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Fernwort papers presented at a meeting of fern students, held in New York city, June 27, 1900, under the auspices of the Linnaean fern chapter ... online

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8 I










Issued December 20, 1900.

Printed for the -Linnaean Fern Chapter.




WILLIAM R. MAXON, U. S. National Museum,

Washington, D. C.

Vice-Presiden t
B. D. GILBERT, Clayville, N. Y.

Miss MARGARET SLOSSON, Andover, Mass.

JAMES A. GRAVES, Susquehanna, Pa.

This Chapter, founded in 1893, has as its aim
the study of ferns by correspondence, and the publi-
cation of the results of such study. It comprises
nearly 125 members from all parts of the United
States, and from Canada and Great Britain. All who
are interested in ferns are cordially invited to join.
Dues are one dollar annually ; the initiation fee of
one dollar includes the dues of the first year. Mem-
bers receive without cost the Fern Bulletin quar-
terly and Annual Reports ; other publications either
without cost or at a rate considerably lower than
available to other than Chapter members. For full
information address either the President or Secretary.



O ,

^ / \/w n 2




the Chapter. Issued December 20. 1900



In the three seasons that have elapsed since the appearance of
Dodge's "Ferns and Fern Allies of New England," the study of
our local species of the genus hoetes has progressed so far that it
seems advisable at this time to present some of the results, and
state some of the problems, in order to elicit the co-operation of
collectors in clearing up some of the unsettled questions. As the
settling of these questions will pave the way to a larger and more
comprehensive work, this paper may be considered as preliminary
only .

It appears advisable to give a history of the genus, so far as our
chosen territory is concerned, and in doing this we can do no better
than to adapt and extend Engelmann's chronological history in his
comprehensive account of the genus, given in Trans. St. Louis
Acad. 4:358-390. 1882.

According to this, the first collection was at Upbridge, Mass.,
by Robbins, in 1831. It was referred to riparia.

1840. Robbins collected lacustris in the same town.

1843. Tuckerman collected lacustris in Echo Lake, N. H.

1845. Robbins found echinospora Braunii in Massachusetts.

1848. Tuckerman found near Boston the species which bears
his name.

1856. Engelmann found echinospora Braunii in Lake Winni-
piseogee, New Hampshire.

1857. E. D. Eaton found Engelmanni in New England.

* For an account of this Meeting, see Fern Bulletin for July, 1900. The first
gathering of those interested in American fern study occurred in Boston,
August 24, 1898, and the papers read were published early in 1899, under the
title, " Papers Presented at the Boston Meeting."


1 1860. Boott discovered echinospora muricata.

1865. Boott rediscovered Tuckermani.

1867. Boott collected echinospora Boottii.

1878. Pringle collected echinospora robusta in Lake Cham-

Engelmann also mentions other New England collections, but
they are not given in his list. After his time there is a hiatus in
publications, but collectors were accumulating undigested material,
and wrongly referring it often. The localities for Braunii have be-
come so numerous that it appears useless to enumerate them.
They occur in all of the New England States.

1887. Kennedy found what is now known as Tuckermani
borealis in Somes stream, Mt. Desert.

1889. Dr. C. B. Graves found echinospora muricata in Con-

1892. Fernald collected Tuckermani and Harveyi at Mt.
Desert, and Rand discovered what is now the type of heterospora
at the same place.

1893. Pringle and Fernald collected the types of hieroglyphica
at St. Francis lakes. Dodge discovered Engelmanni at Newbury.

1894. Dodge discovered Tuckermani and Braunii at New-

1895. Coville discovered Tuckermani borealis at Kennebago
lakes, and hieroglyphica at the Rangeleys. Raynal Dodge and
myself found Tuckermani at Essex, Mass., and severally discov-
ered it at Amesbury. I also found it at Kingston, N. H., and dis-
covered muricata at Kingston and East Kingston, where I found
the types of Eatoni and one specimen of Dodgei in a pond running
into Kingston. Graves collected Gravesii in Connecticut.

1896. I discovered Dodgei at Kingston in abundance, foveo-
lata, Eatoni, echinospora robusta, muricata and Tuckermani at
Epping, N. H., and the last at Nottingham. Dr. Graves collected
Tuckermani in southern Connecticut.

During the years 1895 and 1896 I found Engelmanni to be com-
mon in Rockingham county, N. H., and echinospora Braunii in
many places.

1898. Harvey discovered the type of Harveyi at Pushaw pond,
Oldtown, Maine. I found Eatoni at Amesbury, and Dodge at New-
bury, Mass.

