A.Fructification æthalioid, capillitium poorly defined; columella rudimentary or noneAmaurochætaceæB.Fructification of distinct sporangia, capillitium well defined; the columella generally prominent, long and abundantly branched throughoutStemonitaceæC.Sporangia distinct; capillitium developed chiefly or only, from the summit of the columellaLamprodermaceæ
A.AMAUROCHÆTACEÆ
Fructification æthalioid, an inch or two in diameter, in form varying with the habitat and place; capillitium dendroid, consisting of rather stout branches which rise irregularly more or less vertically from the hypothallus, branch repeatedly, often anastomose to form a network, especially toward the periphery; spores black.
A single genus—
1. AmaurochæteRostafinski
The genusAmaurochaeteas defined by Rostafinski and the genusReticulariaas represented byR. lycoperdonBull. stand, the expression, perhaps, of not dissimilar histories. Whether in regressive orprogressive series, each to-day presents a case of arrested development. Each in æthalioid fructification, reveals a mass of involved individual (?) sporangia, so imperfectly developed that their outlines can be inferred rather than anywhere, with absolute definiteness, certainly ascertained. Perhaps, because similar sporangia in the group to which either belongs, do come under other circumstances, to more perfect individual form and function—perhaps for this reason we may look upon these æthalia as exhibiting a suspended performance; the sporangia have failed to go forward to what was evidently a possible, though apparently not an essential destiny in form and figure. For the care and dispersal of the spores, achievement must surely be somewhat impaired. Whatever the measure of such inefficiency, among theStemonitales Amaurochaeteshows the acme, asReticulariaamong the brown-spored forms.
InAmaurochaetethe individuality of anything like separate sporangia is less clear. The view afforded, however, by a good vertical section of a well-developed colony or cushion is interestinglyarborescent. Ragged, dendroid stems arise, dissipated above into a network most intricate, a "pleached arbor" if you please. The resemblance of the overhead net to that presented by a stemonitis or comatricha is very striking.
Key to the Species of Amaurochæte
A.Capillitium rigid, irregular spores rough1.A. fuliginosaB.Capillitium soft, woolly, cincinnate, spores as inA2.A. tubulina
1.Amaurochæte fuliginosa(Sowerby)Macbr.
Plate V., Figs. 8, 8a.
Fructification aethalioid, varying in form and size, if on the upper side of the substratum, pulvinate, if below pendent and almost stipitate, covered with a delicate cortex, at first shining, soon dull, black, fragile, and early dissipated; hypothallus long-persisting, supporting the capillitium, which is extremely variable, irregular, and for itsperfection dependent upon the form assumed by the æthalium, and the conditions of weather, etc., under which it matures, sometimes, especially when prostrate, in a very much depressed æthalium, spreading into long fibrous threads, again under better conditions rising in columella-like forms, supporting a peripheral net; spores dark brown or black, irregularly globose, spinulose, 12.5–15 µ.
Common in Europe, and probably not uncommon in this country wherever pine forests occur. Specimens before us are from New England and New York, Ohio, Carolina, Colorado. Canada.
Sowerby, in his comment on plate 257,Eng. Fungi, says: "It appears to consist of branching threads affixed to the deal and holding a dense mass of sooty powder. Over the whole is a thin, deciduous pellicle." This description seems to be applicable to nothing else. The figure amounts to little. Fries recognizes the English description, as does Rostafinski, but both authors adopt the later name given by Albertini and Schweinitz, simply because of the excellent detailed description found in theConspectus.
2.Amaurochæte tubulina(Alb. & Schw.)Macbr.
Plate XX., 6 and 6a.
Plasmodium at first transparent then white then rosy, ashen or grey finally deepening to jet-black; the æthalium even, thin, variable in extent from one to ten centimeters, covered by a distinct but thin transparent cortex, papillate, extended laterally but a short distance beyond the fructification, fragile, soon disappearing; hypothallus long-persistent, thin, silvery, supporting the capillitium as if by stipes, short slender columns, irregular plates, expansions, etc.; the capillitium an intricate network, very abundant, elastic, on fall of the peridium appearing like tiny tufts of wool, the meshes large, but formed as inStemonitis, persistent, dull black; spores, under the lens, dull olivaceous black, minutely roughened, 12–14 µ.
