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THE NORTH AMERICAN SLIME-MOULDS
The Macmillan Company
New York . Boston . Chicago . Dallas
Atlanta . San Francisco
Macmillan & Co., Limited
London . Bombay . Calcutta
Melbourne
The Macmillan Co. of Canada, Ltd.
Toronto
[Illustration: PHYSARUM NOTABILE (Enlarged one half)
In the field; sporangia in varied magnification, due to inequality in
background.]
THE NORTH AMERICAN SLIME-MOULDS
A Descriptive List of
All Species of Myxomycetes
Hitherto Reported from the Continent of
North America
With Notes on Some Extra-Limital Species
by
THOMAS H. MACBRIDE
State University of Iowa
New and Revised Edition
New York
The Macmillan Company
London: Macmillan and Co., Ltd.
1922
All rights reserved
Copyright, 1899,
By The Macmillan Company.
Copyright, 1922,
By The Macmillan Company.
The Clio Press
Iowa City, Iowa, U. S. A.
. IN . MEMORIAM .
. SAMUELIS . CALVINI .
. SCIENTIAE . NATURALIS . IN . UNIVERSITATE . IOWENSI .
. NUPER . PROFESSORIS .
. PRAECEPTORIS . COMITIS . AMICI .
. HUNC . LIBRUM .
. GRATO . ANIMO . DEDICAT .
. DISCIPULUS .
"Ihr naht euch wieder schwankende Gestalten,
Die frueh sich einst dem trueben Blick gezeigt."
GOETHE.
"Diese Kinder der Natur, welche aus einer ungeformten Gallert, und einem
unsichtbaren Saamen entstehen, sind im stande, in dem sie sich nach und
nach entwickeln und ihre scheinbar nachlaessige Bildung genau beobachten
(lassen), eben so sehr als die schoenste Pflanze, einem empfindenden
Herzen die tiefe Achtung und das paradiesische Vernuegen zu verschaffen,
welches einzig die Betrachtung der Heere der Natur und ihre
gleichbleibende Erhaltung durch eine ewige Kraft hervorbringen kann."
A. J. G. C. BATSCH 1783.
TABLE OF CONTENTS
PAGE
PREFACE ix
PREFACE TO SECOND EDITION xiii
BIBLIOGRAPHY xv
INTRODUCTORY 1
THE MYXOMYCETES 17
ADDENDA 282
INDEX OF GENERA 289
INDEX OF SPECIES 290
PLATES, WITH EXPLANATIONS 301
CORRIGENDA
The indulgent student will please notice the following for the new
edition _North American Slime Moulds_--
On p. 63, No. 17, read _Physarum megalosporum_ Macbr. Last line should
read 1917 Physarum _melanospermum_ Sturgis, _Mycologia_, Vol. IX, p.
323.
On p. 67, last line but one, at the end, read, p. 323.
On p. 67, insert just before No. 23, Vicinity of
Philadelphia,--_Bilgram_.
On p. 327, Plate XIII, lacks numbers. These may readily be supplied by
consulting descriptive text.
On p. 344, in explanation figure 2, last word read hour.
On p. 346, for name of species read _Fuligo rufa_ Pers., p. 28.
PREFACE TO THE FIRST EDITION[1]
The present work has grown out of a monograph entitled _Myxomycetes of
Eastern Iowa_, published by the present author about eight years ago.
The original work was intended chiefly for the use of the author's own
pupils; but interest in the subject proved much wider than had been
supposed, and a rather large edition of that little work was
speedily exhausted. At that time literature on the subject in
question--literature accessible to English readers--was scant indeed.
Cooke's translation of Rostafinski, in so far as concerned the species
of Great Britain, was practically all there was to be consulted in
English.
In 1892 appeared in London Massee's _Monograph of the Myxogastres_, and
two years later in the same world's centre the trustees of the British
Museum brought out Lister's _Mycetozoa_. Although these two English
works both claim revision of the entire group under discussion, the
latter paying special attention to American forms, nevertheless there
still seems place for a less pretentious volume which for American
students shall present succinct descriptions of North American species
only. The material basis of the present work consists of collections now
in the herbarium of the State University of Iowa. In accumulating the
material the author has had the generous assistance of botanists in all
parts of the country, from Alaska to Panama, and the geographical
distribution is in most cases authenticated by specimens from the
localities named. The descriptions, in case of species represented in
Europe, are based upon those of European authors; for forms first
described in this country, the original descriptions have been
consulted. A bibliography follows this preface.
In reference to the omnipresent vexed question of nomenclature, a word
is perhaps necessary. De Candolle's rule, "The first authentic specific
name published under the genus in which the species now stands," may be
true philosophy, but it is certainly an open question how that rule
shall be applied. If an author recognized and defined a given species in
times past, and, in accordance with views then held, assigned the
species to a particular genus, common honesty, it would seem, would
require that his work be recognized. To assume that any later writer who
may choose to set to familiar genera limits unknown before shall thereby
be empowered to write all species so displaced his own, as if, forsooth,
now for the first time in the history of science published or described,
is not only absolutely and inexcusably misleading, but actually
increases by just so much the amount of _debris_ with which the taxonomy
of the subject is already cumbered.
In face of a work so painstaking and voluminous as that of Rostafinski,
and in view of the almost universal confusion that preceded him, it
would seem idle to change for reasons purely technical the nomenclature
which the Polish author has established. Especially is this true in the
case of organisms so very perishable and fragile as those now in
question where comparative revision is apt to result in uncertainty. We
had preferred to leave the Rostafinskian, _i. e._ the heretofore current
nomenclature, untouched; but since other writers have preferred to do
otherwise, we are compelled to recognize the resultant confusion.
Slime-moulds have long attracted the attention of the student of nature.
For nearly two hundred years they find place more or less definite in
botanical literature. Micheli, 1729, figures a number of them, some so
accurately that the identity of the species is hardly to be questioned.
Other early writers are Buxbaum and Dillenius. But the great names
before Rostafinski are Schrader, Persoon, and Fries. Schrader's judgment
was especially clear. In his _Nova Genera_, 1797, he recognizes plainly
the difference between slime-moulds and everything else that passed by
the name of fungus, and proposed that they should be set off in a family
by themselves,[2] but he suggested no definite name. Nees (C. G.) also
made the same observation in 1817, and proposed the name _Aerogastres_;
but he cites as type of his aerogastres, _Eurotium_, and includes so
many fungi, that it seems unsafe now to approve his nomenclature.
Schrader also has left an excellent account of the cribrarias, the basis
of all that has since been attempted in that genus.
Persoon, in his _Synopsis_, 1801, attempts a review of all the fungi
known up to that time. His notes and synonymy are invaluable, enabling
us to understand the references of many of the earlier authors where
these had otherwise been indefinite if not unintelligible. He makes a
great many changes in nomenclature, and excuses himself on the ground
that he follows, in this particular, illustrious examples!
Unfortunately, so do we all!
Fries, in his _Systema Mycologicum_, 1829, summed up in most wonderful
way the work of all his predecessors and the mycologic science of his
time. In reading Fries the modern student hardly knows which most to
admire, the author's far-reaching, patient research, the singular acumen
of his taxonomic instinct, the graceful exactness of the Latin in which
his conclusions are expressed, or the delicate courtesy with which he
touches the work even the most primitive, of those his predecessors or
contemporaries. Nevertheless in our particular group even the
determinations of Fries are not conclusive. He himself often confesses
as much. The microscopic technique of that day did not yield the data
needful for minute comparison among these most delicate forms.
It remained for DeBary and Rostafinski to introduce a new factor into
the description of species, and by spore-measurement and the delineation
of microscopic detail to supply an element of definiteness which has no
parallel in the work of any earlier student of this group. Under these
conditions the revision undertaken by Rostafinski was of a most heroic
sort. His work was almost a new beginning; and while in nomenclature he
was inclined to follow the Paris Code, yet the inadequacy of the earlier
descriptions often made such a course impracticable. The synonymy of
Rostafinski is largely that of Fries, and upon this the Polish author
attempts to apply the law of priority. In the historical note, _wzmianka
historyczna_, accompanying the description of each specific form, he
generally states the reason for the nomenclature he adopts, whether
selected from the mass of supposed synonymy or introduced by himself _de
novo_. Unfortunately, Rostafinski is sometimes purely arbitrary in his
selections. He sometimes changes a specific or even generic name,
otherwise correctly applied, simply because in primary etymological
significance the name seems to him inappropriate. In such cases it is
proper to restore the earlier name. Nevertheless Rostafinski is still
our most trustworthy guide.
Of course, where later investigations have served to obliterate the
once-thought patent distinctions between supposed genera or species, it
is proper to unite such forms under the older determinable titles and
this we have attempted. But wherever in the present work a name has been
changed, the name of the earlier author will be found in parenthesis,
followed immediately by that of him who made the change, and in general,
recent practice, especially as expressed in the rules of the various
codes, has determined the puzzling questions of nomenclature.
In justification of the use of _Myxomycetes_ as a general title it may
be said that in this case prevalent usage is not inconsistent with a
rational application of the rules of priority. The Friesian designation
_Myxogastres_ was applied by its author in 1829 to the endosporous
slime-moulds as a section of gasteromycetous fungi. Four years later
Link, perceiving more clearly the absolute distinctness of the group,
substituted the name _Myxomycetes_. In the same year Wallroth adopted
the same designation, but strangely confused the limitations of the
group he named. Wallroth seems to have thought _Myxomycetes_ a synonym
for _Gasteromycetes_ Fries. In 1858 DeBary applied the title _Mycetozoa_
to a group which included the then lately discovered _Acrasieae_ with
the true slime-moulds, both endosporous and exosporous. For all except
the _Acrasieae_ DeBary retained the old appellation, Myxomycetes.
Rostafinski adopted DeBary's general name, but changed its application.
As it has been shown, since DeBary's time, that the _Acrasieae_[3] have
no true plasmodium, and are therefore not properly, or at least not
necessarily, associated with the slime-moulds, there appears no
necessity for the term _Mycetozoa_, and the question lies between
_Myxogastres_ and _Myxomycetes_. Of these two names the former, as we
have seen, has undoubted priority, but only as applied to the
endosporous species. The same thing was true of Link's designation until
DeBary redefined it, but having been taken up by DeBary, redefined and
correctly applied, Myxomycetes (Link) DeBary must remain the undisputed
title for all true slime-moulds, endosporous and exosporous alike.
In arranging the larger divisions of the group the scheme of Rostafinski
has been somewhat modified in order to give expression to what the
present author deems a more natural sequence of species. The highest
expression of myxomycetan fructification is doubtless the isolated
sporangium with its capillitium. This is reached by successive
differentiations from the simple plasmodium. The aethalium may be
esteemed in some instances a case of degeneration, in others of arrested
development. In any event in the present arrangement, aethalioid forms
are first disposed of, leaving the sporangiate species to follow from
plasmodiocarpous as directly as may be.
The artificial keys herewith presented proceed on the same plan and are
to be taken, as such keys always are, not as definitive in any case, but
simply as an aid to help the student more speedily to reach a probably
satisfactory description.
FOOTNOTES:
[1] _The North American Slime Moulds_, 1899.
[2] Schrader, _Nova Plantarum Genera_, 1797, pp. vi-vii.
[3] Cf. Edgar W. Olive, _Monograph of the Acrasieae_; Boston, 1902.
PREFACE TO THE SECOND EDITION
The first edition of this little book having been exhausted long ago,
the writer in this second issue takes opportunity to correct sundry
errata, typographical and other, and at the same time to incorporate
such new information in reference to individual species and to the
subject entire as the researches of more recent years may afford.
To Miss Gulielma Lister, of London, the writer expresses his sense of
deep obligation for much assistance in settling difficult matters of
nomenclature and identification; it will be found as a result that in
most instances the same thing in the two volumes, English and American,
appears under the same name. There are still differences; these result
in most cases from different points of view, different estimates or
emphasis of characteristics in these ever elusive objects.
To Professor Torrend, formerly of Lisbon, the writer is indebted for a
set of European types, and to Professor Bethel, pathologist of Denver,
for rich material from the fertile mountains of Colorado and California.
To Professor Morton Peck, of Oregon, we are indebted for many notes of
the color of plasmodia and for collections of Pacific coast forms. Mr.
Bilgram, of Philadelphia, read the manuscript of the genus _Physarum_
and has contributed many rare species. To Dr. Sturgis, of Massachusetts,
we are indebted for material from both east and west.
The present volume is intended especially for American readers and is
accordingly particularly devoted to a discussion of species so far
reported on the western continent; nevertheless it has seemed wise to
include a brief description of some other forms as well, and reference
to many extra-limital species now generally recognized will be found
here and there in connection with the more extended treatment of related
American forms.
_February twenty-eight, 1921._
At the last moment, nearly all plates and drawings of the first
edition disappeared! necessitating a quick renewal of drawings
and plates. This may in part explain lack of uniformity, and
various minor irregularities sure to grieve the intelligent
student.
BIBLIOGRAPHY
The following are the principal works consulted in the prosecution of
the investigations here recorded:--
1763. Adanson, M. Familles des Plantes.
1805. Albertini--see under Schweinitz.
1841. Annals and Magazine of Natural History. London, various volumes:
1841, Ser. I., vol. vi.; 1850, Ser. II., vol. v.
1887. Annals of Botany, vols. i-xxxi.
1783. Batsch, A. J. G. C. Elenchus Fungorum; with Continuatio I. 1786;
Continuatio II. 1789.
1775. Battara, A. Fungorum Agri Arimensis Historia.
1860. Berkeley, M. J. Outlines of Fungology.
1789. Bolton, J. History of Funguses about Halifax.
1851. Bonorden, H. F. Mycologie.
1875. Botanical Gazette, The. Various volumes to 1921.
1843. Botanische Zeitung. Various volumes to 1898.
1892. Bulletin Laboratories Nat. Hist. Iowa, vol. ii.
1873. Bulletin Torrey Botanical Club. Various volumes to 1898.
1791. Bulliard, P. Histoire des Champignons de la France.
1721. Buxbaum, J. C. Enumeratio Plantarum.
1863. Cienkowski, L. Zur Entwickelungsgeschichte der Myxomyceten.
1893. Celakowsky, L. Die Myxomyceten Boehmens.
1871. Cooke, M. C. Handbook of British Fungi.
1877. Cooke, M. C. Myxomycetes of Great Britain.
1877. Cooke, M. C. Myxomycetes of the United States.
1837. Corda, A. I. C. Icones Fungorum.
1854. Currey, F., in Quart. Journal Microscopical Science.
1848. Curtis, M. A. Contributions to the Mycology of North America; Am.
Journal of Science and Arts.
