On the Trail of Equisetum for Four Thousand Miles

American Fern Society

On the Trail of Equisetum for Four Thousand MilesAuthor(s): John H. SchaffnerSource: American Fern Journal, Vol. 16, No. 3 (Jul. - Sep., 1926), pp. 81-92Published by: American Fern SocietyStable URL: http://www.jstor.org/stable/1544267 .

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ON THE TRAIL OF EQUISETUM ON THE TRAIL OF EQUISETUM

and covered 4,000 miles. How times have changed in a

quarter century! No Notholaena was seen the past summer, but it may still be present, since no careful search was made, no ladder being available to reach the tops of the large concretions at the west end. The powdery cloak-fern is also an odd little plant. From above it appears like a very diminutive Pellaex. But the waxy- powdery material on the underside of the leaves gives the plant a chalky appearance, making recognition easy.

There are, therefore, four species to the credit of this peculiar sandstone formation, which is an exceedingly good showing for a region where one may walk for many miles, with never a sight of a fern, usually to be rewarded finally by but one or two species growing around sdme far-isolated spring or on some special overhanging, shaded ledge of a creek or deep ravine. Who can show a more remarkable fern habitat than that presented by this field of Titan marbles of the prairie?

COLUMBUS, OHIO

On the Trail of Equisetum for Four Thousand Miles'

JOHN H. SCHAFFNER

Knowing that most of the species of Equisetum of eastern North America are to be found in abundance in the region of the Great Lakes and having read the allur- ing literature of the Michigan Tourist Associations, the writer decided to make an extensive tour in the summer of 1925 with his family, by automobile, with the primary purpose in view of studying horsetails in their native

1Papers from the Department of Botany, The Ohio State Uni- versity, No. 167.

and covered 4,000 miles. How times have changed in a

quarter century! No Notholaena was seen the past summer, but it may still be present, since no careful search was made, no ladder being available to reach the tops of the large concretions at the west end. The powdery cloak-fern is also an odd little plant. From above it appears like a very diminutive Pellaex. But the waxy- powdery material on the underside of the leaves gives the plant a chalky appearance, making recognition easy.

There are, therefore, four species to the credit of this peculiar sandstone formation, which is an exceedingly good showing for a region where one may walk for many miles, with never a sight of a fern, usually to be rewarded finally by but one or two species growing around sdme far-isolated spring or on some special overhanging, shaded ledge of a creek or deep ravine. Who can show a more remarkable fern habitat than that presented by this field of Titan marbles of the prairie?

COLUMBUS, OHIO

On the Trail of Equisetum for Four Thousand Miles'

JOHN H. SCHAFFNER

Knowing that most of the species of Equisetum of eastern North America are to be found in abundance in the region of the Great Lakes and having read the allur- ing literature of the Michigan Tourist Associations, the writer decided to make an extensive tour in the summer of 1925 with his family, by automobile, with the primary purpose in view of studying horsetails in their native

1Papers from the Department of Botany, The Ohio State Uni- versity, No. 167.

81 81



AMERICAN FERN JOURNAL

haunts. The trip was begun from Columbus, Ohio, on July 5.

Just north of Columbus, Indian Springs ravine was passed where Equisetum praealtum and E. arvense grow in great abundance. Here many interesting observations on our two most common species have been made for many years. E. praealtum covers the wide ravine bottom in various places like a thick stand of tall grass, com- peting successfully with bluegrass, blackberries, and other plants, since most of the trees have been cut down years ago and the ground has not been pastured. The

largest patch covers more than an acre of ground. The plants are usually from 2 to 4 feet high and rigidly erect. The branches developed on erect stems are also rigidly erect, growing up close to the main stem. At whatever angle the main stem is bent over the branches developing on it are perpendicular to the surface of the earth. The branches produce small cones abundantly. The main stems often produce zones of nodes without appreciable internodal development or with very short internodes. This development is entirely sporadic and may occur at several points on the same stem. Occasionally dichotom- ous tips with two cones are produced but they are difficult to find. In winter the internodal cavities are often turgid with water which in cold weather freezes into solid cylin- ders of ice. Many stems take on a dull reddish color on the side exposed to the winter sun. This color is not anthocyanin but' is due to a red pigment in the chloro- plasts. The main blooming period is in June, although isolated cones may be shedding their spores- from May to September. Young cones of various sizes can be collected at any season of the year. Very few cones are produced in the drier parts of the patch. The mature cones vary in size all the way from minute structures no larger than

