg. Stipes 5' — 8' long, stramineous; fronds 12' — 15' long, 4' — 8' broad ; pinnae numerous, tapering to a slender point, sharply serrate ; sori nearest the midrib; veins with 2 — 4 veinlets. {P.glycyrrhiza D. C. Eaton.) California to British Columbia. ** Sori smaller, often minute. 3. P. plumula H. B. K. Stipes i' — 4' long, black, slender; fronds narrowly lanceolate, 9' — 18' long, i' — 2' broad; pinnae nu- merous, narrow, entire, blunt, lower gradually reduced ; surfaces naked except the black wiry rachis ; veinlets forked, bscure. Florida. 4. P. pectinatum L. Stipes rigid 2' — 6' long ; fronds el- liptical-lanceolate, 1° — 2^° long, 2' — 6' broad, cut to the rachis into horizontal, entire or toothed pinnae, the lower ones much reduced ; rachis naked or finely villose ; veinlets pellncid, once or twice forked , sori in long rows, of medium size. Florida. §2. GONIOPHLEBIUM Blume. Veins forming ample regular areola {almost imperceptible in No. 5), each with a single distinct free included veinlet, bearing a sorus at its terminus. * Under surface squamous. 5. P. polypodloides (L.) Hitch. Rootstock creeping, cov- ered with small brown scales; stipes i' — 4' long, erect, densely scaly; fronds 2'— 6' long, i' — \\' broad, cut to the rachis into entire pinnae; texture coriaceous ; sori small; veins indistinct, C/*. incanum Swz.) Virginia to Illinois, and southward^ POLYPODIACEM. ^ 1.0 I.I 1.25 IM 12.5 m m 10 m 12.2 2? Ki 'i NIMU U 11.6 • : V] /] /: 7 Hiotographic Sciences Corporation 23 WIST MAIN STRIET WEBSTER, N.Y. 14580 (716)S72-4503 ^ o [V ^ ^ ^ 84 OUR NATIVE FERNS AND THEIR ALLIES. i lar arches. Areolae similar, containing two or more sori. Name from Gr. Ka/nit^Xoi, curved, and vevfiov, a nerve. A tropical genus of about 50 species. I. C. phyllitldis (L.) Presl. Rootstock stout, scaly; stipes short or none; fronds simple, i* — 3* long, i' — ^4' broad, the point acute, lower part gradually narrowed ; texture rigid, cori- aceous ; areolae in rows of 6 — 12 from midrib to edge. Florida. V. PHYMATODES Presl. Sori round, naked, dorsal, various in position. Veins forming fine, copious, irregular areolae with free veinlets spreading in various directions. Name from Gr. 90 OUR NATIVE FERNS AND THEIR ALLIES, \ \ 3. A. Jordan! C. Muell. Stipes rather stout, nearly black, polished; fronds ovate or deltoid-pyramidal, bi — tripinnate; pinnules and upper pinnae ample, smooth, or nearly so, rounded or even reniform, upper margin rounded, slightly incised ; sort 2—5, transversely linear-oblong, subcontinuous. {A. entargina- tum of former edition not Hook.) tt Fronds pilose, with whitish hairs. 4. A. trioholepis Fee. Stipes smooth, polished, deep black ; fronds oval ; pinnules roundish, moderately long-stalked ; sori few (3 — 7), of unequal size; indusia very velvety. {A, 4