This species differs from the preceding, already well known, especially in the capillitial characters. In the older species the capillitial branches fray out, and are only sparingly united into a net extremely lax. In the present form the net is the thing, common to all sporangia. The total effect is to lend to the blown-out æthalium a woolly appearance, entirely unlike that of its congener under the same conditions. But until fructification is quite mature, the presence of the collaborating sporangia below is indicated, suggested, by the papillose upper surface.
The amaurochetes are remarkable in that they appear upon coniferous wood, logs or lumber, to all appearance undecayed. The species just described developed abundantly in August on the recently decorticated logs ofPinus ponderosa, on the south-western slopes of Mt. Rainier, Washington. In logging operations in the locality referred to, the trees are felled often at considerable distance from the mill. They are not infrequently large, 75–120 cm. in diameter. The logs are dragged along the ground, the transportation facilitated by removal of the bark from the new fallen trunk. In a few weeks' time, affected by alternate rain and sun, the whole surface becomes marked with hundreds of minute, almost invisible cracks, and it is in the larger of these that the plasmodium of the present species has its habitat. Hardly any mycologic phenomenon is more surprising than to see plasmodia rising to fructification, scores at a time, upon a surface, new and white, showing otherwise no evidence of any decomposition. Doubtless the persisting cambium, the unused starches, sugars, the wood of the season yet unlignified, afford easily accessible nutrition.
When this form was first examined in the laboratory its distinctness was immediately seen. It was without doubt Fries' cribrose reticularia; nobody questions that. Under this name, citing Fries' description, specimens were sent out to herbaria as Harvard. Further study of the records, however, soon convinces one familiar with the ontogeny of the case that we are here face to face with the species, described by Alb. & Schw. in their fineConspectus. Their account of the form, evidently often taken and now described with great care, is entirely clear when read in presence of the facts. It is here submitted,as less easy of access but essential, if the reader would appreciate the present disposal of the species.
"S. TubulinaNOBIS
"S. magna pulvinata subhemisphaerica, stylidiis gregariis circinantibus, capillitiis elongatis cylindraceis in massam pulveraceam fuscam connatis, apicibus obtusis, prominulis, lucidis nigris.
"The size indeed, the circumscribed form, the capillitiums conjoined into a single body—indue this (form) with an appearance peculiar to a degree; however, should anyone prefer to call it a very remarkable variety of the preceding (S. fasciculata), we shall not strenuously refuse. At first glance it looks like a tubulina. After the fashion of its kind, the beginning is soft and milky. The diameter generally an inch and a half to two inches, the height four to six lines; the form perfectly round, or more rarely somewhat oblong. The hypothallus, stout, pellucid silvery, betimes iridescent, when turned to the light, easily separable from the substratum, bears the columellae, dusky, thin, hair-like, aggregate and yet entirely free, and everywhere circinately convergent, depressed by the superimposed burden, hence decumbent: ... the capillitium loosely interwoven, coalesces to a common mass whose smooth and shining surface shows above, regularly disposed minute papillae, the apices of individual sporangia.
"Far from infrequent, on decorticate pine, ofLycogala atruma constant companion"!
It goes of course without saying, that for the authors quoted,Lycogala atrumisAmaurochaete atraRost.A. fuliginosa(Sow.) of more recent students, described and perfectly figured in the volume cited.
It is surprising that they did not enter the present species also as a lycogala. But the stemonitis relationship this time impressed them rather than the æthalial; besides they were misled by theS. fasciculataof Gmelin and Persoon, a composite which the genius of Fries hardly availed to disentangle twenty-five years later.