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ON THE TRAIL OF EQUISETUM

pin heads to the perfectly developed cones three fourths inch long and one half inch in diameter. The smaller cones do not develop spores, in fact the merest rudiments of sporangia are produced. The intermediate sizes are usually semi-sterile and produce imperfect spores. This variation in the development of the cones in every gradation from vegetative tip to sterile, semi-sterile, and fertile structures is characteristic of various Equiseta and is, no doubt, the basis for the reports of supposed hybrids and many special varieties. These semi-sterile shoots in- dicate the low position of Equisetum in the evolutionary scale of the vascular plants. In the higher plants there is usually a definite vegetative shoot or a definite flower. There is no intergrading series. In the lower Equiseta the number of whorls of sporophylls and also the number of sporophylls in a whorl are variable, just as is also the case with the number of sheath segments and teeth. In the higher Equiseta the tendency is to have the numbers more constant.

Having passed our favorite wild Equisetum garden, the main observation in northwestern Ohio and southern Michigan was the widespread abundance of E. arvense. It grows in all sorts of situations-along brooks and

streams, in meadows and woods, in sandy places, and especially on railroad embankments. It may be regarded as the highest species and the most successful one, with the widest distribution over the earth. The habit of its body is as diverse as its habitat. In the woods it is often a tall, erect plant with regular whorls of simple branches standing out at right angles to the main stem or sometimes slightly drooping. The branches may be com- pounded to various degrees. In the open the plant often forms dense tufts or reclining, irregularly branched masses. On sand the form is frequently a perfect mat with prostrate stems and branches radiating from the

83




center and closely applied to the substratum. There are all gradations from fertile to sterile shoots but the intermediate forms are rare and it was much too late in the season to find any.

In several places near Ann Arbor, E. praealtum was noticed and after turning north toward the region of the conifer forest, which originally covered the Great Lakes area, the search for horsetails began in earnest. At Johnson's Landing, in Genesee county, E. laevigatum was found in fruit, in abundance, along with E. arvense. Northwest of Midland, E. praealtum was collected on the high river banks. At Clare, farther on, E. arvense was taken simply as a sou-enir of the locality.

East of Houghton Lake, some distance north of Pru- denville, a great abundance of E. fluviatile was found. Both types of shoots, the unbranched and those with whorls of branches, were present. In central Ohio the plants usually have the whorls of branches. As one goes northward the simple-shoot type seems to become more abundant-the so-called variety E. limosum. Both types of shoot come from the same rhizomes, so the unbranched form should not be called a variety. In Ohio the cones of E. fluviatile show abundant proliferation, frequently with a second cone at the tip. These proliferated cones are usually imperfect. At Houghton Lake the tall sterile plants showed the slender, spiral and coiling tips which are frequently developed in this species. The coils may be more or less circinnate or spiral, or they may be merely flexuous or hooked or bent in various ways. These coils are often intertwined or coiled about some other plant. The coiling tip is usually from one to four inches long. There is probably no twining by contact. I have seen these slender coiling tips at various times but never in such abundance and with such perfect spi- rals and coils.

84




At Gaylord, in Otsego county, the banks of a railroad cut going through the forest were covered with large E. praealtum. On examination, they were found mostly to be decidedly bent and curved. This was in striking contrast to the rigidly erect condition characteristic of the

species. I tried to discover the reason for the curved stems and concluded that they might have been bent

through the gradual melting of heavy snow, either drifted or thrown upon the plants by snow ploughs.

Coming to the south end of Burts Lake, in Cheboygan county, I examined a wooded marshy shore with old fallen trees and decaying logs, and soon found abundant patches of E. scirpoides. This low tufted plant with its numerous dark-green, slender, curly flexuous shoots is of special interest in many ways. The wavy-curly tufts show extreme specialization away from the straight erect growth of its more primitive relatives. The stems turn rather whitish when old, which contrasts strongly with the dark green living portions. The shoots are quite uni- formly grooved and the sheaths three-toothed. The six ridges appear to represent three original ones, which have become double by the development of a central groove in each ridge. The cones are very small, as one would ex- pect on such small shoots. The constancy of the ridges and teeth is in striking contrast to the variability of these structures in E. laevigatum and E. praealtum.