1859. De Bary, A. H. Die Mycetozoen.
1866. De Bary, A. H. Morphologie der Pilze, Mycetozoen und Bacterien.
1802. De Candolle, A. P. Flore Francaise.
1719. Dillenius, J. J. Catalogus Plantarum circa Cissam nascentium.
1813. Ditmar, L. P. F., Sturm, Deutschlands Flora, 3te Abtheil; Die
Pilze Deutschlands.
1878. Ellis, J. B. North American Fungi. _Exsiccati. et seq._
1818. Ehrenberg, C. G. Sylvae Mycologicae Berolinenses.
1761. Flora, Danica, vol. i.; also vols. iii. iv. v.
1817. Fries, Elias M. Symbolae Gasteromycetum.
1818. Fries, Elias M. Observationes Mycologicae.
1829. Fries, Elias M. Systema Mycologicum.
1873. Fuckel, I. Symbolae Mycologicae.
1791. Gmelin, C. C. Systema Naturae, Tom. II., Pars. ii.
1823. Greville, R. K. Scottish Cryptogamic Flora.
1872. Grevillea, various volumes to 1897.
1751. Hill, Sir John. A History of Plants.
1795. Hoffman, G. C. Deutschlands Flora.
1773. Jacquin, N. I. Miscellanea Austriaca.
1885. Journal of Mycology and _seq._
1878. Karsten, Mycologia Fennica.
1809. Link, H. F. Nova Plantarum Genera.
1753. Linne, C. Systema Naturae--to 1767.
1894. Lister, Arthur. The Mycetozoa; 1911, Second Edition, revised by
Gulielma Lister.
1892. Massee, George. Monograph of the Myxogastres.
1729. Micheli, P. A. Nova Plantarum Genera.
1892. Morgan, A. P. Myxomycetes of the Miami Valley--to 1895.
1816. Nees, Ch. G. D. Das System der Pilze und Schwamme.
1837. Nees, T. F. L. et A. Henry. Das System der Pilze.
1869. Peck, Charles H. Reports N. Y. State Museum Nat. History--to 1898.
1795. Persoon, C. H. Observationes Mycologicae, Pars prima.
1799. Persoon, C. H. Observationes Mycologicae, Pars secunda.
1797. Persoon, C. H. Tentamen Dispositionis Methodicae Fungorum.
1801. Persoon, C. H. Synopsis Methodica Fungorum.
1844. Rabenhorst, L. Deutschland's Kryptogamenflora.
1884. Raciborski, M. Myxomycetes Agri Krakov. Genera, Species et
Varietates novae.
1888. Raunkiaer, C. Myxomycetes Daniae.
1769. Retzius, A. J. In Handlungen, Kon. Svensk. Vet. Acad.
1890. Rex, George A. In Proceedings Philad. Acad. of Nat. Sciences--to
1893.
1873. Rostafinski, J. Versuch eines Systems der Mycetozoen.
1875. Rostafinski, J. Sluzowce Monografia.
1778. Roth, A. W. Tentamen Florae Germanicae.
1888. Saccardo, P. A. Sylloge Fungorum, vol. vii., _et seq._
1841. Sauter, A. Flora, vol. xxiv., p. 316.
1762. Schaeffer, J. C. Fungi qui in Bav. et Pal. nascuntur--to 1774.
1797. Schrader, H. A. Nova Genera Plantarum.
1890. Schroeter, J. Myxomycetes, in Engler u. Prantl Pflanzenfamilien.
1885. Schroeter, J. Kryptogamenflora von Schlesien, die Pilze.
1801. Schumacher, C. F. Enumeratio Plant. Saell. crescentium.
1805. Albertini, I. and Schweinitz, L. D. de. Conspectus Fungorum.
1822. Schweinitz, L. D. de. Synopsis Fungorum Car. Sup.
1834. Schweinitz, L. D. de. Synopsis Fungorum in America Boreali.
1797. Sowerby, J. English Fungi--to 1809; 3 vols.
1760. Scopoli, J. A. Flora Carniolica--to 1772.
1797. Trentepohl, K. Observations Botanicae,--to Roth, Catalecta
Botanica, Fasc. i.
1833. Wallroth, C. F. Flora Cryptogamica Germaniae.
1787. Willdenow, K. L. Florae Berolinensis Prodromus.
1886. Wingate, Harold, Jour. Mycol. ii., 125.
1889. Wingate Harold, In Proc. Acad. Nat. Sci. Philad.
1890. Wingate, Harold--in Revue Mycologique.
1873. Woronin u. Famintzin, Ueber Zwei neuen Formen von Schleimpilzen.
1885. Zopf, W. Die Pilzthiere oder Schleimpilze.
To these may be added the many contributions on the general subject, as
these are found in all sorts of current botanical literature; cited
everywhere in this volume as occasion offered.
INTRODUCTORY
The Myxomycetes, or slime-moulds, include certain very delicate and
extremely beautiful fungus-like organisms common in all the moist and
wooded regions of the earth. Deriving sustenance, as they for the most
part do, in connection with the decomposition of organic matter, they
are usually to be found upon or near decaying logs, sticks, leaves, and
other masses of vegetable detritus, wherever the quantity of such
material is sufficient to insure continuous moisture. In fruit, however,
as will appear hereafter, slime-moulds may occur on objects of any and
every sort. Their minuteness retires them from ordinary ken; but such is
the extreme beauty of their microscopic structure, such the exceeding
interest of their life-history, that for many years enthusiastic
students have found the group one of peculiar fascination, in some
respects, at least, the most interesting and remarkable that falls
beneath our lens.
The slime-mould presents in the course of its life-history two very
distinct phases: the _vegetative_, or growing, assimilating phase, and
the _reproductive_. The former is in many cases inconspicuous and
therefore unobserved; the latter generally receives more or less
attention at the hands of the collector of fungi. The vegetative phase
differs from the corresponding phase of all other plants in that it
exhibits extreme simplicity of structure, if structure that may be
called which consists of a simple mass of protoplasm destitute of
cell-walls, protean in form and amoeboid in its movements. This phase
of the slime-mould is described as plasmodial and it is proper to
designate the vegetative phase in any species, as the _plasmodium_ of
the species. It was formerly taught that the plasmodium is unicellular,
but more recent investigation has shown that the plasmodial protoplasm
is not only multinuclear but karyokinetic; its cells divide and
redivide, as do the _reproductive_ cells of plants and animals
generally. Nevertheless, in its plasmodial phase, the slime-mould is
hardly to be distinguished from any other protoplasmic mass, may be
compared to a giant amoeba, and justifies in so far the views of those
systematists who would remove the slime-moulds from the domain of the
botanist altogether, and call them animals. The plasmodium is often
quite large. It may frequently be found covering with manifold
ramifications and net-like sheets the surface of some convenient
substratum for the space of several square feet.
The substance of the plasmodium has about the consistency of the white
of an egg; is slippery to the touch, tasteless, and odorless. Plasmodia
vary in color in different species and at different times in the same
species. The prevailing color is yellow, but may be brown, orange, red,
ruby-red, violet, in fact any tint, even green. Young plasmodia in
certain species are colorless (as in _Diderma floriforme_), while many
have a peculiar ecru-white or creamy tint difficult to define. Not only
does the color change, sometimes more than once in the course of the
life history of the same species, but it may be the same for several
forms, which in fruit are singularly diverse indeed, so that the mere
color of the plasmodium brings small assistance to the systematist. In
fact, the color depends no doubt upon the presence in the plasmodium of
various matters, more or less foreign, unassimilated, possibly some of
them excretory, differing from day to day.
In its plasmodial state, as has been said, the slime-mould affects damp
or moist situations, and during warm weather in such places spreads over
all moist surfaces, creeps through the interstices of the rotting bark,
spreads between the cells, between the growth-layers of the wood, runs
in corded vein-like nets between the wood and bark, and finds in all
these cases nutrition in the products of organic decomposition. Such a
plasmodium may be divided, and so long as suitable surroundings are
maintained, each part will manifest all the properties of the whole.
Parts of the same plasmodium will even coalesce again. If a piece of
plasmodium-bearing wood be brought indoors, be protected from
desiccation by aid of a moist dark chamber, not too warm (70 deg. F.), the
organism seems to suffer little if any injury, but will continue for
days or weeks to manifest all the phenomena of living matter. Thus,
under such circumstances, the plasmodium will constantly change shape
and position, can be induced to spread over a plate of moist glass, and
so be transferred to the stage of a microscope, there to exhibit in the
richest and most interesting and abundant fashion the streaming
protoplasmic currents. As just indicated, the plasmodia follow moisture,
creep from one moist substance to another, especially follow nutritive
substrata. They seem also to secure in some way exclusive possession. I
have never seen them interfered with by hyphae or enemies of any sort,
nor do they seem to interfere with one another. Plasmodia of two common
species, _Hemitrichia clavata_ and _H. vesparium_ are often side by side
on the same substratum, but do not mix, and their perfected fruits
presently stand erect side by side, each with its own characteristics,
entirely unaffected by the presence of the other. On the other hand, it
is probable that some of the forms which, judged by their different
fructifications, and by this alone, are to us distinct, may be more
closely related than we suspect, and puzzling phases which show the
distinctive marks supposed to characterize different species are no
doubt sometimes to be explained on the theory of plasmodial crossing;
they are hybrids.
Under certain conditions, low temperature, lack of moisture, the
plasmodium may pass into a resting phase, when it masses itself in heaps
and may become quite dry in lumps of considerable size, and so await the
return of favorable conditions when former activity is quickly resumed.
Sometimes the larger plasmodia pass into the resting phase by undergoing
a very peculiar change of structure. In ordinary circumstances the
abundant free nuclei demonstrable in the plasmodium afford the only
evidence of cellular organization. In passing now into the condition of
rest, the whole protoplasmic mass separates simultaneously into numerous
definite polyhedral or parenchymatous cells, each with a well-developed
_cellulose wall_.[4] When the conditions essential to activity are
restored, the walls disappear, the cellulose is resorbed, and the
plasmodium resumes its usual habit and structure.
The plasmodial phase of the slime-mould, like the hyphal phase of the
fungus, may continue a long time; for months, possibly for years. The
reason for making the latter statement will presently appear. But
however long or short the plasmodial phase continue, the time of fruit,
the reproductive phase, at length arrives. When this time comes, induced
partly by a certain maturity in the organism itself, partly no doubt by
the trend of external conditions, the plasmodium no longer as before
evades the light, but pushes to the surface, and appears usually in some
elevated or exposed position, the upper side of the log, the top of the
stump, the upper surface of its habitat, whatever that may be; or even
leaves its nutrient base entirely and finds lodging on some neighboring
object. In such emergency the stems and leaves of flowering plants are
often made to serve, and even fruits and flowers afford convenient
resting places. The object now to be attained is not the formation of
fruit alone, but likewise its speedy desiccation and the prompt
dispersal of the perfected spores. Nothing can be more interesting than
to watch the slime-mould as its plasmodium accomplishes this its last
migration. If hitherto its habitat has been the soft interior of a
rotten log, it now begins to ooze out in all directions, to well up
through the crevices of the bark as if pushed by some energy acting in
the rear, to stream down upon the ground, to flow in a hundred tiny
streams over all the region round about, to climb all stems, ascend all
branches, to the height of many inches, all to pass suddenly as if by
magic charm into one widespread, dusty field of flying spores. Or, to be
more exact, whatever the position ultimately assumed, the plasmodium
soon becomes quiescent, takes on definite and ultimate shape, which
varies greatly, almost for each species. Thus it may simply form a flat,
cake-like mass, _aethalium_, internally divided into an indefinite
number of ill-defined spore cases, sporangia; or the plasmodium may take
the form of a simple net, _plasmodiocarp_, whose cords stand out like
swollen veins, whose meshes vary both in form and size; or more commonly
the whole protoplasmic mass breaks up into little spheroidal heaps which
may be sessile directly on the substratum, or may be lifted on tiny
stems, stipitate, which may rest in turn upon a common sheet-like film,
or more or less continuous net, spreading beneath them all, the
_hypothallus_. In any case, each differentiated portion of the
plasmodium, portion poorly or well defined, elongate, net-like,
spheroidal, elliptical, or of whatever shape, becomes at length a
sporangium, spore-case, receptacle for the development and temporary
preservation of the spores.[5]
The slime-moulds were formerly classed with the gasteromycetous fungi,
puff-balls, and in description of their fruiting phase the terms
applicable to the description of a puff-ball are still employed,
although it will be understood that the structures described are not in
the two cases homologous; analogous only. The sporangium of the
slime-mould exhibits usually a distinct _peridium_, or outer limiting
wall, which is at first continuous, enclosing the spores and their
attendant machinery, but at length ruptures, irregularly as a rule, and
so suffers the contents to escape. The peridium may be double, varies in
texture, color, persistence, and so forth, as will be more fully set
forth in the several specific descriptions. The peridium blends with the
hypothallus below when such structure is recognizable, either directly,
when the sporangium is sessile, or by the intervention of a _stipe_. The
stipe may be hollow, may contain coloring matter of some sort, or may
even contain peculiar spore-like cells or spores; is often furrowed, and
in some cases shows a disposition to unite or blend with the stalks of
neighboring sporangia. In many cases the stipe is continued upward, more
or less definitely into the cavity of the sporangium, and there forms
the _columella_, sometimes simple and rounded, like the analogous
structure in the _Mucores_, sometimes as in _Comatricha_, branching
again and again in wonderful richness and complexity.
Each sporangium is at maturity filled with numerous unicellular spores.
These are usually spherical, sometimes flattened at various points by
mutual contact; they are of various colors, more commonly yellow or
violet brown, are sometimes smooth (?), but generally roughened either
by the presence of minute warts, or spines, or by the occurence of more
or less strongly elevated bands dividing reticulately the entire
surface. The spores are in all cases small 3-20 mu, and reveal their
surface characters only under the most excellent lenses.