The last named author, as we see, wrote firstLachnobolus, thenReticularia. He calls the interwoven capillitium—lachne, wool, a"pilam tactu eximie elasticam," etc. He read the description in theConspectus, but carried away the stemonitis suggestion dominant there, as we have seen, putS. tubulinaA. & S. as an undeveloped phase ofS. fusca, which, of course, it is not. It needed not the authority of Rostafinski,Mon., p. 197, to assure us this. The earlier authors describe the species in course of development to complete maturity, and clinch the story by declaring the form a constant companion of the commonly recognized amaurochete, so fixing the relationship for us by habitat also.
These men made a mistake, of course, in placing their species among the stemonites at all. They did much better however than Fries who called it a reticularia. It was also a mistake to citeS. fasciculata,—the small fasciculate tufts ofS. fuscaandS. axiferaoffering by the aggregate habit only faint resemblance,—a possible refuge for those who would prefer another disposition of their species distinct (aliena) though it is.
Since Fries' day the species has been overlooked although the genus has received more than once attention. ZukalHedwigia, XXXV., p. 335, describesA. speciosaas a new species. This Saccardo writes down, Syll. Fung., VII., p. 399,S. tubulinaA. & S., admitting, however, at the same time, that as fine an authority as Raciborsky refuses to call Zukal's species either a stemonite or an amaurochete, thinks it deserving generic appellation of its own.
However,A. speciosaZuk. need not here concern us. Neither in his description nor figures does Zukal at all approach the form we study. His species is not an amaurochete; the size of the spores suggest that, to say nothing of the capillitial structure.
In the same volume VII., the distinguished author introduces another amaurochete,A. minorSacc. & Ellis,Mich.II., p. 566. This is American; sent from Utah by our famous pioneer collector Harkness. A specimen is before us: it is a lepidoderma! in shining, scaly armor dressed; vid. underL. carestianum.
Since the distribution of Washington material, as mentioned, our species reappears at various points in western Europe, points in England, etc., and will no doubt now share, hereafter as a century ago, the habitat so long conceded to the long familiar older type.
B.STEMONITACEÆ
Capillitium abundant, springing usually as dissipating branches from all parts of the columella; the sporangia generally definite and distinct, though sometimes closely placed and generally rising from a common hypothallus.
Key to the Genera of the Stemonitaceæ
A.Fructification æthalioid; capillitium charged with vesicles1.BrefeldiaB.Sporangia distinct, or nearly so.a.Stipe and columella jet-black.1. Capillitium so united as to form a surface net2.Stemonitis2. Capillitial branch-tips free3.Comatrichab.Stipe and columella whitish; calcareous4.Diachaea
1. BrefeldiaRostafinski
Sporangia occupying in the æthalium several layers, those of the median, and especially of the lowest layers, furnished with columellæ which blend beneath; capillitium threads in the lowest layers arising from the columella, in the upper extending radiately between the individual sporangia, and united at the sporangial limits by means of rather large inflated sacs.
The genusBrefeldiais, like some others, difficult to dispose of in any scheme of classification where linear sequence must be followed. Rostafinski placed it in an order by itself. Its relationships are on the one hand withAmaurochaeteandReticularia, and on the other with theStemonitales, though easily distinguished from either. It is intermediate toAmaurochaeteandStemonitis, and withal, as it appears to us, a little nearer the latter, as the limits of the individual sporangia are inBrefeldiapretty well defined.
1.Brefeldia maxima(Fr.)Rost.
Plate V., Figs. 7, 7a, 7b, andPlates XXI.,XXII.
Æthalium large, four to twenty cm, papillate above, violet-black at first, then purple or purple-brown, developed upon a widespread,silver-shining hypothallus; sporangia in favorable cases distinct, indicated above by the papillæ; columellæ obscure, black; capillitium abundant, the threads uniting by multifid ends to surround as with a net the peculiar vesicles; spore-mass dark violet-black, the individual spores paler by transmitted light, distinctly papillose, 12–15 µ.