A single little plant of typical E. variegatum was found in this swampy woods. Thinking that there would be an abundance of it further north no further search was made for it. Great was my surprise when I was un- able to find it again during the entire journey. I looked in every conceivable place and habitat, but it eluded me. All of the other ten species of eastern North America were seen at various times but no more E. variegatum.

85




One of the excuses for making the trip was to be able to visit the University of Michigan Biological Station on Douglas Lake in order to study Equisetum nelsoni. Through the kindness of Dr. John H. Ehlers, of the Uni- versity of Michigan, and Dr. George E. Nichols, of Yale University, I was able to visit habitats where various species grew in abundance. We collected E. scirpoides, E. nelsoni, E. arvense, E. praealtwm, E. silvaticum, E. laevigatum, and E. palustre. Besides these E. fluviatile and E. variegatum are also found in the region but were not collected. A careful study of E. nelsoni in the field convinced the writer that it is well entitled to specific rank.

In a boggy swamp in an abandoned road passing through a forest, near the shore of Burts Lake, was a wonderful stand of Equisetum palustre. A great number of forms was collected growing in various habitats and in various degrees of light and shade. The forms

ranged from tall plants with regular whorls of branches, through bushy and irregularly branched forms, to small unbranched shoots of various sizes. The species has a

very wide range of habit. This patch of E. palustre pro- vided an unusual opportunity for studying the cones which were in full bloom (July 10). The mature cones are of all sizes from as large as pin heads on very slender

tips to the normal large cones on robust shoots. There is in E. palustre a complete gradation from fertile shoots through semi-sterile shoots to the ordinary sterile shoots. The semi-sterile small cones are very common. I am now convinced of what I had tentatively concluded from a study of herbarium material, both from Europe and America, that E. litorale is not a hybrid nor a species but merely the ordinary semi-sterile shoots of E. palustre. It is probable that all of the supposed hybrids of Equisetum, based on defective sporangia and spores, are

86




all merely examples of semi-sterile shoots which develop from the same rhizomes as the normal fertile shoots. Morphological studies on the cones of Equisetum must also take account of this condition of affairs, for it is probably the basis for the diverse types of sporangium development which have been reported. One could probably obtain a normal, developmental series by selecting the young cones from shoots with large robust tips. Any cones taken from slender-tipped shoots will probably be semi-sterile or sterile.

After crossing the straits of Mackinac we made our first camp about thirteen miles north of St. Ignace. Be- sides the ever-present E. arvense, E. scirpoides was abundant. While walking along a north-facing bluff of the Carp River I saw my first E. pratense, growing in its native habitat. This is a most delicate and beautiful species of a pale gray-green color, with symmetrical whorls of simple thread-like branches which put one in mind of spider webs arranged in tiers. It is very easily distinguished from E. arvense, both in the field and in herbarium specimens, not only because of its filmy appearance but especially by the triangular white teeth which are without the subulate point characteristic of E. arvense.

At Sault Ste. Marie, E. silvaticum, E. fluviatile and E. palustre were collected. E. fluviatile was mostly of the unbranched form and many shoots showed the coiling tips. A great variety of forms of E. palustre was growing in grassy meadows, in ditches and on the sandy shore of St. Mary's River-tiny tufted forms with unbranched shoots, tall unbranched specimens, and the usual run of irregular shoots and shoots with regular whorls of branches. The cones also showed all gradations in the various habitats, from perfect cones to the minut-

87




est sterile strobili. Here was also collected a specimen with a spiral sheath, extending around the stem for over an inch, with a spiral row of lateral branches. These forms in which the sheath develops as a spiral are occa- sional in E. palustre and can often be found by diligent search.

On the shore of Mille Coquins Lake, E. palustre was abundant and in full bloom. The cones were again in all grades of size and perfection from those normal and fertile through the various degrees of semi-sterile ones over to the completely sterile tips. At Rapid River Falls Park, northwest of Rapid River, in Delta County, Equi- setum pratense was rather common and besides the typical form, low irregularly branched tufted specimens were found growing on the bare sand of the roadside. How-

ever, it is apparently not so variable in form as E. arvense. The locality also yielded the following species: E. scirpoides, E. silvaticum and E. arvense. At Muni- sing, on the Lake Superior shore, E. laevigatum was collected.