Associated with the spores in the sporangium occurs the _capillitium_.
This consists of most delicate thread-or hair-like elements, offering
great variety both in form and structure. The threads composing the
capillitium are not to be regarded, even when free, as cells, nor even
of cellular origin; probably, as would appear from the researches of
Strasburger and Harper, all forms of capillitial threads arise in
connection with vacuoles in the protoplasmic mass. "Whether the thread
is hollow or solid, simple or branched, free or connected with the
peridium or a columella,--these are entirely secondary conditions,
depending on the extent and form of the vacuoles."[6] They may occur
singly or be combined into a net, they may be terete or flat, attached
to the peridial wall or free, simple or adorned with bands or spires and
knobs in every variety, uniform or profusely knotted and thickened at
intervals, and burdened with calcic particles. In many cases, the
capillitium contributes materially to the dispersal of the spores; in
others, it doubtless contributes mechanically to the support of the
peridial wall, and renders so far persistent the delicate sporangium.
For more exact description the reader is again referred to the specific
delineations which follow.
The transition from phase to phase requires, as intimated, no great
length of time. _Tilmadoche polycephala_ completed the transition from
vegetative to fruiting phase in less than twelve hours.
The germination of the spores ensues closely upon their dispersal or
maturity and is unique in many respects.[7] The wall of the spore is
ruptured and the protoplasmic content escapes as a zoospore
indistinguishable so far from an amoeba, or from the zoospore of our
chytridiaceous fungi. This amoeboid zoospore is without cell-wall,
changes its outline, and moves slowly by creeping or flowing from point
to point. At this stage many of the spores assume each a flagellate
cilium, and so acquire power of more rapid locomotion. The zoospores,
whether ciliate or not, thus enjoy independent existence and are capable
of continuing such existence for some time, assimilating, growing, and
even reproducing themselves by simple fission, over and over again. This
takes place, of course, only in the presence of suitable nutrient media.
Nevertheless the spores of many species germinate quickly simply in
water, and a drop suspended in the form of the ordinary drop-culture on
a cover-glass affords ample opportunity. In the course of time, usually
not more than two or three days, the swarm spores cease their activity,
lose their cilia, and come to rest, exhibiting at most nothing more than
the slow amoeboid movement already referred to. In the course of two
or three days more, in favorable cases, the little spores begin to
assemble and flow together; at first into small aggregations, then
larger, until at length all have blended in one creeping protoplasmic
mass to form thus once again the plasmodium, or plasmodial phase with
which the round began. Small plasmodia may generally be thus obtained
artificially from drop-cultures. Such, however, in the experience of the
writer, are with difficulty kept alive. Hay infusions, infusions of
rotten wood, etc., may sometimes for a time give excellent results.
The spores of _Didymium crustaceum_ were sown upon a heap of leaves in
autumn. An abundant display of the same species followed in the next
June; but, of course, the intervening phases were not observed. The most
satisfactory studies are obtained by plasmodia carefully brought in
directly from the field. A plasmodium that appeared suddenly and passed
to fruit on agar in a petri dish offers a valuable suggestion for
further research.
With such a life-history as that thus briefly sketched, it is small
wonder that the taxonomic place of the slime-moulds is a matter of
uncertainty, not to say perplexity. So long as men studied the ripened
fruit, the sporangia and the spores, with the marvellous capillitium,
there seemed little difficulty; the myxomycetes were fungi, related to
the puff-balls, and in fact to be classed in the same natural order. The
synonymy of some of the more noticeable species affords a very
interesting epitome of the history of scientific thought in this
particular field of investigation. Thus the first described slime-mould
identifiable by its description is Lycogala epidendrum (Buxbaum) Fries,
the most puff-ball looking of the whole series. Ray, in 1690, called
this _Fungus coccineus_. In 1718, Ruppinus described the same thing as
_Lycoperdon sanguineum_; Dillenius at about the same time, as _Bovista
miniata_; and it was not until 1729, that Micheli so far appreciated the
structure of the little puff-ball as to give it a definite, independent,
generic place and title, _Lycogala globosum_ ..., etc.[8]
But Micheli's light was too strong for his generation. As Fries, one
hundred years later quaintly says, ... "immortalis Micheli tam claram
lucem accendit ut succesores proximi eam ne ferre quidem potuerint."
Notwithstanding Micheli's clear distinctions, he was entirely
disregarded, and our little Lycogala was dubbed _Lycoperdon_ and _Mucor_
down to the end of the century; and so it was not till 1790 that Persoon
comes around to the standpoint of Micheli and writes _Lycogala miniata_.
Fries himself, reviewing the labors of his predecessors all, grouped the
slime-moulds as a sub-order of the gasteromycetes and gave expression to
his view of their nature and position when he named the sub-order
_Myxogastres_. In 1833, Link, having more prominently in mind the
minuteness of most of the species collocated by Fries, and perceiving
perhaps more clearly even than the great mycologist the entire
independence of the group, suggested as a substitute for the sub-order
_Myxogastres_, the order _Myxomycetes, slime-moulds_. Link's decision
passed unchallenged for nearly thirty years. The slime-moulds were set
apart by themselves; they were fungi without question and, of course,
plants.
If the hypha is the morphological test of a fungus, then it is plain
that the slime-moulds are not fungi. No myxomycete has hyphae, nor indeed
anything at all of the kind. Nevertheless, there are certain parasitic
fungi, _Chytridiaceae_ for example, whose relationships plainly entitle
them to a place among the hyphate forms that have no hyphae whatever in
the entire round of their life-history. These are, however, exceptional
cases and really do not bear very closely on the question at issue.
Physiologically, the fungi are incapable of independent existence, being
destitute of chlorophyl. In this respect the slime-moulds are like the
fungi; they are nearly all saprophytes and absolutely destitute of
chlorophyl. Unfortunately this physiological character is identically
that one which the fungi share with the whole animal world, so that the
startling inquiry instantly rises, are the slime-moulds plants at all?
Are they not animals? Do not their amoeboid spores and plasmodia ally
them at once to the amoeba and his congeners, to all the monad,
rhizopodal world? This is the position suggested by DeBary in 1858, and
adopted since by many distinguished authorities, among whom may be
mentioned Saville Kent, of England, and Dr. William Zopf, of Germany, in
_Die Pilzthiere_, 1885. Rostafinski was a pupil of DeBary's. However,
his volume on the slime-moulds was written after leaving the laboratory;
and no doubt with the suggestion of his master still before his mind, he
adopts the title Mycetozoa, as indicating a closer relationship with the
animal world, but our leading authority really has little to say in
regard to the matter.[9]
Dr. Schroeter, a recent writer on the subject, after showing the
probable connection between the phycochromaceous Algae and the simplest
colorless forms, namely, the _Schizomycetes_, goes on to remark: "At the
same point where the Schizomycetous series take rise, there begin
certain other lines of development among the most diminutive
protoplasmic masses.... Through the amoebae one of these lines gives
rise on the one hand to rhizopods and sponges in the animal kingdom, on
the other to the _Myxomycetes_ among the fungi." This ranges the
Myxomycetes, in origin at least, near the _Schizomycetes_.
The brilliant studies of Dr. Thaxter, resulting in the discovery and
recognition of a new group, a new order of the schizomycetes, strikingly
confirm the judgment of Schroeter.[10] Here we have forms that strangely
unite characteristics of both the groups in question. If on the one hand
the _Myxobacteria_ are certainly schizomycetes, on the other they just
as certainly offer in their developmental history "phenomena closely
resembling those presented by plasmodia or pseudo-plasmodia...." Now the
schizophytes certainly pass by gradations easy to the filamentous algae,
and so to relationship with the plants, and the discovery of the
_Myxobacteriacae_, brings the myxomycetes very near the vegetable
kingdom if not within it.
All authorities agree that the myxomycetes have no connection in the
direction of upward development, "keinen Anschluss nach oben," if then
their only relationship with other organisms is to be found at the
bottom (centre) of the series only, it is purely a matter of
indifference whether we say plant or animal, for at the only point where
there is connection there is no distinction.
But why call them either animals or plants? Was Nature then so poor that
forsooth only two lines of differentiation were at the beginning open
for her effort? May we not rather believe that life's tree may have
risen at first in hundreds of tentative trunks of which two have become
in the progress of the ages so far dominant as to entirely obscure less
progressive types? The Myxomycetes are independent; all that we may
attempt is to assert their near kinship with one or other of life's
great branches.
The cellulose of the slime-mould looks toward the world of plants. The
aerial fructification and stipitate habit of the higher forms tends in
the same direction. The disposition to attach themselves to some fixed
base is a curious characteristic of plants, more pronounced as we ascend
the scale; but by no means lacking in many of the simplest, diatoms,
filamentous algae, etc., and it is quite as reasonable to call a
vorticella, or a stentor, by virtue of his stipitate form and habit, a
plant as to call a slime-mould an animal because in one stage of its
history it resembles an amoeba. The total life of an organism in any
case must be taken into account.[11] At the outset plants and animals
are alike; there is no doubt about it; they differ in the course of
their life-histories. The plasmodium is the vegetative phase of the
slime-mould. It needs no cell-walls of cellulose, no more than do the
dividing cells of a lily-endosperm; both are nourished by organic food
and resort to walls only as conditions change. The possession of walls
is an indication of some maturity. In the slime-mould the assumption of
walls is indeed delayed. Walls at length appear and when they do come
they are like those of the lily; they are cellulose. The myxomycetes may
be regarded as a section of the organic world in which the forces of
heredity are at a maximum whatever those forces may be. Slime-moulds
have in smallest degree responded to the stimulus of environment. They
have, it is true, escaped the sea, the fresh waters in part, and become
adapted to habitation on dry land, but nothing more. It is instructive
to reflect that even in her most highly differentiated forms the channel
which Nature elects for the transmissal of all that heredity may bestow,
is naught else than a minute mass of naked protoplasm. Nature reverts,
we say, to her most ancient and simple phases, and heredity is still
consonant with apparent simplicity; apparent we say, for as becomes
increasingly evident, nothing that lives is simple!
The fact is the Myxomycetes constitute an exceedingly well-defined
group, and the question of relationship in any direction need not much
perplex the student. Least of all is the question to be settled by
anybody's dictum, which is apt to be positive inversely in proportion to
the speaker's acquaintance with the subject. No one test can be applied
as a universal touchstone to separate plants from animals. Such is
simply _petitio principii_. Nor is there any advantage at present
apparent in attempts to associate slime-moulds with other presumably
related groups. Saville Kent's effort to join them with the sponges was
not happy, and Dr. Zopf's association of the slime-moulds and monads
appears forced, at best; for when it comes to the consideration of the
former, their systematic and even morphological treatment, he is
compelled to deal with them by themselves under headings such as
"Eumycetozoen," "Hoehere Pilzthiere," etc. One rather commends the
discreetness of DeBary, whose painstaking investigations first called
attention to the uncertain position of the group. After reviewing the
results of all his labors DeBary does not quite relegate the
slime-moulds to the zoologist for further consideration, but simply
says:[12] "From naked amoeba, with which the Mycetozoa (=Myxomycetes)
are connected in ascending line, the zoologists with reason derive the
copiously and highly developed section of the shell-forming
Rhizopoda.... And since there are sufficient grounds for placing the
rhizopods outside the vegetable and in the animal kingdom, and this is
undoubtedly the true position for the amoebae, which are their earlier
and simpler forms, the Mycetozoa, which _may_ be directly derived from
the same stem, are at least brought very near to the domain of zoology."
Notwithstanding all the controversy in regard to the matter, the study
of the slime-moulds still rests chiefly with the botanists. A simple
phylogenetic scheme for thallophytes is offered in the Strasburger text
as follows:--
THALLOPHYTA
1. SCHIZOPHYTA
BACTERIA
CYANOPHYCEAE
2. FLAGELLATA
{ MYXOMYCETES
{ PERIDINEAE
_a_ { CONJUGATAE
{ HETEROCONTAE
{ CHLOROPHYCEAE
_b_ { CHARACEAE
3. RHODOPHYCEAE
4. FUNGI
About 500 species of slime-moulds have been described. Saccardo
enumerates 443, inclusive of those denominated doubtful or less
perfectly known. These 443 species are distributed among 47 genera, of
which 15 are represented by but a single species each,--monotypic. In
the United States there have been recognized about 300 species. Of those
here described, some are almost world-wide in their distribution, others
are limited to comparatively narrow boundaries. The greater number occur
in the temperate regions of the earth, although many are reported from
the tropics, and some even from the arctic zone. Schroeter found
_Physarum cinereum_ at North Cape. Our Iowa forms are much more numerous
in the eastern, that is, the wooded regions of the state. _Physarum
cinereum_ has however been taken on the untouched prairie, and on the
western deserts, as also _Physarum contextum_ on the decaying stem of
_Calamagrostis_, far from forest.
As to the economic importance of our myxomycetes, there is no long
chapter to write. Fries says: "Usu in vita communi parum admodum sese
commendant, sed in oeconomia naturae certe non spernendi. Multa
insectorum genera ex eorum sporidiis unica capiunt nutrimenta." However
this may be, there is one species which has come to light since Fries's
day which is the source of no inconsiderable mischief to the
agriculturist. _Plasmodiophora brassicae_ occasions the disease known as
"club-root" in cabbage, and has been often made the subject of
discussion in our agricultural and botanical journals.[13] Aside from
the injurious tendencies, possible or real, of the forms mentioned, I
know not that all other slime-moulds of all the world, taken all
together, affect in any slightest measure the hap or fortune of man or
nation. And yet, if in the economic relations of things, man's
intellectual life is to be considered, then surely come the uncertain
myxos, with their fascinating problems proffered still in forms of
unapproachable delicacy and beauty, not without inspiration.
COLLECTION AND CARE OF SLIME-MOULD MATERIAL
On this subject a word may here be appropriate. As just now intimated,
specimens may be taken at the appropriate season in almost any or every
locality. Beginning with the latter part of May or first of June, in the
Northern states, plasmodia are to be found everywhere on piles of
organic refuse: in the woods, especially about fallen and rotting logs,
undisturbed piles of leaves, beds of moss, stumps, by the seeping edge
of melting snow on mountain sides, by sedgy drain or swamp, nor less in
the open field where piles of straw or herbaceous matter of any sort
sinks in undisturbed decay. Within fifty years tree-planting in all the
prairie states has greatly extended the range of many more definitely
woodland species, so that species of _Stemonitis_, for instance, are
common in the groves on farms far into Nebraska and Dakota. In any
locality the plasmodia pass rapidly to fruit, but not infrequently a
plasmodium in June will be succeeded in the same place by others of the
same species, on and on, until the cold of approaching winter checks all
vital phenomena. The process of fruiting should be watched as far as
possible, and for herbarium material, allowed to pass to perfection in
the field.