1899. Harvey found Tuckermani botealis at Oldtown, Maine,
hieroglyphica and muricata at Moosehead lake. 1 traced Eatoni,



foveolata, and muricata to Newmarket, N. H. Underwood found
some peculiar plants, not clearly referable to any published species,
at Goshen, Conn. The spores are either abortive or unripe, and
unsatisfactory for study.""

It is hoped collectors will call my attention to the many gaps in
this list, that I may fill them.

A critical study, however, shows the remarkable fact that ri-
paria and lacustris, tho formerly considered common, are very rare
and have not been collected in recent years. It is true they have
been reported from various localities, but all such specimens which
have been accessible to me, have proven to be either Engelmanni,
Tuckermani, Tuckermani borealis, or an undescribed species.
It is unsafe to draw conclusions until some of the old material has
been re-examined, which I hope will be in the near future ; but
there is a growing probability that the former has rarely been met
north of the Delaware, and that the latter in its typical state is not
American. I cannot agree with Dodge in his determinations as
regards these species, and identify them mostly as Tuckermani.
Incidentally, his saccharata from the Merrimac is immature Braunii.

There is one other point in his work to which I take exception,
and that is the theory of hybridity of our species. Tho this may
well occur, I have never seen a case which I thought called for that
explanation. Eatoni, as is well known, usually bears only female
sporangia, only one plant in two or three hundred bearing male
spores, and then the sporanges occupy no regular zone, but are
intermixed with the others. It is evident, then, that this would be
a very good plant to experiment upon, as the macrospores could be
easily obtained without microspores. This has been done by Mr.
T. C. Palmer, who, I hope, will soon publish the results. I will
anticipate this, however, by stating that while straight cultures
yielded plants, attempted crosses between Eatoni and Dodgei or
Engelmanni were negative. So we have presumptive proof that
hybridity is extremely rare, if not altogether absent. Mr. Dodge
based his presumption on spore characters only, a very unsatisfac-
tory character, inasmuch as the sculpture of spores of all species is
very variable, in the echinosporas often confluent into walls, espe-
cially in muricata; while in Tuckermani spores are often found
mixed among normal ones that have the crests, or some of them,
broken up into spinules. As these spores have been found in

* Additional specimens from I/ynn, Conn., collected by Dr. Graves, prove
this to be an undescribed species. It is inserted under the name of /. Gravssii.

sporanges with normal ones, I think them explainable better on the
theory of accidental variation, than on that of hybridity, especially
as they are found where the species do not grew in juxtaposition.
My investigations on this point have been extensive, covering
thousands of plants, and my conclusion is that hybridity in this
genus, though not impossible, nor even improbable so far as ex-
ternals go, is at least extremely rare among New England species.

I wish here to pay tribute to the general excellence of Mr.
Dodge's treatment of the genus, a more practical account never
having been written ; and for the beginner, I can conceive of no bet-
ter directions than are contained in his notes. I wish I could say
as much for Britton and Brown's more pretentious "Illustrated
Flora," but to anyone with more than the most superficial knowl-
edge of the genus, their treatment is very disappointing.

Considerable labor has been expended by Durieu, Braun, and
Engelmann, to devise some system of classification that would hold
water, but with ill success, as they themselves testify ; for while it
is comparatively easy to classify species of a limited area, the sys-
tem fails when applied to the genus as a whole. Perhaps the most
natural grouping is into the three divisions Aquaticae or Submersae,
Amphibiae, and Terrestres, and if applied with sufficient looseness
will hold. But Submersae in Europe are bilobed and without
stomata, consisting of the species lacustris and echinospora. When
applied to the genus as a whole, however, we find Gunnii and elatior
of Tasmania three lobed, and also find our varieties of echinospora
with stomata, thus placing them in a different group from the type,
as indeed they appear from habit to fall. The system fails, too,
when taken in its original sense in the division Amphibiae. All
Old World species of this group are three lobed, and this was made
use of in the classification ; but not one of our numerous species of
eastern America is three lobed, and only four from North America,
Nuttallii, Cubana, Orcuttii and minima, are so constituted. The
only works treating of the genus as a whole, do not tend to clear
up the matter. Baker, in "Fern Allies," treats the genus very
superficially, while Motelay's more pretentious work is full of inac-
curacies, nearly two pages being given to " errata," and these cov-
ering a small percentage of those contained in the work. Though
a help in many ways, the critical student hesitates to place much
weight on statements accompanied by such errors.