A very remarkable species and one of the largest, rivalled byFuligoonly. To be compared withReticularia, which it resembles somewhat externally, and with some of the larger specimens ofEnteridium. The plasmodium at first white with a bluish tinge is developed abundantly in rotten wood, preferably a large oak stump, and changes color as maturity comes on, much in the fashion ofStemonitis splendens, leaving a widespread hypothallic film to extend far around the perfected fruit-mass. In well-matured æthalia, "Jove favente," the sporangia stand out perfectly distinct, particularly above and around the margins. Closely and compactly crowded, they become prismatic by mutual pressure, and attain sometimes the height of half an inch or more. In the centre of the fructification, next the hypothallus, the sporangia are very imperfectly differentiated. Many are here horizontally placed, and perhaps supplied with an imperfectly formed peridium,—if so are to be interpreted the lowest parts of the capillitial structure, the long, branching, ribbon-like strands which lie along the hypothallus. Some of these branch repeatedly with flat anastomosing branchlets, ultimately fray out into lengthened threads, and perish after all the superstructure has been blown away. From every part of the structure so described, but more especially from the margins, are given off in profusion the strange cystiferous threads, so characteristic of this genus. These are exceeding delicate filaments, attached at one end, it may be, to a principal branch, at the other free or united to a second which again joins a third, and so looping and branching, dividing, they form a more or less extended network, a capillitium in which are entangled the myriad spores. Each filament bears at its middle point (or is it the meeting point of two?) a peculiar plexus which embraces several large cysts or vesicles whose function or further homology does not readily appear.
From the base of the fructification rise also ascending branches which are black, terete, and not infrequently branched as if to formthe capillitium of a stemonitis. These ascending branches are in many cases, probably in all, real, though as yet imperfectly developed, columellæ. They rise, at least in many cases, directly from the hypothallus, each is central to an individual sporangium, rises to about two-thirds its height, but never attains the summit. The sporangia are so crowded that many are choked off below, never reach the top of the æthalium. In such cases the columella may cease at the sporangium-top. The columella bears cystiferous threads sparingly, if at all; nevertheless these abound in the peripheral portions of the sporangium all the way up, and are especially noticeable beyond the level of the top of the columella. Many are so arranged that the plexus with its vesicles occupies a place in the plane separating adjacent sporangia, suggesting the possibility that we have here to do with an imperfectly developed surface-net and peridium. In this view the cysts would represent the meeting-point of two opposite radial capillitial threads rather than the middle of one. This accords with Rostafinski's observations and drawings. The cysts, then, belong morphologically to the peridium or sporangium wall. It is a stemonitis whose sporangia have never been perfectly differentiated, a case of arrested development. See further underStemonitis confluens.
Rostafinski really offers the first definitive description. Fries probably distinguished it, but his description would not indicate the fact except for the added note wherein appears the reason for discarding an apparently older name, viz., that given by Link. But neither Link nor Sowerby distinguished by description or figureBrefeldiafromAmaurochaete.
Throughout the northern forest; Maine to Vancouver Island: not common.
2. Stemonitis(Gleditsch)Rost.
Sporangia distinct, though often closely aggregate, cylindric, stipitate; columella prominent; capillitium well developed by repeated lateral and apical branching of the columella, at length assuming at the surface the form of a distinct net which supports an evanescent peridium.
The genus is marked by its surface-net supported at the tips of the dichotomously branched divisions of the columella. Over the net is spread, theoretically at least, the peridial film supported by very short points projecting from the net,—the peridial processes; the peridium, however, is seldom seen; in some cases, certainly, is never developed. Rostafinski first defined the genus as employed by recent writers. Gleditsch simply renamed Micheli'sClathroidastrum; all writers subsequent included species of other genera.
The taxonomy of this genus is of the most difficult. Macroscopic, defining characters are few, and even these sometimes uncertain. Microscopic distinctions also tend to be illusive, variable in such fashion that often at the critical point the most exact description fails. All that may be done at present is to recognize two or three definite types and then cautiously differentiate among these with the light we have, until more general study of the group brings to service a wider range of observation with more comprehensive record on which judgment may better be sustained.