Specimens collected should be placed immediately in boxes in such a way
as to suffer no injury in transport; beautiful material is often ruined
by lack of care on the part of the collector. Once at the herbarium,
specimens may be mounted by gluing the supporting material to the bottom
of a small box. Boxes of uniform size and depth may be secured for the
purpose. Some collectors prefer to fasten the specimen to a piece of
stiff paper, of a size to be pressed into the box snugly, but which may
be removed at pleasure. Every pains must in any case be taken to exclude
insects. Against such depredators occasional baking of the boxes on the
steam radiator in winter is found to be an efficient remedy.
For simple microscopic examination it will be found convenient to first
wet the material with alcohol on the slide, then with a weak solution of
potassic hydrate, to cause the spores and other structures to assume
proper plumpness. A little glycerine may be added or run under the cover
if it is desired to preserve the material for further or prolonged
study. For permanent mounting nothing in most cases is better than
glycerine jelly. As a preparation, the material should lie for some time
in Haentsch's fluid,[14] opportunity being given for evaporation of the
alcohol and water. When the material shows the proper clearness and
fulness, it may be mounted in jelly in the usual way. Kaiser's formula
gives beautiful results. After mounting, the preparation should be
sealed with some good cement, as Hollis's glue.
FOOTNOTES:
[4] DeBary, _Morphology and Biology of the Fungi,_ p. 428.
[5] See, however, _Ceratiomyxa_, p. 18, following.
[6] Harper in _Botanical Gazette_, Vol. XXX., p. 219.
[7] The following germination periods are furnished by
Dr. Constantineanu (_Inaugural Dissertation ueber die
Entwickelungsbedingungen der Myxomyceten_; Halle, 1907).
_Reticularia lycoperdon_ 30 to 60 min.
_Fuligo ovata_ 30 to 90 min.
_Stemonitis splendens_ 5 to 6 hrs.
_Perichaena depressa_ 5 to 8 hrs.
_Amaurochaete atra_ 6 to 10 hrs.
_Arcyria incarnata_ 8 to 10 hrs.
_Lycogala epidendrum_ to 60 hrs.
_Physarum didermoides_ 1 to 10 da.
_Dictydium cancellatum_ 1 to 20 da.
These records are for sowings in drop cultures, in distilled water, kept
at temperature of 65 deg.-70 deg. F. (18 deg.-20 deg. C.).
Our own experiments have been made both with distilled water and
tap-water with the advantage in favor of the latter. _Dictydium
cancellatum_ germinates in tap-water at temperature 70 deg.-80 deg. F. in 12-15
hours fresh from the field. _Fuligo ovata_ spores were all swarming in
about one hour at the same temperature. Jahn (_Myxomycetenstudien; Ber.
der Deutschen Bot. Ges._ Bd. XXIII., p. 495) finds that the germination
in some cases as _Stemonitis_ species, is hastened by wetting, then
drying, then wetting again.
Pinoy thinks microbes aid in germination (_Bull. Soc. Myc. de France_ T.
XVIII.).
[8] The plasmodium in this case chances to be red, scarlet, etc.
[9] "Die Myxomyceten sind ebenso den Pilzen wie den echten Thieren
verwandt."--Rostafinski; closing sentence of the _Versuch_, thesis for
his doctorate at the University of Strasburg, 1873.
[10] _Botanical Gazette_, XVII., pp. 389, etc.; 1892.
[11] Researches of Olive, _Trans. Wis. Acad. Sci., Arts and Let._, XV.,
Pt. 2, p. 771, and of Jahn, _Ber. d. Deutsch Bot. Ges._ XXVI., p. 342,
and XXIX., p. 231, demonstrate synapsis, and accordingly some form of
alternation among the slime-moulds. From the protracted and painstaking
investigation of the German author it appears that in _Didymium_ at
least, and probably _Badhamia_ synapsis immediately precedes
spore-formation as in _Ceratiomyxa_; that the amoeboid issue of the
spores are haploid; the nuclei of the plasmodium, diploid; that the
ordinary vegetative plasmodium is accordingly sporophytic. That is, the
sporophytic phase is dominant, as in higher plants.
[12] Cf., 1884, _Ver. Morph. u. Biol. der Pilz. Mycet. u. Bact._, p.
478. Italics, in quotations, ours.
[13] See _Journal of Mycology_, Washington, D. C., Vol. VII., No. 2;
also _Bulletin No. 66, Agric. Station of Vermont_. See also Bull. _33
Arizona Agric. Ex. Station_: An Inquiry into the Cause and Nature of
Crown-Gall. J. W. Tuomey. Also _Bull. Torrey Bot. Club_, Vol. 21, p. 26,
where it appears that club-root may attack crucifers generally.
Professor B. M. Duggar in _Fungous Diseases of Plants_, pp. 97-102,
gives to club-root an illustrated chapter.
[14]
Haentsch's Fluid:--
Alcohol 90% three parts
Water two parts
Glycerine one part
THE NORTH AMERICAN SLIME-MOULDS
THE MYXOMYCETES (_Link_) _DeBary_
Chlorophyl-less organisms whose vegetative phase consists of a naked
mass of multinuclear protoplasm, the _plasmodium_; reproduced by spores
which are either free or more commonly enclosed in sporangia, and which
on germinating produce ciliated or amoeboid zoospores, whose
coalescence gives rise to the plasmodium.
The Myxomycetes are,--
_A._ _Parasites_, in the cells of living plants PHYTOMYXINAE
_B._ _Saprophytes_, developed in connection with decaying
vegetable matter:
_a._ With free spores EXOSPOREAE
_b._ With spores in receptacles or sporangia MYXOGASTRES
Sub-Class PHYTOMYXINAE _Schroeter_
1889. _Phytomyxinae Schroeter, Engl. u. Prantl._, I., i., pp. 1 and 5.
The parasitic Myxomycetes affecting plants include but few (four or
five) species, distributed among four genera. All are parasites in the
cells of particular hosts; their vegetative phase is plasmodial and
their spores are formed by the simultaneous breaking up of the
plasmodium into an indefinite number of independent cells. But a single
genus need here concern us,--
=Plasmodiophora= _Woronin_
1879. _Plasmodiophora_ Woronin, _Pringsh. Jahrb._, XI., p. 548.
Parasitic in the parenchymatous cells of the roots of living plants,
causing noticeable enlargement of the affected organ, producing at
length galls, knots, and various deformities and distortions. Spores
spherical, smooth, colorless, 16 mu.
I. PLASMODIOPHORA BRASSICAE _Woronin_.
1879. _Plasmodiophora brassicae_ Woronin, _op. cit._
This species, typical of forms so far reported in this country, infests
the roots of cabbages,[15] and produces a very serious disease of that
vegetable. In England the malady has long been known under the names
"clubbing," "fingers and toes," etc. The roots affected swell greatly,
and at length resemble sometimes the flexed fingers of the human hand;
hence the English name. As the disease progresses, the roots speedily
rot away, to the serious injury of the leaf-bearing portion of the
plant. In badly affected fields, sometimes one-half of the crop is
utterly destroyed. Careful search continued through several years has
not availed to bring this species to my personal acquaintance.
For a full account of the parasitism of this species and its
distribution in the United States see _Jour. Myc._, VII., p. 79; also
_Bull._ 66, Agric. Sta. of Vermont.
Sub-Class EXOSPOREAE _Rost._
1873._ Exosporeae_ Rostafinski, _Versuch_, p. 2.
Spores developed, superficially, outside the fructification, which
consists of sporophores, membranous, or slender and branching; spores
white, stalked. A single genus,--
=Ceratiomyxa= _Schroeter_
1889. _Ceratiomyxa_ Schroeter, _Engl. u. Prantl_, I., i., p. 16. For
further synonymy, see under first species.
Sporangia none; spores superficial, borne on erect papillae or pillars,
or even on the inside of minute depressions or pits; each spore
surmounting a delicate pedicel or stalk. The spores on germinating give
rise to amoeboid zoospores, which undergo repeated divisions, later
become ciliate, and at length again amoeboid to blend into genuine
plasmodia. At maturity the plasmodium gives rise to numerous minute
divisions, each of which may lengthen in a direction perpendicular to
the surface and bear a spore at the tip.
The homologies between the structures just described and the
fructification of the ordinary slime-mould are somewhat obscure, if
indeed any really exist. Are these minute reproductive bodies
spores?--their behavior on germination is unique; are they
sporangia?--the arrested development they exhibit is none the less
puzzling. Perhaps the sporiferous pillars represent incipient stipes,
the spores the uncombined fragments of what might otherwise have
coalesced at the summit of the pillar to form a true sporangium.[16]
Several species have been recognized, all referable probably to one or
two, or at most, four forms. That universally recognized alike in the
literature of the past and in recent studies is,--
1. CERATIOMYXA FRUTICULOSA (_Muell._) _Macbr._
PLATE I., Figs. 7 and 7 _a_.
1729. _Puccinia ramosa, bifurcata_, etc. Micheli, p. 213, Tab. 92,
Fig. 2.
1775. _Byssus fruticulosa_ Mueller, in _Fl. Dan._, t. 718, Fig. 2.
1778. _Tremella hydnoidea_ Jacquin, _Misc._, Vol. I., t. 16.
1783. _Clavaria puccinia_ Batsch, _Elench. Fung._, p. 139, Fig. 19.
1791. _Puccinia byssoides_ Gmelin, _Syst. Naturae_, p. 1462.
1791. _Clavaria byssoides_ Bulliard, _Champ. de la France_, t. 415,
Fig. 2.
1794. _Isaria mucida_ Pers., Roemer, _N. Mag. Bot._, I., p. 121.
1801. _Isaria mucida_ Pers., _Syn. Meth._, p. 688.
1805. _Ceratium hydnoides_ Alb. & Schw., _Consp. Fung._, p. 258.
1811. _Ceratiomyxa porioides_ (A. & S.) Schroet., _Mycet._, p. 26,
_var._
1829. _Ceratium hydnoides_ Fries, _Syst. Myc._, III., p. 294.
1872. _Ceratium hydnoides_ Wor. & Fam., _Mem. Acad. Imp._, Petersburg.
1887. _Ceratium hydnoides_ DeBary, _Comp. Morph. Fung._, p. 432.
1889. _Ceratiomyxa mucida_ Schroeter, _Engl. u. Prantl Nat. Pflanz._,
I., i., p. 16.
1893. _Ceratiomyxa mucida_, Pers., Macbr., _Bull. Nat. Hist. Iowa_,
II., p. 114.
1894. _Ceratiomyxa mucida_ Schroet., Lister, _Mycetozoa_, p. 25.
Plasmodium in rotten wood, white or nearly transparent; when fruiting,
forming on the substratum mould-like patches composed of the minute
sporiferous pillars, generally in clusters of three or more together;
spores white, ovoid, or ellipsoidal, smooth, 10-12x6 mu.
Very common, occurring in summer on shaded rotten logs, especially
after warm showers and in sultry weather. Easily distinguishable from
all similar moulds by the absence of mycelium or of anything like a
hypha. In Europe the plant seems to be in autumn exceedingly common.
Micheli not only described the form but figured it, nearly two hundred
years ago. Micheli's figure is good, as is that of Mueller, _Fl. Dan._,
l. c. Mueller referred the species to a Linnean genus _Byssus_, which
seems to have included Algae rather than anything else, if one can
determine its limits at all. The same thing is true of _Tremella_; but
this name is now otherwise applied, as are all the other generic names
down to _Ceratium_, Alb. & Schw. But this had been by Schrank
preoccupied, 1793. See the reference above for 1889. As for specific
name, there seems no reason to depart from the rule of priority, since
Mueller's work is determinative.
_Ceratiomyxa arbuscula_, Berk. & Br., apparently a form of this, is
cited from Toronto by Miss Currie. Massee gives it recognition; Lister
as varietal. The sporophores are inclined to be simple, stipitate and
dendroid.
_C. filiforme_ of the English authors latest named is a wonderful thing
and deserves a paragraph here, if not recognition as a distinct species.
It occurs rarely; but once it appears, attracts attention. As in the
historic species, the sporifers are white, stand more or less erect, but
are every way finer and larger. Each individual sporifer rises like a
stiff stem, as of white thread, 2-3 mm. high; at top a tuft of fruiting
branchlets, more or less distinct. All taken together, we have a dense
mat completely concealing the substratum and spreading out sometimes
over an area of surprising extent, several centimetres square.
Common everywhere in summer on decaying sticks and wood of every
description, especially in wet places. Alaska to Nicaragua, and probably
around the world.
2. CERATIOMYXA PORIOIDES (_Alb. & Schw._) _Schroeter._
1805. _Ceratium porioides_ Alb. & Schw., _Consp. Fung._, p. 359.
1829. _Ceratium porioides_ Fries, _Syst. Myc._, III., p. 295.
1873. _Ceratium porioides_ Fam. & Wor. _Acad. Imp._, XX., 3, p. 5.
1889. _Ceratiomyxa porioides_ Schroet., _Engl. u. Prantl_, I., i.,
p. 16.
1894. _Ceratiomyxa mucida_ Schroet. var. _porioides_ Lister,
_Mycetozoa_, p. 26.
1899. _Ceratiomyxa porioides_ Alb. & Schw. (Schroet.), Macbr., _N.
A. S._, p. 19.
1911. _Ceratiomyxa porioides_ Alb. & Schw., Schroet., _List.
Mycet._, p. 26, _var._
Entire fructification confluent forming a mucilaginous mass, porose.
Pores ample, angulate, at length radiate-dentate. Spores as in the
preceding. Plasmodium yellow.