The group Terrestres is, it seems to me, unnecessarily limited
to the two species with persistent leaf bases. Nuttallii, fiutleri, and


Montezumas of America, placed beside Duriaei or hystrix of
Europe, could not be separated by superficial examination, this one
point aside ; and some forms of Duriaei and hystrix, those growing
in damp places, habitually have no scales. On the other hand, the
three species mentioned can be separated at a glance from melano-
poda, Mexicana, or Pringlei. Leaf sections also are identical in
the Terrestres, the leaves being short, setaceous, with very small
cavities, and correspondingly wide dissepiments, and with four
stout bast-bundles (three in Nuttallii}, while the others named, as
well as most European, African, and Asian species which I have
examined, have stout leaves, four primary, and several accessory
bast-bundles, narrower dissepiments, and correspondingly larger
cavities. I leave a further discussion of these points for a future
occasion, especially as they bear only remotely on the immediate
subject in hand. I would remark, however, that Engelmann is at
variance, and I think rightly so, with European authors in his con-
ception of the subdivisions.

Probably the main reason that our species of Isoetes are not
better known, both in habit and distribution, is the difficulty novices
experience in finding them. Their unattractive appearance and the
tedious process of analysis by use of a compound microscope, re-
quired to determine them with accuracy, have also conspired to
prevent general study. No character given can be expected to be
true in all phases. The leaves are long or short, according to the
stage of the water ; they are stout or slender, and the trunk large
or small, to correspond to the general vigor of the plant. The
outer sporanges are usually nearly round, while the inner are two to
four times as long as broad. The velum varies greatly in some
species, in certain forms of echinospora varying from ^-% indusi-
ate, and I have seen specimens from Maine and Amesbury, Mass. ,
where a pin head would cover the opening. I should have consid-
ered this a good variety, had not fresh specimens from the same
place possessed a normal velum. The spores vary greatly in size,
and one may always expect to find some larger than the descrip-
tion allows. Location appears to regulate this. Juckermani, for
instance, is normal in this respect in most Massachusetts localities,
while New Hampshire and Maine plants usually measure 10-20 //

I know of no character more unreliable than the sculpture of
the spores. It is always easy to recognize Engelmanni in New
England, but southward even this breaks up into several varieties.

Eatoni usually has its peculiar convoluted ridges, but one often finds
a spore which is cristate. Dodgei shows the same tendency to
vary, and to a greater extent Tuckermani,foveolata, and the echino-
sporas. In the latter one may at times find a spore, or perhaps sev-
eral in a sporangium, with crests running across the upper faces,
while even the spines below are variable. Other spores grown
with them, will have very long, slender, forked spines. Indeed, I
do not at present feel competent to fix the limits of the varieties,
but leave them where Engelmann did ; or, where specimens have
been referred to one or another of the varieties, the arrangement is
tentative only.

I would remark, however, that while most specimens from my
neighborhood answer well to the descriptions of muricata and
Boottii, northern and mountain specimens usually have scattered,
short, blunt, flat spinules, most nearly corresponding to Braunii,
which, as originally described, is a mountain form.

The leaves of all species bear a central bast-bundle, but none of
ours have more than four peripheral ones, and only four have them
at all, Engelmanni, Eatoni, Dodgei and Gravesii. In well devel-
oped plants of the first, all four are pretty constant, but young or
starved plants are liable to have only one or two, perhaps none.
Eatoni is oftener without than with them.

All our species habitually have bi-lobed trunks, but tri-lobed
ones are not infrequently found, and in one species, Tuckerwani,
I find about 25 per cent, tri-lobed, and have found one four- and
another seven-lobed. While these plants appear to have been
chiefly tri-lobed from the seedling, occasionally one is found which
is apparently in process of transformation, One notable case was
a plant oifoveolata, collected at Newmarket, N. H., in 1899, which
bore a distinct lesion on one side, in which roots were forming.
The cortex between the lobe and lesion had not yet given way, but
in the natural course of events would soon have decayed, and prob-
ably the root-bearing area would have persisted, the plant thus be-
coming tri-lobed. Indeed, in the four- and seven-lobed Tucker-
inani specimens, there was evidence that a similar accident had be-
fallen them, and we might conjecture from this that the lower part
of the trunk is capable of becoming root-bearing, provided the cor-
tex be ruptured. Nothing conclusive has yet been observed on this
point and it needs a series of experiments to demonstrate the truth
or falsity of the hypothesis.