At Michigamme, just below the beautiful state camp site and park, E. fluviatile covered great areas of marshy ground. It was nearly all of the "limosum" type with unbranched shoots. Only here and there would a shoot appear with the typical whorls, although many would have a few isolated short branches at some of the nodes. Dense tufts of small delicate shoots of this species were also collected on the bare sand. They would be difficult to identify by their morphological characters, especially when dried.

West of Michigamme, along a sandy roadside cut through the conifer forest, Equisetum silvaticum oc- curred in abundance. There were all gradations of form from prostrate mats on the bare sand through irregularly

88




branched and tufted individuals to the well developed shade forms with symmetrical whorls of branches. Those in the dry exposed sand were of a decided yellowish tint.

Shirley Hibbard has called the wood horsetail "the most elegant of all the plants upon the face of the earth. " In typical shoots the branches of the whorls spread out horizontally in broad circular disks with slightly drooping margins forming a series of horizontal platforms, which usually become smaller toward the top, giving the whole plant the appearance of a symmetrical conifer. The whorls of branches are horizontal with the surface of the earth even if the stem is inclined. They form a striking contrast to the rigidly perpendicular branches of E. praealtum. In its perfect form it is certainly a graceful and elegant plant, but is not so delicate and ethereal in aspect as E. pratense.

At Millicent Lake, near Iron River, Wisconsin, E. arvense, E. praealtum, E. silvaticum, and E. fluviatile were found. The E. fluviatile was largely of the simple- stemmed form.

North of Duluth, at Twin Harbor, Minnesota, E. fluviatie, both the branched and the unbranched form, was collected and an assortment of E. arvense. Among the special forms of the latter species were mats and a most delicate, symmetrical, erect type, four to eight inches high, with whorls of short rigid branches standing out nearly at right angles and from one third inch to one inch long. Many such forms have received formal names but I believe this is a waste of energy and a great detri- ment to systematic botany. We certainly do not want to name our individual plants in the way we give pet names to our cats, dogs and canaries. There has been much nam- ing of fluctuating branches in Equisetum. It makes one shudder to think of what would happen to our manuals

89




if systematists should begin to describe and give formal names to all the diverse types and forms of branches to be found on our trees and other branching species.

After passing northward along the lake shore as far as the Manitou River we turned southward and at Farm- ington, Minnesota, found a large patch of E. kansanum in a little prairie on sandy loam. Accordingly, this species was studied for the gradation series of semi- sterile shoots and cones. Every gradation was collected, ranging from semi-sterile shoots with cones as large as small pin heads to the normal, spore-producing strobili. As stated above, the presence of defective cones, sporangia, or spores in Equisetum is no indication of hybridity. The way to find out what hybrid Equiseta would be like is to produce them under controlled conditions.

Near Harris, Minn., both E. praealtum and typical E.

laevigatum were collected. Further on, near St. James, E. laevigatum was growing among prairie grass on a

typical Andropogon prairie, much as one finds E. kansanum in southern Nebraska and northern Kansas, E. arvense was also growing as a normal member of the

prairie association. Two days were spent at Lake Okoboji, Dickinson

County, Iowa, and thanks are due to Dr. Geo. W. Martin, of the Iowa State University, for aid in locating horsetails of this region. E. arvense and E. kansanum were

collected, the latter both in typical prairie and on the lake shore. E. praealtum and E. laevigatum have been

reported and no doubt occur in the locality but were not seen by the writer. After leaving Lake Okoboji, E. kansanum was collected in a gravel and sand pit near Denni-

son, Iowa, and west of Omaha, Nebraska, vigorous E.

laevigatum was found iit a railroad cut. South of Lin-

coln, E. kansanum was abundant in a natural prairie.