Of these two species Fries remarks: "... Duae sunt distinctissimae, inter
has vero longa formarum intermediarum series." Famintzin and Woronin not
only concur, but consider it were more fitting to place the present
species in a distinct genus, as _Polyporus_ is set off from _Hydnum_. A
species based upon the color of the vegetative phase only, unconfirmed
by any subsequent differential character in the fruit would seem
somewhat hazardous. The color of the plasmodium is incident probably to
varied nutrient environment. Pores, however, are usually in evidence.
Iowa, Tennessee, Missouri, etc.; probably common everywhere.
Sub-Class MYXOGASTRES (_Fries_) _Macbr._
1829. Sub-order _Myxogastres_ Fries, _Syst. Myc._, III., p. 67.
1833. Sub-order _Myxomycetes_ Link, _Handb. der Gew._, 3, p. 405.
1833. Sub-order _Myxomycetes_ Wallroth, _Fl. Crypt._, II., p. 333, in
part.
1858. Class _Mycetozoa_ DeBary, _Bot. Zeitung_, 1858, pp. 357-365, in
part.
1889. Class _Myxogastres_ Schroeter, _Engl. u. Prantl_, Nat. Pflanz.,
I., i., p. 16.
1892. Class _Myxogastres_ (Fries) Massee, _Monograph_, p. 28.
1894. Class _Mycetozoa_ Lister, _Mycetozoa_, p. 21, in part.
Except as just described, the slime-moulds present abundant, minute,
unicellular spores, enclosed in sporangia more or less perfectly
defined, and attended by peculiar thread-like structures, free or
variously attached and conjoined, the so-called _capillitium_.
So far as known, the spores on germination give rise to zoospores, at
first amoeboid, later ciliate, again amoeboid, conjugating in pairs,
then, in some cases, at least, coalescing and dividing indefinitely to
form the plasmodial or vegetative phase.[17]
=Key to the Orders of the Myxogastres=
Spore-mass black or violaceous, rarely ferruginous Series A
Spore-mass never black; usually some shade of brown or
yellow, rarely purplish or rosy, etc. Series B
SERIES A
1. Capillitium present, delicate, thread-like;
sporangia calcareous more or less throughout I. PHYSARALES
2. Capillitium present, thread-like, arising
usually as anastomosing branches from a
well-developed columella, which in a single
genus contains lime; sporangia otherwise
non-calcareous II. STEMONITALES
SERIES B
3. Capillitium none, or very imperfectly developed; spores of some
shade of brown, rarely purplish III. CRIBRARIALES
4. Capillitium the inwardly produced irregular extremities of plates
or tubules, which by their interweaving outwardly make up the
aethalial wall; spores pale, ashen IV. LYCOGALALES
5. Capillitium made up of more or less distinctly sculptured threads,
parietal or free, simple, branched, or reticulate; spores
commonly yellow V. TRICHIALES
This sequence is meant to convey the idea that the presence of lime is
indicative of differentiation less complete. That the plasmodium should
at the outset eliminate, by refusing the unnecessary lime, is indicative
of higher rank than that the lime should be carried until the last and
then be crystallized out, or excreted by simple desiccation. The
circumstance that the excreted lime may sometimes serve a protective
purpose in the fruit, does not vitiate the general principle. In Series
B the differentiation reaches a climax in the sculptured capillitium of
the trichias.
ORDER I
=PHYSARALES=
Spores violaceous-black. The capillitium usually delicate and
thread-like; peridium and capillitium, one or other or both, more or
less extensively surcharged with lime. Peridium simple or double.
Fructification various.
This order is recognizable by several characteristics, but is especially
marked by the peculiar calcareous deposits which affect the capillitium
or peridium, now one, now the other, more often both.
As here defined, the order Physarales includes two distinct families; of
the one _Physarum_, of the other _Didymium_, is type.
=Key to the Families of the Order Physarales=
_A._ Fructification often calcareous throughout;
capillitium intricate _Physaraceae_
_B._ Calcareous deposits, when present, affecting the
peridium only, or sometimes the stipe, in the
typical genus plainly crystalline; capillitium
simple
_Didymiaceae_
A. PHYSARACEAE
=Key to the Genera of the Physaraceae=
_A._ Fructification aethalioid 1. _Fuligo_
_B._ Fructification plasmodiocarpous or of distinct
sporangia.
_a._ Peridium evidently calcareous.
i. Capillitium calcareous throughout 2. _Badhamia_
ii. Capillitium largely hyaline.
O Sporangia globose, etc.;
dehiscence irregular 3. _Physarum_
OO Sporangia vasiform or more or less
tubular
+ Dehiscence by a lid or more
or less circumscissile 4. _Craterium_
++ Dehiscence irregular,
peridium introverted 5. _Physarella_
_b._ Peridium apparently limeless, at least outside.
i. Plasmodiocarpous 6. _Cienkowskia_
ii. Sporangia distinct 7. _Leocarpus_
C. Extra-limital.
_a._ Sporangia stipitate, saucer-shaped, following
No. 3. _Trichamphora_
_b._ Sporangia elongate allantoid, etc., following
No. 1. _Erionema_
=1. Fuligo= (_Haller_) _Pers._
1753. _Mucor_ Linn., _Sp. Pl._ II., No. 1656 (?).
1768. _Fuligo_ Haller, _Hist. Helv._, Nos. 1233-1235, in part.
1801. _Fuligo_ Haller, _Pers. Syn._, p. 159.
1809. _Aethalium_ Link, _Diss._, I, p. 42.
1829. _Aethalium_ Fries, _Sym. Myc._, III., p. 92.
Sporangia undefined, obscurely woven in and out among each other forming
usually a cushion-shaped aethalioid mass. The outer layer sterile, often
calcareous, forming a fragile crust, more or less defined. The middle
layer sporiferous with calcigerous capillitium. The lowest layer a
membranous hypothallus.
The identity of this genus seems to have been recognized first by
Haller, _op. cit._, but by Persoon more closely defined and illustrated.
Link simply translated the name into Greek, for reasons less evident
now, and in this was followed by Fries. Haller's designation is now
probably securely fixed.
The sporigerous median structure of the fructifications, under whatever
specific name or names, is entirely confused. Sporangial walls, if ever
such there were, are hardly as such recoverable, seemingly _indicated_
only, in the changes to which the aethalium submits as in the ripening
the sporogenic plasm passes on to spores.
In the present state of our knowledge the forms of this genus present
withal a most perplexing problem. Are they simply phases of a single
species, or are they in style and in structure sufficiently constant in
their admitted variety, to claim specific rank and separate description?
To follow the example of Greville and recognize in all the literature of
two hundred years varied descriptions of a single type,--this were
perhaps the easier and speedier disposal of the case. Fries thought so
to treat the problem but was unable to keep faith with his own decision;
for no sooner he states the genus monotypic than he proceeds forthwith
to offer four varieties, a. b. c. d., viz. those by Persoon and others
duly recognized as species.
Recent students all, however, seem to find convenience in specific
division. All seem disposed to honor Dr. Peck's _Fuligo ochracea_
whether or not by the name he gave; and of other varieties some seem
impressed by the constancy of one, some of another characteristic, thus
indicating that to careful observers all over the world there are
differences that may be recognized, that have been recognized again and
again. If there are two species there are certainly more. Out of the
gatherings of many years one may set in order not less than five
variations in the fruiting of _Fuligo_, five distinct types of
fructification, to all appearing sufficiently constant for specific
recognition.
It will be said, has been said, was said by Fries, that these variations
are insignificant, "pendent ex aeris constitutione"; but as a matter of
fact the several types now in question may be found on the same day, so
that evidently something other than the atmospheric environment must
determine.
Again it is said that the differences are in external form or color
only, the spores in all cases almost if not quite the same. This is
true; but specific characters are _surface_ characters in fact: a
species morphologically is merely the form in which a _kind_ or _genus_
presents itself. If the presentation be constant, for our convenience we
say so, in bestowing a name. Whether in our present treatment the
convenience is purely personal, students may decide.
However it all may be, there are in this part of the world many varying
presentations of _Fuligo_ capable of illustration and description; the
same forms, perhaps, which have attracted the notice of the more acute
mycologists in the older history of the subject. Some of these forms we
here venture to describe, with such annotation as may show something of
present knowledge.
=Key to the Species of Fuligo=
_A._ Aethalium 1 cm. or less; spores spherical 1. _F. muscorum_
_B._ Aethalium larger, or plasmodiocarpous, even
sporangi-form, crust white, smooth, even,
spores elliptical 2. _F. cinerea_
_C._ Aethalia larger, 2 cm. or more.
1. Cortex yellow, etc., not white; spores 6-8 mu 3. _F. septica_
2. Cortex nearly or quite wanting; spores
10-12 4. _F. intermedia_
3. Cortex white, a foamy crust; spores 15-25 5. _F. megaspora_
1. FULIGO MUSCORUM _Alb. & Schw._
1894. _Fuligo muscorum_, Alb. & Schw. Lister, _Mycetozoa_, p. 67.
1875. _Licea ochracea_ Peck, N. Y. _Rep._, XVIII., p. 55.
1879. _Fuligo ochracea_ Peck, N. Y. _Rep._, XXXI., p. 56.
1894. _Fuligo muscorum_, Alb. & Schw., Lister, _Mycetozoa_, p. 67.
1911. _Fuligo muscorum_ Alb. & Schw., Lister, _Mycetozoa, 2nd ed._,
p. 87.
Plasmodium orange-yellow. Aethalium globoid, very small, 1 cm. or less,
the cortex very thin, greenish yellow; sporangial walls not evident;
capillitium well-developed, the numerous calcareous nodes fusiform or
often branching, and connected by rather short, transparent internodes;
spores coarsely warted, 10-11 mu.
This form seems to differ from _F. septica_ chiefly in its constant
diminutive habit of fruiting, in its delicate cortex, and in its spores,
brighter, larger, and more coarsely warted. The descriptions and figure
by Schweinitz seem referable to nothing else. First reported by
Albertini and Schweinitz from Germany; by Schweinitz from the Carolinas;
then by Dr. Peck described as a _Licea_ from New York. It seems less
commonly collected in the United States.
2. FULIGO CINEREA (_Schw._) _Morg._
PLATE X., Figs. 3, 3 _a_, and 3 _b_, and Plate XXIII.
1831. _Enteridium cinereum_ Schw., _N. A. F._, No. 2365.
1875. _Physarum ellipsosporum_ Rost., _Mon. App._, p. 10.
1884. _Aethaliopsis stercoriformis_ Zopf., _Pilzthiere_, p. 150.
1894. _Fuligo ellipsospora_ Lister, _Mycetozoa_, p. 67.
1896. _Fuligo cinerea_ (Schw.) Morg., _Cin. Soc. Nat. Hist._, p. 105.
1899. _Physarum ellipsosporum_ Rost., Macbr. _N. A. S._, p. 27.
1911. _Fuligo cinerea_ Morg., List., _Mycetozoa_, 2nd ed., p. 88.
Plasmodium milk-white, watery. Plasmodiocarp long and widely effused,
anon winding, here and there reticulate, always applanate; sometimes in
form an aethalium, the peridial cortex membranous, firm, thick, and
white. Capillitium well-developed, furnished with lime. Spores
thin-walled, ellipsoidal, violaceous, plicate-rugose, 14-16 x 11-12 mu.
Not common. Found occasionally in shaded situations on piles of rotting
straw or in the woods, especially on detritus of the bracken. The spores
are many of them ellipsoidal; some are spherical; all are decidedly
spinulose, perhaps might appear plicate-rugulose when dry or shrunken.
Calcareous nodules very large and irregular, white.
Schweinitz, _loc. cit._, described this form as _Enteridium cinereum_.
Rostafinski referred it to the genus _Physarum_, but was obliged to
adopt also a new specific name, as that suggested by Schweinitz was
already in use in the genus _Physarum_. Zopf, _Die Pilzthiere_, p. 149,
founds a new genus on what seems to be the same form as here considered.
This he publishes as _Aethaliopsis stercoriformis_ Z. Massee regards the
specimens discovered by Zopf as belonging to the genus _Fuligo_, and
Lister regards Rostafinski's type as _Fuligo_, and includes Zopf's
material under the Rostafinskian species.
This has been described as properly an American form; Lister cites other
far localities.
3. FULIGO SEPTICA (_Linn._) _Gmel_.
1753. _Mucor septicus_ Linn., _Sp. Pl._ II., No. 1656 (?).
1763. _Mucor ovatus_ Schaeff., _Fung. Bav._, p. 132, Fig. 192.
1791. _Fuligo septica_ (Linn.) Gmel., _Syst. Nat._, p. 1466.
1826. _Fuligo varians_ Sommf., _Fl. Lapl. Sup._, p. 231.
1809. _Aethalium flavum_ Link, _Diss._, I., p. 42.
1829. _Aethalium septicum_ Fr., _Syst. Myc._, III., p. 93.
1875. _Fuligo varians_ Sommf., Rost., _Mon._, p. 134.
1892. _Fuligo varians_ Sommf., Macbr., _Bull. Lab. Nat. Hist. Ia._
II., p. 160.
1894. _Fuligo septica_ (Linn.) Lister, _Mycetozoa_, p. 66.
1899. _Fuligo ovata_ (Schaeff.) Macbr., _N. A. S._, p. 23.
1911. _Fuligo septica_ Gmel., Lister, _Mycetozoa, 2nd ed._, p. 86.
This remarkable and universal species presents as stated many forms and
phases. Of these five have been selected as representative.
1. Form _a._ Plasmodium yellow; cortex yellow, or
orange-brown, strongly calcareous friable; form
indefinite _F. ovata_
2. Form _b._ Cortex less calcareous porose, yellowish brown,
fructification definite, pulvinate _F. rufa_
3. Form _c._ Cortex smooth, persistent; fructification
small, less than two inches _F. laevis_
4. Form _d._ Plasmodium yellow; cortex none; capillitium
yellow, fructification thin, sometimes wide-spread _F. flava_
5. Form _e._ Plasmodium violaceous, dark; cortex almost
none; whole mass reddish or violet _F. violacea_
1. Form _a._ _Fuligo ovata_ (Schaeff.) Pers.
Plasmodium bright yellow; aethalium pale brown, or yellowish-ochraceous,
of variable size and shape, one to many cm. in diameter, and one to two
cm. thick, enclosed by a distinct calcareous crust, which varies in
texture, thickness, and color; capillitium well developed but variable
in color, form, and extent; spore-mass dull black, sooty; spores
spherical, purplish brown, nearly smooth, 7-9 mu.