In the following arrangement I have based the key on charac-
ters which will most easily allow the determination of species with-
out recourse to the compound microscope. Though not perfect, I
hope it will be helpful to the general student,

Group i.

Plants habitually under water in the driest seasons, growing in
sand or gravel. Bast absent, and stomata few or none.


Leaves 1.5 to 2mm. in diameter, stiffly erect, stomata none.

1015 cm. long, spores (averaging under 600 //) covered with
short crests, rarely a little reticulate below. i. lacustris.

5-8 cm. long, spores larger (averaging over 600 n, at times
over 1000 /*), of various shapes, more densely cristate,
crests anastamosing but not reticulated. 2. heterospora .

Leaves spiral or recurved, stomata few or none.

Leaves very slender, under i mm. in diameter, reddish when
young, becoming olive-green, 8-15 cm. long, stomata in
single series over air cavities, or none, macrospores wavy
crested above, more or less reticulate below, 600 /j. or less
in diameter. 3. Tuckerniani.

Leaves dull green, stouter, about imm., not spiral, slightly
recurved, stomata not seen, spores larger, averaging above
650 /*. 33. Tuckermani borealis.

Leaves stouter, over i mm. in diameter, shorter, 8-iocm.,
rigidly recurved, spores less than 600 fj., loosely covered
with vermiform wrinkles. 4. hieroglyphica.

Leaves very stout, 2.5-3mm., short, 5-6cm., purple bronze in
color, rigid, recurved, spores as in No. 3. 5. Harveyi.

Group 2.

Plants growing usually in mud on borders of ponds or rivers,
inundated most of the year, but fruiting as the water recedes. Sto-
mata abundant but peripheral bast-bundles none. Never far from
water, and always in very damp soil. Amphibiae.

Spores reticulated below, jagged cristate above, much as in
No, 3, but leaves stouter and erect. 6. riparia.

Spores smaller, averaging 440 /*, covered with very small
pits. 7- foveolata.

Spores covered with spinules, stomata absent.

8. echinospora, not American.

Stomata present. of the spines short, broad, usually retuse, leaves 25 or
less. 8a. echinospora Braunii.

Like No. 8a, but leaves 25-70, and more densely stomatose.
8b. echinospora robusta.

Spines long and slender, sharp or forked, leaves stiffly erect.

Sc. echinospora Boottii.

Spines often mixed with short crests, otherwise as in No. 8a,
spores larger, leaves long and slender.

8d. echinospora muricata.

Group 3.

Plants growing in ditches or near the borders of ponds at higher
levels than the preceding, emersed during the greater part of the
summer. Stomata many, and bast-bundles (in our species) four.

Spores smaller (450 ^ or less).

Closely set with irregular, thick, rough, anastamosing

ridges. 9. Eatoni.

Densely covered with short, truncate columns. 10. Gravesii

Spores larger (averaging over 550 ft) with irregular, usually scat-
tered, variously anastamosing crests, ii. Dodgei.

Spores medium ( averaging about 450 fi } , regularly honeycomb-
Plants larger, with stouter leaves, and four bast-bundles,

12. Engehnanni.

Plants weaker, leaves 15 or less, often without bast-bundles.
I2a. Engehnanni gracilis.


Quite common in clear waters in northern Europe. Variously
reported from North America, but most reports have proven erro-
neous. The nearest approach to it I have yet met, was collected
by Coville in Kennebago lake, Maine, in 1895. I have referred his
plants doubtfully to Tuckermani borealis.

2. I. heterospora n. sp.

Trunk bi-lobed, leaves 50-75, 5-8cm. long, very stiffly erect,
nearly 2mm. in diameter, tapering to a sharp point, wanting stomata
or bast : velum J/j-% indusiate : sporangium spotted, often thickly
so, with dark cells ; macrospores normally 540-675 //, but occasion-
ally i loo or even 1134 n, densely covered with thick, jagged, con-
voluted crests, often honeycomb-reticulated below ; microspores
30.8-39.6 ft, averaging 35x27 p, dark brown, papillose.

This species differs from locus fris in its shorter, more numerous,
tapering leaves, spotted sporangium, papillose microspores, and
larger macrospores.

It is remarkable for the shape and size of the spores. Nearly
all the sporanges of my plants are unripe, but good spores are found
among the roots. In these the sculpture opens into a network.
Normal spores are found, but most of them bear evidence of having
grown singly or in pairs in the mother cells, as does Selaginella ru-
pestris. These are spherical or hemispherical, without commisures
and with no equatorial belt, or with it misplaced, often inclosing a
very small space at one end. The microspores also vary, some-
times reaching 44 p.