We have before us many and beautiful forms of this genus yet unstudied. Some of these doubtless have already found place in our growing taxonomic literature; some apparently undescribed; all to wait wider leisure or perhaps a younger hand.
The entire life-history of every form is none too much if we would set out with any hope of accuracy the genetic relationships for which taxonomy stands. Recently European students are making the color of the plasmodium a basis for species-discrimination, which is good so far. But plasmodic characters are at present unserviceable generally, for two reasons; they vary in the same species; and unfortunately, when most needed, they are unknown and inaccessible. The student is generally confronted by forms mature, the plasmodic stage already past.
Key to the Species of Stemonitis
A.Sporangia connately united.a.Spores verruculose1.S. confluensb.Spores reticulate2.S. trechisporaB.Sporangia at maturity distinct.a.Spore-mass grayish black.1. Larger, 8–12 mm. spores distinctly reticulate or warted, but sometimes nearly smooth3.S. fusca2. Spores reticulate and spinulose.i. Spores adherent, clustered4.S. uviferaii. Sporangia very tall, 15–20 mm., rigid5.S. dictyosporaiii. Sporangia short, jet- or violet-black6.S. nigrescensb.Spore-mass rich brown.1. Columella central.i. Sporangia shorter, 5–6 mm., spores banded7.S. virginiensisii. Sporangia 8–10 mm.; spores verruculose8.S. webberiiii. Sporangia tall, 15–20 mm. or more9.S. splendens2. Columella eccentric, sporangium in cross-section, angular10.S. fenestratac.Spore-mass ferruginous; sporangia in tufts.1. Spores smooth or nearly so.i. Sporangia pale, small, 3–5 mm., crowded, stipe unpolished11.S. smithiiii. Sporangia ferruginous; columella regular12.S. axiferaiii. Sporangia ferruginous; columella proliferate just below the apex13.S. flavogenitaiv. Sporangia, spore-mass, dusky-purplish or brown.O On dead wood.o Scattered, apex blunt14.S. pallidaoo Clustered, acuminate15.S. carolinensisOO On living leaves, preferably, spore-mass brown16.S. herbatica
1.Stemonitis confluensCooke & Ellis.
Plate XI., Figs. 4, 4a, 5.
"Sporangia fasciculate, confluent on a persistent hypothallus, dark fuscous; peridia very fugacious; stipes united at the base, erect, furcate; spores large, brown, globose. On oak bark.
"The stems are branched in a furcate manner and confluent at the base, forming a compact tuft. The capillitium is membranaceous at the angles; spores very large compared with allied species, being 12 µ. The specimens were too fully matured for more satisfactory description."
Such is the original description of this unique and interesting species. The sporangia occur in close-set tufts or clusters, are distinct, separate at their tips and bases only; perhaps not always at base. The capillitium rises by branching from the columella, rather more prolific than usual, and combines to form a distinct superficial net of large even meshes. From the outer arcs of the bounding net spring rather long acute processes which should support the peridium. This, however, is altogether rudimentary. In most places there is no sign of peridium at all, but here and there between contiguous sporangia opposite processes unite and at their point of union a tiny circular disk of the peridial membrane appears. At intervals, therefore, over the entire sporangium are seen these small brown disks, each about equalling in diameter the size of the average mesh. At other points the sporangia do not seem at all coalescent, but where the opposing processes do meet the union is perfect and the little disk seen edgewise looks like some delicate counter strung upon a wire.
The interest attaching to this in view of what has been said aboutAmaurochaeteandBrefeldiais obvious.
Under the lens the spores and capillitium are concolorous, dark fuscous, the spores distinctly verruculose, about 12.5 µ.
The original gathering here described was from New Jersey; twenty years later Mr. Ellis was so fortunate as to find again fine specimens all on oak bark. The sporangia are quite small, only 3 mm. high, when blown out concolorous with the habitat.
2.Stemonitis trechispora(Berk.)Torr.
Plate XX., Figs. 11, 11a, 11b, 11c.