Under this name may be placed our most common form. Rising with an
abundant yellowish creamy plasmodium from masses of decaying vegetation,
lumber, sawdust, half buried logs, it creeps about with energy
unsurpassed, coming to rest only in some position specially exposed, as
the top of a log or stump, the face of a stone or post, or even the high
clods of a cultivated field! The fructification is large, yellow, or at
most pale ochraceous, the surface when mature extremely friable like dry
foam. Bulliard figures this phase well on Plate 424, Fig. 2, and calls
it _Reticularia_ (_Fuligo_) _hortensis_, from its affecting the soils of
gardens. More than thirty fructifications have appeared at one time,
varying in size from one to twenty cm. in a field of potatoes, well
tilled, and less than an acre in extent! Such is life's perennial
exuberance on this time-worn old world of ours!
Schaeffer's plate CXII represents probably the same thing. So also
Bolton's plate, CXXXIV. Sowerby's Fig. 2 on plate 199, and figures 1 and
2 on Greville's plate 272 possibly also depict this form. Persoon calls
this _F. vaporaria_ because it frequents hotbeds and the like, and
believes this to represent the "_untuosus flavus_" of Linnee, although
he thinks Schaeffer's specimens do not. The calcareous internal structure
is white.
2. Form _b_, _F. rufa_ Pers.
This type of Fuligo is very different from the preceding in form, habit,
and color. In form it is much more definite, usually thick, well-rounded
and with some solidity. The interior fructification is gray throughout,
much less expanded than in _a_; in fact does not resemble _a_ at all!
The cortex is porose but firm, orange at first, but becoming tawny with
age, even in the herbarium. Bulliard figures it well, plate 380, Fig. 1,
and Sowerby's Fig. 1 on plate 399 is also good, as are also Greville's
figure 3 on plate 272 showing the two colors referred to. Not uncommon
in the forest from June till September, but far more rare than _a_:
always well-marked, with no other forms associated.
3. Form _c_, _F. laevis_ Pers.
This is a still more specialized type of the group. The fructification
is usually small, smooth, about an inch in diameter and sometimes nearly
as thick; the cortex rusty brown, enduring, persisting often when all
the sporiferous grayish mass has been distributed through chinks, or
from below. The figure 2 on plate X. shows this form. This also is a
forest species, is autumnal rather, but may be taken sometimes as early
as July. The cortex is not at all porose or spongy, in color reddish or
brown, fragile indeed, but not to the touch, in the herbarium enduring
for years.
4. Form _d_, _F. flava_ Pers.
PLATE X, Figs. 2, 2 _a_, 2 _b_.
This is hardly _F. flava_ of Persoon; rather of Morgan who uses
Persoon's specific designation. Persoon cites Bolton's fig. CXXXIV,
which is yellow indeed but is the ordinary presentation of _F. septica_.
The form here considered is remarkable for its delicacy; extremely thin,
perhaps one layer only of overlying elongate flexuous sporangia(?),
covered by the merest shadow of a cortex in the form of yellow dust,
soon lost: the capillitial structure yellow throughout; occurring upon
fallen logs in moist dark woods; not common.
5. Form _e_, _F. violacea_ Pers.
Plasmodium (Morgan _teste_) dark red, or wine-colored; the aethalium
thin, two or three inches wide, covered by a cortex at first dull red
and very soft, at length almost wholly vanishing, so that the entire
mass takes on a purple-violet tint, upper surface varied with white;
capillitium rather open, the more or less inflated, large, irregular
nodes joined by long, slender, delicate, transparent filaments; spores
dark violet, minutely roughened, spherical, about 7.5 mu.
Ohio, Tennessee. Probably everywhere, but not distinguished from 1.
Professor Morgan, who gave the genus under consideration much attention,
regarded _F. violacea_ as a form particularly well-defined. What the
value of plasmodic color as a specific character in general, and how far
such character is in the present case definitive, because constant, are
points yet to be determined.
4. FULIGO INTERMEDIA _Macbr. n. s._
Aethalium two to three cm. in greatest diameter, .5-1 cm. thick, covered
with a thin, fragile, but not calcareous, greyish or brownish cortex;
the spore-mass grey or violaceous-grey, firm, not at all sooty, the
sporangia intricate, their walls more or less calcareous; capillitium
not conspicuous; spores globose, pale purple, slightly roughened, 10-12
mu.
This form has been repeatedly sent me from Denver, Colorado, by
Professor Bethel. I have refrained from publishing it, still anxious to
believe that all fuligos on the face of the earth were of one species.
In the species next following it must be admitted that the
spore-variations are too wide to remain comfortably under shelter of a
single specific name. The present species is not _F. septica_, neither
is it _F. megaspora_; it is _F. intermedia_.
Colorado; Iowa.
5. FULIGO MEGASPORA _Sturg._
1913. _Fuligo megaspora_ Sturg., _Col. Coll. Pub._, p. 443.
Aethalium pulvinate one to three inches in diameter, covered with a thick
spongy incrustation of lime, white or yellowish toward the base:
sporangia convolute, the walls membranous, brittle, charged throughout
with round white granules of lime, 1.5-2 mu in diameter: columella none:
capillitium of delicate, colorless, anastomosing tubules, bearing toward
the center large, white, branching calcareous nodules; spores spherical,
or somewhat oval, dark purple-brown, rough-tuberculate, 15-20 mu.
This species differs as pointed out by Professor Sturgis, chiefly in the
character of the spores, their unusual size and roughness.[18]
Colorado; Africa!--_Robert Fries._
=EXTRA-LIMITAL=
=Erionema= _Penzig_
1898. _Erionema_ Penzig, _Die Myx. d. Fl. v. Beutenzorg_, p. 36.
Sporangia plasmodiocarpous but distinct, cylindrical; capillitium
intricate, elastic; nodules few.
1. ERIONEMA AUREUM _Penzig_
1898. _Erionema aureum_ Penz. _l. c._
Sporangia elongate, clustered, pendulous, yellow or grayish yellow,
generally stipitate on long flaccid stalks, or sessile and interlacing:
stipes yellow, blending with the hypothallus; capillitium intricate,
expanding at maturity after the manner of _Arcyria_ to several times the
sporangial length, the nodules small, yellow; spores nearly smooth,
violaceous-brown, 5-6 mu.
This unique form is near the fuligos which it resembles, especially when
sessile, in its intricate sporangia. The spores also are those of the
common _Fuligo septica_. The habit is however entirely different. Mr.
Fetch describes clusters in Ceylon, hanging free, four to six cm. in
length!
=2. Badhamia= (_Berkeley_) _Rost._
1852. _Badhamia_ Berkeley, _Trans. Linn. Soc._, XXI., p. 153.
1875. _Badhamia_ Rostafinski, _Monograph_, p. 139.
Sporangia simple; peridial wall simple, thin, breaking irregularly;
capillitium formed of abundant, richly anastomosing tubules, filled
throughout their entire length with calcareous granules; the nodes often
feebly represented; stipe poorly developed or wanting entirely;
columella, except in forms sometimes assigned to the sub-genus
_Scyphium_, poorly developed or none; spores frequently adherent in
clusters.
The whole genus calls for careful and protracted study; and the
present so-called species are like something new on the world;
as full of vagaries as though but just entered upon their
phylogenetic race.
This genus is closely related to _Physarum_, but differs in having the
capillitium calcareous throughout. Forms occur and are included here, in
which the capillitium, especially in some parts, is physarum-like,
physaroid. Nevertheless, the distinctions hold good as a rule, and are
at once diagnostic.
In capillitial differentiation the badhamias are definite and beautiful.
The net in a typical species, as _B. papaveracea_, is throughout
uniformly evenly tubular, the calcareous deposits delicate in the
extreme, presenting, as the spores disappear, an elegant trabecular
structure as if to support the persisting peridium if not the original
content. In other forms the capillitium is physaroid, with swollen
nodes, but heavily calcareous but not quite throughout. _Badhamia_,
_Physarum_, _Tilmadoche_, _Craterium_ present a consistent group, of
which _Physarum_ is the generalized expression.
Berkeley's idea of the genus was expressed as follows: "Peridium naked
or furfuraceous. Spores in groups, enclosed, at first, in a hyaline
sack." Rostafinski, while accepting Berkeley's generic name, redefined
it, emphasized the calcareous capillitium, and made reference to the
spore-adherence only to assert that Berkeley's description was, in this
particular, based on mistaken observation. In some species, the spores
do, in fact, show a tendency to cling together, a characteristic which
Badham was perhaps first to notice; but that this is occasioned by their
being surrounded by a sac or common pellicle has not been proved nor
even suggested, by any subsequent investigator. Berkeley's genus was
therefore founded upon a slight mistake; but we may conserve his rights
in the premises if we write _Badhamia_ (Berk.) Rost., and so keep
history straight.
=Key to the Species of Badhamia=
_A._ Spores ovoid or ellipsoidal
_a._ Spores free 1. _B. ovispora_
_b._ Spores adherent 2. _B. versicolor_
_B._ Spores spherical
_a._ Sporangia yellow
i. Spores free 3. _B. decipiens_
ii. Spores adhering 4. _B. nitens_
_b._ Sporangia grey, spores free
i. Always sessile 5. _B. panicea_
ii. Stalked, at least some of them
O Stipe when present black
+ Globose, small .5 mm. 6. _B. affinis_
++ Larger, spores strongly spinulose 7. _B. macrocarpa_
+++ Discoidal or annulate 8. _B. orbiculata_
OO Stipes membranous yellowish
+ Stipes long, sporangia iridescent 9. _B. magna_
++ Stipes short or none; iridescent 10. _B. foliicola_
_c._ Sporangia grey, spores adherent
i. Stipe when present yellowish
+ Wall iridescent, spores uniformly
marked 11. _B. utricularis_
++ More calcareous, spores strongly
marked on one side 12. _B. capsulifera_
+++ Colorado, spores anon barred 13. _B. populina_
ii. Stipe when present black 14. _B. papaveracea_
_d._ Sporangia brown, lilacine
i. Sessile 15. _B. lilacina_
ii. Stipitate, columellate 16. _B. rubiginosa_
1. BADHAMIA OVISPORA _Racib._
1884. _Badhamia ovispora_ Racib., _Myx. Ag. Cracov._, XII., p. 72.
Sporangia sessile depressed-globose or plasmodiocarpous, white or
ochraceous, covered by dense calcareous scales; capillitium white, the
lime-granules sometimes aggregate at the center to form a
pseudo-columella; spores not adhering, brownish-purple ellipsoidal, 8 x
10-10 x 15 mu.
Reported from Bohemia, England, Pennsylvania.
2. BADHAMIA VERSICOLOR _Lister_.
1901. _Badhamia versicolor_ List., _Jour. Bot._, XXXIX., p. 81.
1911. _Badhamia versicolor_ List., _Mycetozoa 2nd ed._, p. 35.
Sporangia scattered or clustered, minute, .3-.5 mm., grey or
flesh-colored, sessile, the calcareous deposits slight; capillitium
white or apricot-colored; spores ovoid, 8 x 10-9 x 12 mu, clustered,
purplish, and warted at the broader end, elsewhere colorless and smooth.
This little species, as it comes to us, is grey, very uneven in size,
.2-.5 mm. and generally irregular in form and habit, perhaps scarce
mature. The capillitium is white, physaroid. The spores furnish the
distinguishing character. Sometimes globose, about 9-10. They are most
of them definitely and permanently affected in shape by the fact of
cluster-association, narrower in the direction of the cluster center.
The indications are that these may become globose with maturity.
Colorado,--_Bethel_; Scotland.
3. BADHAMIA DECIPIENS (_Curtis_) _Berk._
1848. _Physarum decipiens_ Curtis, _Am. Jour. Sci._, VI., p. 352.
1873. _Badhamia decipiens_ Berk., _Grev._, II., p. 66.
1873. _Physarum chrysotrichum_ Berk. & C., Grev. II., p. 66.
1876. _Badhamia chrysotricha_ (Berk. & C.) Rost., _App._, p. 4.
Sporangia gregarious, depressed-spherical or ovate, sessile,
occasionally plasmodiocarpous, dull yellow, roughened by the rather
large numerous calcareous scales; columella none; capillitium dull
orange, strongly calcareous, only slightly widened at the nodes;
spore-mass black; spores pale violet, minutely spinulose, free, 10-12 mu.
Among badhamias this and the next species are at once distinguished by
the color. If the brief description (_Grev._, II., p. 66) can be
regarded as defining anything, this is the same as _P. chrysotrichum_
Berk. & C. It resembles somewhat _P. serpula_ Morg., but differs
externally in color and in the surface scales, which are not perceptible
in the _Physarum_. The present species also resembles _Cienkowskia
reticulata_ (Schw.) Rost., but has a different capillitium. See under
that species.
Chiefly eastern and American. New England, Pennsylvania, Ohio, South
Carolina; reported recently also from Sweden and Germany.
4. BADHAMIA NITENS Berk.
1852. _Badhamia nitens_ Berk., _Trans. Linn. Soc._, XXI., p. 153.
1863. _Badhamia inaurata_ Currey, _Trans. Linn. Soc._, XXIV., p. 156.
1873. _Badhamia nitens_ Berk., Rost., _Mon. App._, p. 3.
Sporangia gregarious or closely crowded, globose or depressed-globose,
.5-1 mm. in diameter, yellow or greenish yellow, rugulose, sessile;
capillitium yellow, forming an open net with occasional thickenings at
the nodes; spores clustered, delicately roughened, violaceous-brown,
10-12 mu.
This much resembles the preceding species except in the clustered
spores, and more commonly aggregate habit. The spores, as usual when
clustered, are conspicuously echinulate on the outer side. This did not
escape the notice of the author of the species, _op. cit._
Colorado, Oregon. Reported from West Indies, Ceylon, various parts of
Europe.
5. BADHAMIA PANICEA (Fries) Rost.
1829. _Physarum paniceum_ Fries, _Syst. Myc._, III., p. 141.
1873. _Badhamia panicea_ (Fr.) Rost., Fuckel, _Sym. Myc. Nachtr._,
2, p. 71.