Deer brook beach, Jordan Pond, Mt. Desert Island, Maine,
Rand. Quoted in Redfield and Rand's Flora as Braunii. South
shore of Jordan Pond, September 10, 1894, Rand.

Type in the A. A. Eaton herbarium.


Formerly considered rare, this species appears to be quite com-
mon in New England, and any large pond with sandy shores may
be expected to yield it, especially if a little silt has been deposited
on the sand. It is usually found in "submersed pastures," the belt
where littoral vegetation extends beneath the surface of the water
for a short distance after the lowest stage of water has been reached.
I quite accidentally discovered in 1895 that this species was tri-lobed
in many instances, and some localities yield 25 per cent, of tri-lobed
plants. Baker (Fern Allies, p. 126) appears to be the first to men-
tion stomata in this species, though the fact was discovered by Mr.
Dodge and myself quite independently. Plants taken from inun-
dated situations usually have none, but when growing on the bor-
ders of ponds, a few leaves may usually be found which show them.

Distinguishable at sight by its very slender, spiral or recurved,
reddish leaves. Spores of northern specimens are much larger than
those from central Massachusetts, so far as seen.

At Pautuckaway Pond, Nottingham, N. H., and Kimball's
Pond, Amesbury, Mass., I find a few plants which in the field have
been taken for this species, but the leaves are larger, the sporangia
dark spotted, as much so as most specimens of Howellii. It ap-
pears to be a new species, but material is inadequate for descrip-

Mt. Desert, Rand, Fernald: Oldtown, Maine, Harvey. Com-

mon in Rockingham county, N. H., and known from several ponds
in Essex county, and near Boston, Mass., ; also from North Ston-
ington, and Ledyard, Conn., Graves.

3a. I. Tuckermani borealis n. var.

Trunk bi-lobed, leaves 1020, 3-8cm. long, imm. thick, slightly
recurved at tip : sporanges sometimes showing a few spots ; mac-
rospores 600-783 /u, sculptured as in the species, but the markings

This might perhaps as well be considered a variety of lacustris,
but the smaller leaves, wider velum, spotted sporanges and reticu-
lated spores, are characters enough to separate it, even if that spe-
cies were not so extremely rare.

Somes stream, Mt. Desert, August 13, 1887, Kennedy; pond
north of Long Pond, Mt. Desert, September 22, 1892 (with Harveyi
and Tuckermani } Fernald; Somes stream, September 30, 1893,
Rand; Kennebago lake (leaves only 2.5-5011. long), January 12,
1895, Coville, no. 78 ; Somes stream, Mt. Desert, September 14,
1895, Rand; Pushaw Pond, Oldtown, Maine, August 21, 1899,
Hervey. I also occasionally find it in Lamprey river, at Epping,
New Hampshire.

Type in the herbarium of A. A. Eaton ; co-types in the U. S.
National, and University of Minnesota herbaria.

4. I. hieroglyphica n. sp.

Aspect of Tuckermani : trunk bi-lobed : leaves 10-20, 6-7. 5011.
long, recurved, i-2mm. in diameter, blunt at tip : velum ^ indusi-
ate, sporangium unspotted ; macrospores 486-590 /a, polished, cov-
ered with vermiform, subconfluent and somewhat reticulated ridges,
becoming naked next the equator ; microspores 31-44 v, averaging
39 p, distinctly verrucose.

The spores are unique in appearance ; the ridges are very bold,
and the rest of the surface unmarked. I have seen nothing like it
from any quarter, but Motelay's illustration oiflaccida spores gives
a fair idea of their appearance, though utterly unlike the species he
intends to represent.

St. Francis lakes, Maine, type, Pringle; also Fernald from the
same place ; Moosehead lake, Harvey; Rangeley lakes, Coville.

Type in the herbarium of A. A. Eaton ; co-types in the U. S.
National, and the University of Minnesota herbaria.

5. I. Harveyi n. sp.

Trunk deeply 2- or occasionally 3-lobed, 1.6-3 cm - in diameter :
leaves 50-140, short (5-6cm.), very stout and fleshy, 2.5-3111111. in di-
ameter, strongly recurved, with an abrupt sharp point, purple-bronze
in color, often reddish in drying, without bast or stomata : ligula

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Online LibraryAmerican Fern SocietyFernwort papers presented at a meeting of fern students, held in New York city, June 27, 1900, under the auspices of the Linnaean fern chapter ... → online text (page 1 of 5)