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We finally arrived at Morganville, Clay County, Kan-

sas, where the writer first became interested in Equi- setum because the plants were so different from every- thing else in the region, and because the most common species present would not fit the key and descriptions in the manual. Many years after these plants were de- scribed as E. kansanum. The original habitat of the type locality has been turned into a pasture for a number of

years and the cattle have browsed the plants so persis- tently that only small tufts could be found. The species was, however, abundant on ungrazed, mixed Andropogon furcatus-A. scoparius prairie in various places. E. laevigatum is abundant in the bottom land of the Repub- lican River and often persists for years in cultivated soil. Both E. arvense and E. praealtum have been collected in the past in Clay County but none were found.

After staying for a month on my farm, a trip was made through western Kansas. After leaving Clay County and passing through Grainfield in Gove County and Dighton in Lane County not a single Equisetum was seen in the 572 miles traveled until we reached Emporia in Lyon County, except at one place. West of Ness City, Ness County, I observed a high, steep, north-facing bluff along a creek which looked like a suitable habitat and on investigation it was found to harbor a considerable quan- tity of E. kansanum growing among the Andropogon scoparius. Of course, with special search some Equi- setum would have been found at other places, but Equi- seta and all other pteridophytes are exceedingly rare. Three years ago the writer collected E. laevigatum on a sandy bank of the South Fork Solomon River near Penokee, Graham County, and in Colorado near the foot- hills E. kansanum is often abundant along the sand-filled stream channels. At Emporia, Lyon County, E. praeal- turn was found to be common on river banks.

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AMERICAN FERN JOURNAL AMERICAN FERN JOURNAL

At Marshall, Missouri, E. arvense and E. laevigatum were abundant in a sandy creek bottom, and at Warren-

ton, Warren County, giant specimens of E. praealtum were collected, growing in a ditch in the flood plain of a creek. Some of these plants measured five feet eight inches tall and were one half inch in diameter. Similar

large specimens were seen in the Missouri River valley near St. Charles. Near Effingham, Illinois, E. arvense, E. laevigatum and E. praealtum were abundant, the latter sometimes measuring one half inch in diameter and

slightly over six feet in height. Having collected all of the common species many times

and feeling quite certain that only two other species could

possibly come into the region bordering the road home, namely, E. fluviatile and E. kansanum, no further time was taken for botanizing.

The final drive was made on Sept. 7 from Teutopolis, Illinois, to Columbus, Ohio, where we arrived safely in the evening, the speedometer indicating that from the

departure to the return something over 4,000 miles had been covered, yet we had barely reached the half-way line across the continent.

COLUMBUS, OHIO

The West Virginia Locality of the southeastern Relative of Woodsia scopulina

EDGAR T. WHERRY AND FRED W. GRAY

In a recent publication' Professor M. L. Fernald in- cludes the following footnote to the discussion of the distribution of Woodsia scopulina:

Persistence of plants in unglaciated areas of boreal America. Mem. Am. Acad. Arts Sci. 15: 250. 1925.

At Marshall, Missouri, E. arvense and E. laevigatum were abundant in a sandy creek bottom, and at Warren-

ton, Warren County, giant specimens of E. praealtum were collected, growing in a ditch in the flood plain of a creek. Some of these plants measured five feet eight inches tall and were one half inch in diameter. Similar

large specimens were seen in the Missouri River valley near St. Charles. Near Effingham, Illinois, E. arvense, E. laevigatum and E. praealtum were abundant, the latter sometimes measuring one half inch in diameter and

slightly over six feet in height. Having collected all of the common species many times

and feeling quite certain that only two other species could

possibly come into the region bordering the road home, namely, E. fluviatile and E. kansanum, no further time was taken for botanizing.

The final drive was made on Sept. 7 from Teutopolis, Illinois, to Columbus, Ohio, where we arrived safely in the evening, the speedometer indicating that from the

departure to the return something over 4,000 miles had been covered, yet we had barely reached the half-way line across the continent.

COLUMBUS, OHIO

The West Virginia Locality of the southeastern Relative of Woodsia scopulina

EDGAR T. WHERRY AND FRED W. GRAY

In a recent publication' Professor M. L. Fernald in- cludes the following footnote to the discussion of the distribution of Woodsia scopulina:

Persistence of plants in unglaciated areas of boreal America. Mem. Am. Acad. Arts Sci. 15: 250. 1925.

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On the Trail of Equisetum for Four Thousand Miles