Sporangia gregarious or aggregated in closely compacted clusters,
globose or hemispherical, sessile, the peridium thin, transparent,
thickly dotted with white calcareous scales; stipe none; columella none,
although a pseudo-columella sometimes appears, formed by a more dense
development of the capillitium near the centre of the sporangium below;
capillitium abundantly developed, quite uniformly thickened, but showing
an occasional delicate connecting thread, the nodes also somewhat
flattened and enlarged; spore-mass black; spores by transmitted light,
bright violaceous-brown, minutely roughened, 10-13 mu. Plasmodium is said
to be white.
In America this seems to be a purely western species. Specimens are
before us from western Iowa and from Colorado, South Dakota, Nevada, and
Southern California. It is very well marked, though liable perhaps to be
mistaken at first sight for sessile phases of _P. notabile_ or _P.
cinereum_. The capillitium is, however, at once determinative. Colorado;
_Bethel_. Europe generally.
6. BADHAMIA AFFINIS _Rost._
1875. BADHAMIA AFFINIS Rost., _Mon._, p. 143.
Sporangia aggregated, cespitose and sessile, or sometimes stipitate,
depressed above, flat or umbilicate below, the wall grayish white,
rugulose, and more or less calcareous-scaly; the stipe when present
erect or sometimes nodding, black or brownish black; hypothallus scanty;
columella none; capillitium not abundant, white, the nodes somewhat
expanded; spores globose, minutely roughened, violet-brown, large, 16-17
mu.
Chiefly on moss, the pale ashen sporangia generally very small, mounted
on the tips of the leaves, sometimes sessile, sometimes with a distinct
black stipe in which case the peridium is distinctly umbilicate.
Specimens from Kansas referred here have the stipe pale, rugose, long,
about twice the sporangium; habitat bark.
Rare. New York, Ohio, Kansas; more recently reported from Scotland and
Japan.
There is nothing new to be added here; nor appears any other place to
which such material as we have may be referred. New collections no doubt
will one day appear, when the identity may, let us hope, be made secure.
Meantime we have a form closely related which may be entered as
BADHAMIA IOWENSIS _Macbr. n. s._
Sporangia gregarious or loosely scattered, depressed globose, .4-.6 mm.
in diameter, stipitate, grey, flecked by rather prominent but small
rounded calcareous scales: the stipe short, half the diameter of the
sporangium, black or very dark brown, without hypothallus but widening
above into a shallow expanded base for the sporangia; columella none:
capillitium dull yellow, sometimes white, strongly calcareous,
physaroid, heavy; spores free, dark brown in mass, pale violet by
transmitted light, minutely verruculose, the tiny warts in some areas
more densely placed, producing evident shadowy spots, 10-11 mu.
This interesting little species occurs on the lower surface of fallen
logs, blocks, etc., in colonies of considerable extent, hundreds of
sporangia in a place. The capillitium is comparable to that of _B.
decipiens_ or _B. panicea_; it is physaroid to the extent that an
occasional filament may be found non-calcic, and not typically
badhamioid as in _B. papaveracea_, _B. macrocarpa_. The sporangial base
persists, dark brown, bearing traces of the clumsy capillitium, but no
columella real or simulated. Blackhawk Co., Iowa; _communicavit Dr.
Jessie Parish_. See Plate XX., 1, 1 _a_, 1 _b_.
Reddish or roseate forms sometimes appear in colonies otherwise as
described. It differs from _B. affinis_ in the size and character of the
spores, in color and character of the capillitium, habit and surface
markings.
7. BADHAMIA MACROCARPA (_Ces._) _Rost._
1855. _Physarum macrocarpon_ Cesati, _Flora_, XXXVIII., p. 271.
1875. _Badhamia macrocarpa_ (Ces.) Rost., _Mon._, p. 143.
Sporangia scattered or closely aggregate, crowded globose or
sub-globose, generally sessile, rugulose, white; the peridium
membranous, white above, below yellowish or brown; capillitium not
abundant, thoroughly calcareous, the nodes broad, conspicuous, the
connecting tubules rigid; columella none; hypothallus scant or none;
spore-mass black, spores non-adherent, by transmitted light bright clear
brown, thickly spinulose all over, large spherical, 12-15 mu.
Closely resembles externally _B. panicea_, but is easily distinguished
by larger and remarkably _spinulose_ spores, in this particular
unrivalled in the entire genus. European authors describe both sessile
and stipitate forms. American specimens generally are sessile and for
the most part closely crowded, almost heaped; but--Prof. Bethel finds
this in winter everywhere on fallen rotting stems of Opuntia and on the
bases of dead Yucca leaves, still attached. Associated with the typical
phase and often _occurring alone on the Yucca_ leaves is a discoidal
form which when first sent in (1908) was called var. _gracilis_.
Presented alone to one ignorant of its history and associations, it
would surely pass for a distinct species. This stalked phase is very
delicate; the stipe pale brown, or yellow. See Plate II., Fig. 9. See
also Sturgis _Col. Coll. Pub._ XII., 408.
8. BADHAMIA ORBICULATA _Rex._
PLATE XIV., Fig. 4.
1893. _Badhamia orbiculata_ Rex. _Proc. Phil. Acad._, p. 372.
1894. _Badhamia macrocarpa Rost._, Lister, _Mycetozoa_, p. 34
(in part).
1911. _Badhamia orbiculata_ Rex., Lister, _Mycetozoa, 2nd ed._, p. 37
Sporangia stipitate or sessile, orbicular discoidal, irregularly
elongated or plasmodiocarpous, averaging about 1 mm. in width, generally
stipitate, and when stipitate, flattened or depressed above, plane or
slightly umbilicate below; the peridium simple, more or less translucent
from the varying number of innate granules, sometimes covered with
circular flat masses of lime, gray except the point of attachment to the
stipe which is brown; stipe short, black, rough, plicate; capillitium
dense at the centre, radiant at the periphery where it meets the
sporangial wall, white; spores violaceous black, minutely warted, 12-15
mu.
This is a beautiful species, easily known by its discoidal or almost
annulate sporangia mounted upon short dark black stipes. The stipe in
western collections is sometimes very short, but generally suffices to
raise the sporangium, a little at least, above the substratum. Sessile
and plasmodiocarpous forms do occur with the typical stipitate phase,
but may be regarded here as elsewhere as indicative of incomplete
development. Plasmodium cream-colored, or pale yellow.
Pennsylvania, Ohio, Iowa, Nebraska, South Dakota, Colorado.
9. BADHAMIA MAGNA _Peck._
PLATE XIV., Fig. 1.
1871. _Dictydium magnum_ Peck, _Rep. N. Y. State Mus._, XXIV., p. 84.
1879. _Badhamia magna_ Peck, _Rep. N. Y. State Mus._, XXXI., p. 56.
1894. _Badhamia macrocarpa Rost._, Lister, _Mycetozoa_, p. 34,
in part.
1892. _Bahamia varia_ Mass. _Mon. Myxog._, p. 319, in part.
1894. _Badhamia magna_ Peck, List., _Mycetozoa_, p. 33.
1899. _Badhamia capsulifera_ (Berk.) Macbr., _N. A. S._, p. 68.
1911. _Badhamia magna_ Peck, Lister, _Mycetozoa, 2nd ed._, p. 34.
Sporangia globose or ellipsoid, .7-1 mm., pale iridescent, stipitate;
peridium thin with slight calcareous deposits, rugulose, opening
irregularly, white; stipe long flaccid, straw-colored; capillitium an
elegant uniform net, its threads stiffened by slight deposits of lime,
the nodes little thickened, badhamioid; spores free, dusky with a shade
of violet, minutely spinulose, about 10 mu.
This beautiful species closely resembles some forms of _B. utricularis_
from which it differs chiefly in its unclustered smooth spores. _B.
foliicola_ as recognized here is hardly more than a smaller,
short-stemmed form of this; see species next following.
Not rare in the eastern United States and Canada; Iowa. Seems to take
the place of _B. capsulifera_ of Europe.
10. BADHAMIA FOLIICOLA _Lister_.
1897. _Badhamia foliicola_ List., _Jour. Bot._, XXXV., p. 209.
1911. _Badhamia foliicola_ List., _Mycetozoa, 2nd ed._, p. 34.
"Plasmodium orange." Sporangia smaller, about .5-.6 mm., globose or
ellipsoidal, iridescent-gray, stipitate or sessile, the peridium thin,
rugulose, sparingly calcareous, when empty white; the stipe when present
short but yellowish, of the flaccid sort; capillitium badhamioid; spores
free, delicately spinulescent, dusky-violaceous, about 12-13 mu.
This has been so far collected but once, on the shores of Lake Okoboji.
It was developed, no doubt, on the natural debris of a bur-oak prairie
border, and went to fruit on the leaves, stems, and fruiting spikes of a
species of _Setaria_. It may prove to be different from the _B.
foliicola_ of Europe; future collections and study must reveal that.
Meantime it seems wise to refer it here.
The color of the plasmodium is quoted from Miss Lister; a fact of some
importance only when constant and confirmed by other criteria.
Iowa; Toronto,--_Miss Currie._
11. BADHAMIA UTRICULARIS (_Bull._) _Berk._
1791. _Sphaerocarpus utricularis_ Bull., _Champ._, p. 128, t. 417,
Fig. 1.
1826. _Physarum utriculare_ Chev., _Fl. Paris_, I., p. 337.
1829. _Physarum utriculare_ Fries, _Syst. Myc._, III., p. 139.
1852. _Badhamia utricularis_ (Bull.) Berk., _Tr. Linn. Soc._, XXI.,
p. 153.
Sporangia clustered, spherical or ovoid, large, sessile or mounted on
long, thin, strand-like stalks, blue-gray, violet-iridescent or
cinereous, smooth or more often rugulose; the stipes when present poorly
differentiated, as if thread-like filaments and strips of the
plasmodium, often branched and always reclining or even prostrate;
hypothallus none; capillitium a large-meshed open network of rather
slender tubules, the nodes unequally developed, white with the enclosed
lime; spores not strictly adherent though not without some tendency to
stick together, delicately warted, bright violet-brown, 10-12 mu.
This species resembles _B. capsulifera_, but is distinguished by a more
strongly rugulose less calcareous peridium and a more profuse
development of filamentous stipes, but especially by the character of
the spores. The spores of the present species while inclined, when
mounted in a liquid, to stay together, nevertheless do not coalesce in
heaps as in the related species, nor do they show any differentiation in
the episporic markings, these being uniform over the entire spore.
This is one of the finest and perhaps the most beautiful species of this
fine genus. It is a forest species, generally to be found on trunks of
fallen _Populus_ or _Tilia_ where the fine soft gray colonies often
spread for several inches along the ridges and in crevices of the bark.
Colorado (_Bethel_); Mississippi valley and east.
12. BADHAMIA CAPSULIFERA (_Bull._) _Berkeley_.
1791. _Sphaerocarpus capsulifer_ Bull., _Champ._, p. 139, t. 470,
Fig. 2.
1801. _Physarum hyalinum_ Pers., _Syn. Meth. Fung._, p. 170.
1852. _Badhamia capsulifera_ Berk., _Tr. Lin. Soc._, XXI., p. 153.
1852. _Badhamia hyalina_ Berk., _Tr. Lin. Soc._, XXI., p. 153.
1875. _Badhamia hyalina_ (Pers.) Rost., _Mon._, p. 139.
1875. _Badhamia capsulifera_ (Bull.) Rost., _Mon._, p. 141.
1894. _Badhamia hyalina_ Lister, _Mycetozoa_, p. 30.
1911. _Badhamia capsulifera_ Lister, _Mycetozoa, 2nd ed._, p. 31.
Sporangia clustered or gregarious, sessile or sometimes stipitate,
globose or obovoid, gray or greenish white, snow-white when empty; the
peridium thin, translucent; the stipe, when present, as in _B.
utricularis_, although generally shorter and better developed, yellow or
straw colored; capillitium a very loose, open network of white,
lime-filled tubules, not much expanded at the nodes; columella none;
spore-mass purplish-brown; spores adhering in clusters of five or six to
twenty or more, globose, but affected somewhat by mutual pressure, rough
throughout, the exposed surface in the cluster, more distinctly warted,
10-12 mu.
This is _Badhamia hyalina_ (Pers.) Berk., Rost., _Mon._, p. 139; but
Rostafinski himself admits that the two species, here united, as he
defined them, are very much alike, having "the same spores and
capillitium", differing in the form of the sporangium, an inconstant
feature. Bulliard's name has precedence; his descriptions of this and
the preceding species are remarkable.
The peculiarly adherent spores distinguish the species from _B.
utricularis_; and the sporangia sessile or with short but strand-like
stipes, distinguish it from _B. papaveracea_.
The description above is for the typical European form. Lister expresses
doubt whether this occurs in the United States. The form from Iowa which
is the basis for the inclusion of the species in N. A. S. is, we
believe, nothing else than _B. capsulifera_ (Bull.) Berk. The form
approaches _B. populina_ as this is presented in Colorado. The Iowa
specimens are white, aggregate, superimposed, etc., but have the
capillitium and spores exactly as described for the type. Accordingly
_B. populina_ as this occurs in Colorado has been for years referred to
the Berkeley species. The thicker more strongly calcareous peridia
constitute, as would appear, the principal difference in the forms from
Colorado. See next species.
13. BADHAMIA POPULINA _List._
1904. _Badhamia populina_ List. _Jour. Bot._, XLII., p. 129.
1911. _Badhamia populina_ List. _Mycetozoa, 2nd ed._, p. 32.
Plasmodium white; sporangia sessile, crowded, heaped, large, 1.5 mm.,
rarely stipitate, globose or ovoid, white; stipe when present brown;
capillitial strands broad, calcareous; spores clustered, 16-20 in a
cluster, purple-brown, roughened and sometimes marked by obscure ridges
and bands, 10-12 mu.
Generally distinguishable by its unusually large calcareous, white
sporangia. The peridia are strongly calcareous, shell-like in texture.
In some cases the color is tinted with rose.
This species is very near _B. capsulifera_ as recognized in the United
States. When white the Colorado material corresponds almost exactly with
the forms collected in Iowa, and regarded as representing the species
just named. The Colorado gatherings are more strongly calcareous and the
spores sometimes present the variations named. "The Colorado phase of
the American form."
Colorado,--_Bethel_. Europe?
14. BADHAMIA PAPAVERACEA _Berk. & Rav._
PLATE IX., Figs. 6, 6_a_, and 6_b_.
1873. _Badhamia papaveracea_ Berk. & Rav., _Grev._, II., p. 66.
1894. _Badhamia hyalina_ var. _papaveracea_ Lister, _Mycetozoa_,
p. 30.
1899. _Badhamia papaveracea_ Berk. & Rav., Macbr., _N. A. S._, p. 69.
1911. _Badhamia papaveracea_ Berk. & Rav., List., _Mycetozoa,
2nd ed._, p. 32.
Sporangia gregarious, globose, large, stipitate, iridescent-gray; the
peridium thin, translucent, and containing but little calcareous
deposits, smooth or slightly rugulose; stipe very short, but generally
distinct, black or very dark brown; hypothallus none; capillitium a
network of large meshes with expanded nodes, prominent, white,
persistent after the spores have been blown away; spore-mass deep brown;
spores adherent as in _B. capsulifera_, marked in much the same way, and
about the same size, 10-12.5 mu
Distinguished by its short, dark, stipe and adherent spores.
Not common. New England, Pennsylvania, Ohio, Maryland, South Carolina,
Wisconsin, Iowa.
15. BADHAMIA LILACINA (_Fries_) _Rost._
1829. _Physarum lilacinum_ Fries, _Syst. Myc._, III., p. 141.
1875. _Badhamia lilacina_ (Fries) Rost., _Mon._, p. 145.
1892. _Craterium lilacinum_ Mass., _Mon._, p. 271.
1894. _Badhamia lilacina_ (Fr.) Rost., Lister, _Mycetozoa_, p. 34.
1911. _Badhamia lilacina_ (Fr.) Rost., List., _Mycetozoa_, 2nd ed.,
p. 38.
Sporangia globose, sub-globose, or obconical, sessile, gregarious or
more or less clustered, supported by a thin, continuous, transparent
hypothallus; the peridium smooth pale, lilac-brown without, white
within; stipe none, although some sporangia have a narrowed base;
columella none, the pseudo-columella formed by a more densely aggregated
capillitium near the base; capillitium dense, white, strongly nodulose;
spore-mass black; spores dark, violaceous-brown by transmitted light,
distinctly warted, or reticulate, the reticulations resembling somewhat
those of some of the trichias, as _T. affinis_, 10-15 mu.
Easily recognizable, generally at sight, by its peculiar color. White
forms, however, occur; often lilac-tinted and white from the same
plasmodium. A perfectly white colony seems to be rare. Both colors are
shown in specimens distributed. _N. A. F._, 2494.
Common eastward, Ontario, New England, Pennsylvania, Ohio, etc. Not
reported west of the Mississippi River.
Whatever the color, the spores are in every case positively diagnostic.
The episporic markings are unlike those of any other species in the
present order. Dr. Rex describes some New York forms as provided with a
short but distinct stipe. Such forms resemble externally _Scyphium
rubiginosum_ (Chev.) Rost. The hypothallus is also unique. V. next
species.
16. BADHAMIA RUBIGINOSA (_Chev._) _Rost._
PLATE X., Figs. 1, 1_a_, 1_b_, 1_c_.
1826. _Physarum rubiginosum_ Chev., _Fl. Par._, p. 338.
1872. _Craterium obovatum_ Peck, _Rep. N. Y. Mus._, XXVI., p. 75.
1875. _Scyphium rubiginosum_ (Chev.) Rost., _Mon._, p. 148.
1876. _Badhamia rubiginosa_ (Chev.) Rost., _Mon. App._, p. 5.
1892. _Craterium rubiginosum_ Massee, _Mon._, p. 270.
Sporangia gregarious, obovoid, grayish brown, stipitate, the peridium
simple, membranous, above thin, pale, more or less calcareous below,
more persistent blending with the stipe; stipe erect, reddish brown or
purplish, expanded below into a small hypothallus, above, prolonged
within the sporangia more than half its height as a definite columella;
capillitium very dense, snow white, long persistent with the lower
two-thirds of the sporangial wall; spore-mass dark brown; spores by
transmitted light, dark violet or purple-brown, minutely roughened or
spinulose, not adherent, 12-14 mu.
This is probably the most common badhamia in the country and in the
world. It is found every year, in the woods, on masses of decaying
leaves, especially those of various species of oak. The plasmodium is
yellow. The fructifications are very distinct, not likely to be
mistaken for those of any other species; the stipes constitute a very
prominent feature in every gathering I have seen. Sometimes these are
more or less coalescent, especially toward the base, where they are apt
to be also wrinkled or longitudinally striate; in other specimens the
stipes are well differentiated, long, terete, with little or no
hypothallus.
_Badhamia curtisii_ (Berk.) Rost. is according to Lister (Mon., p. 35) a
sessile phase of this species. The only specimens known are in the
herbarium of Berkeley, now at Kew. The species is based upon a gathering
from S. Carolina. Berkeley thought it a didymium, called it _D.
curtisii_.
Reported from western Europe; the typical form abundant in the forested
regions of eastern N. America, especially in the Mississippi valley.
17. BADHAMIA SUBAQUILA _Macbr._
1899. _Badhamia subaquila_ Macbr., _N. A. S._, p. 64.
Sporangia closely gregarious or crowded, globose or sub-globose,
sessile, brown, the peridium a thin but persistent brown membrane,
rupturing above irregularly and remaining as a cup after spore
dispersal; hypothallus none; capillitium strongly developed, thoroughly
calcareous, the meshes large, the nodular thickenings broad, white;
spores globose, in mass black, by transmitted light brown, very
rough-warted, large, 15-18 mu.
The variety is founded on material sent from Maine by the late Mr. F. L.
Harvey. Professor Harvey, upon the authority of Mr. Morgan of Ohio,
quotes the species, _Bull. Tor. Bot. Club_, 24, 67, as _B. verna_
(Somm.) Rost. But the specimens certainly do not conform to description
of _B. verna_. Here the wall corresponds with what is seen in _B.
rubiginosa_; but the spores are much larger, and the capillitial
structure very different.
Miss Lister regards this a form of No. 16. So far, the original
gathering represents the species; but the woods of Maine are certain one
day to send added information.
Rare. On mossy logs, Maine.
=3. Physarum= (_Persoon_) _Rost._
1794.[19] _Physarum_ Pers., _Rom. Neu. Mag. f. d. Bot._, I., p. 88,
in part.
1795. _Physarum_ Pers., _Ust. Ann. Bot._, XV., p. 5, in part.
1801. _Physarum_ Pers., _Syn. Fung._, p. 168, in part.
1829. _Physarum_ (Pers.) Fries, _Syst. Myc._, II., p. 127, in part.
1875. _Physarum_ (Pers.) Rost., _Mon._, p. 93.
Sporangia plasmodiocarpous, aethalioid or distinct; the peridium usually
simple, sometimes double, irregularly dehiscent, more or less definitely
calcareous; capillitium a uniform irregular net, dilated and calcareous
at the nodes, adherent on all sides to the peridial wall.
This large and cosmopolitan genus is readily recognized by the
characters quoted. It may be added that the capillitial threads are
always exceedingly delicate, probably tubular, but never filled with
lime throughout; the peridium may be almost nude or encrusted with lime,
which, where present, is always amorphous, never crystalline; the
sporangia when distinct may be either sessile or stipitate, and the
stipe in the latter case is often hollow and charged with lime. In
capillitium intermediate between _Leocarpus_ and _Badhamia_, since in
the first the capillitium is unequally calcareous, diverse, while in
_Badhamia_ the capillitium is intricate and calcareous throughout.
As first set up by its founder, the genus included diverse forms, only
one or two of which would be included in the genus as now limited.[20]
Persoon, however, was left to develop the matter to suit himself, and in
successive works gave, under this generic name, more and more prominence
to forms now so referred. Fries, _Syst. Myc._, III., pp. 127 _et seq._,
still better establishes the genus, though still including forms that,
judging from the description, seem to belong elsewhere. Twenty years
later Fries revising somewhat his earlier work thought to improve the
chances of future students by reducing the number of physarums. This he
would do by setting out certain evidently inter-related forms to make a
new genus, _Tilmadoche_.
He named two or three species only, leaving his sucessors to add others
as occasion offered.[21]
Rostafinski approved the good intention of Fries, but in the
_Monograph_, he entirely re-cast the genus as constituted by Fries;
actually called the species 'first cited' a typical physarum! Would not
have it in the new genus at all, first or last; but instead took the
second species of Fries as the type and added several forms, some from
the Friesian list, to make up a respectable group.
Until quite recently writers on the subject have generally approved the
course adopted by the Polish author. The arrangement showed features of
convenience, even if artificial to a degree. Perhaps we gain advantage
in all directions if we treat the original genus _Physarum_ as a whole,
but in the key take advantage of Fries' suggestion. We may write--
=Key to the Species of Physarum=
1. Capillitium irregularly reticulate throughout; calcic
nodes various _Physarum_
2. Capillitium more regular, especially below, furcate;
nodes fusoid _Tilmadoche_
SECTION I. PHYSARUM
I. Fructification not stipitate, more or less plasmodiocarpous.
1. Peridium simple.
_a._ Calcareous deposits yellow 1. _P. serpula_
_b._ Calcareous deposits reddish or orange 2. _P. lateritium_
_c._ Calcareous deposits white, peridium rugulose 3. _P. vernum_
2. Peridium double.
_a._ Fructification flatly compressed 4. _P. sinuosum_
_b._ Fructification less compressed, rounded.
i. Outer peridium white 5. _P. bitectum_
ii. Outer peridium brown or brown-tinged 6. _P. bogoriense_
iii. Outer peridium yellow; capillitium yellow 7. _P. alpinum_
II. Fructification of sporangia more or less distinct.
A. Sporangia sessile, globose, ovoid, reniform, etc.
1. Peridium double.
_a._ Sporangia white, peridium testaceous. 8. _P. diderma_
_b._ Sporangia tinged with yellow.
i. Sporangia as if interwoven,
compressed 9. _P. contextum_
ii. Sporangia more nearly free, distinct.
o Spores pale, inner peridium
brittle 10. _P. conglomeratum_
oo Spores spinulose, dark violet 11. _P. mortoni_
_c._ Sporangia brown, dehiscence revolute 12. _P. brunneolum_
2. Peridium simple, calcareous, flaky.
_a._ Sporangia grey, plasmodiocarpous;
spores dusky, 10-12 forms of 3
_b._ Sporangia grey, more or less dense;
spores violet, 6-7 13. _P. cinereum_
_c._ Calcareous deposits yellow or greenish,
spores 7-9 14. _P. virescens_
_d._ Sporangia rusty or reddish brown,
more or less dense 15. _P. rubiginosum_
_e._ Sporangia minute, lignicolous,
the fructification much extended
upon a hypothallus, lime deposit
tawny 16. _P. instratum_
_f._ Sporangia white, depressed, annulate,
sometimes with short stipes 17. _P. megalosporum_
3. Peridium simple, not flaky,
small .2-.3 mm., heaped 18. _P. confertum_
B. Sporangia, at least some of them, stipitate.
_a._ Sporangia columellate.
i. Columella small, usually conical.
O Sporangium yellow.
o Columella white 19. _P. melleum_
oo Columella yellow 20. _P. citrinum_
OO Sporangium not yellow.
o Capillitial mass persistent.
+ Sporangia globose,
pallid or white 21. _P. globuliferum_
++ Sporangia blue or lilac,
rose, etc. 22. _P. lilacinum_
+++ Sporangia drab or brownish 23. _P. murinum_
++++ Sporangia wine-red 24. _P. pulcherrimum_
oo Capillitial-mass less
persistent; orange 25. _P. pulcherripes_
ii. Columella long, 4-5 the sporangium
non-calcareous. 26. _P. penetrale_
iii. Columella large globose 27. _P. luteo-album_
_b._ Sporangia without columella.
i. Sporangia nucleate, calcareous at center.
O Stipe yellow 28. _P. nucleatum_
OO Stipe white 29. _P. wingatense_
ii. Sporangia non-nucleate.
O Sporangia purple 30. _P. newtoni_
OO Sporangia blue, spotted with red 31. _P. psittacinum_
OOO Grey or white, iridescent betimes.
o Sporangia white, discoidal;
stipe yellow 32. _P. discoidale_
oo Sporangia lightly calcareous,
iridescent, sub-globose,
diam. about = to the stout,
brown, slightly wrinkled
stipe 33. _P. leucophaeum_
ooo Sporangia globose or sub-globose.
x. Small, .5 mm.
+ Stipe erect, clear brown 34. _P. nodulosum_
++ Stipe weak, yellow,
stuffed 35. _P. maculatum_
xx. Larger, lime-capped; stipe
strand-like 36. _P. didermoides_
xxx. Stipe snow-white, fragile 37. _P. leucopus_
xxxx. Stipe generally distinctly fluted
+ Sporangia laterally
compressed, fan-shaped 38. _P. compressum_
++ Sporangia typically
globose, umbilicate
below, connate, etc.,
strongly calcareous 39. _P. notabile_
+++ Sporangia reniform,
concave below _P. affine_,
see under 38
++++ Sporangia larger, to 1 mm.,
nearly limeless,
iridescent 40. _P. tropicale_
oooo Sporangia obovate, compound,
clustered, the stipe fuscous,
fluted, short. 41. _P. nicaraguense_
OOOO Sporangia yellow, rarely iridescent or brown.
o Capillitial nodes white.
x. Stipe also white 42. _P. sulphureum_
xx. Stipe flesh-colored,
spores smaller 43. _P. carneum_
xxx. Stipe red or reddish brown 44. _P. citrinellum_
xxxx. Stipe yellowish, flaccid,
sporangia leocarpine 45. _P. albescens_
xxxxx. Stipe very short or none,
sporangia cylindric, brown 46. _P. variabile_
oo Capillitium nodes yellow or orange-yellow.
x. Badhamioid,
larger,--to .8 mm. 47. _P. auriscalpium_
xx. Physaroid, base persistent 48. _P. oblatum_
ooo Capillitium nodes pure yellow.
x. Capillitial threads yellow 49. _P. galbeum_
xx. Capillitial threads hyaline 50. _P. tenerum_
xxx. Peridium iridescent.
+ Capillitium persistent 51. _P. flavicomum_
++ Capillitium less
persistent, larger 52. _P. bethelii_