Fructification in form of aggregations of more or less coalescent, small, dark-brown or dull black, sessile sporangia; hypothallus continuous, well-developed; columella black, gently tapering to a point beneath the apex, the capillitial branches, irregular, few, but passing into an open rather evenly-meshed net, the mesh several times thespore-diameter, free-ending branch-tips not lacking; the spores by transmitted light distinctly brown, the epispore a beautiful reticulation, a dozen or more cells to the hemisphere, 10–12 µ.
This is entered sometimes as a variety ofS. fuscato which species relationship would seem remote. The differences lie in form, color and structure. The spores alone are distinctive; there are none such, so far, none just like them, elsewhere in the genus. Torrend and Lister both enter the form as varietal; why not set it out, and save questions? The habitat approaches that ofAmaurochaete, but the sporangia are distinct.
For our specimens we are indebted to the kindness of Dr. Roland Thaxter. The specimens were taken in a half-dry marsh, near Cambridge.
Material from Toronto sent by Professor Faull is also provisionally here referred. The form has netted spores, but they are not quite the same. The structure besides is more that of an amaurochaete; it has the peculiar basal webs and band-like stipes at base, stipes that never rise from horizontal to perpendicular and characterizeReticulariaand especiallyBrefeldiaas well as the usual amaurochaete. SeePlate XX., Figs. 9, 9a, 9b.
3.Stemonitis fusca(Roth)Rost.
Plate VI., Figs. 4, 4a, 4b
Sporangia tufted, generally in small clusters 6–8 mm., the individual sporangia slender, cylindric, blue-black or fuscous, becoming pallid as the spores are lost, stipitate; stipe short, about one-fourth the total height, black, shining; hypothallus scanty, but common to all the sporangia; columella prominent, attaining almost the apex of the sporangium, freely branching to support the capillitial net; capillitium of slender dusky threads, which freely anastomose to form a dense interior network, and outwardly at length combine to form aclose-meshed net; spores pale, dusky violet, usually beautifully spinulose-reticulate, but sometimes warted or spinulose only, or nearly smooth, 7–7.5 µ.
As here set out the description is intended to includeS. maximaSchw. of the former edition. Rostafinski, Mon.l. c., describesS. fuscaRoth. as having "spores smooth." Since most American gatherings have reticulated spores, and since Schweinitz described a black American species, his specific name seemed appropriate for all except smooth-spored forms.
In the meantime two things have happened; Mr. Lister has examined the specimens remaining in the Strasburg herbarium and finds them with reticulate spores. The statement quoted from theMonographevidently does not apply toallof Rostafinski's material; but under the circumstances the namefuscamay easily take the field, especially since another discovery makes for the same conclusion. The evidence is good thatS. maximaSchw. was indeed the largest, i. e. perhaps, thetalleststemonitis he ever saw! probably, as his scanty herbarium-remnant shows,S. fenestrataRex!
4.Stemonitis uviferan. s.
Plate XX., Figs. 8, 8a, 8b, 8c.
Sporangia tufted, generally in medium-sized clusters much as inS. fusca. The individual sporangium 7–9 mm. high, dark, slender, brown, becoming dull black or pallid as the spores are lost, stipitate, the stipe about one-fourth to one-third the total height, black polished shining; hypothallus distinct, common to all sporangia, purple-brown, shining; columella distinct, attaining almost the summit of the sporangium but inclined to waver a little at last, in other words, flexuose toward the top, freely branching, the branches rather stout, anastomosing to support the capillitial net; the meshes larger, several times the spore-diameter, the spores sooty-brown, distinctly warted or spinulescent, about 7–8 µ, clustered in groups of four or more.
Mt. Rainier, Washington,—1914.
5.Stemonitis dictyosporaRost.
Sporangia crowded in colonies of unusual size, 4–8 cm., tall, rigid 18–25 mm., slender, erect, stipitate, black throughout; the columella prominent, reaching nearly to the apex, abundantly branched, the branches forming an intricate dark brown capillitium; the net large-meshed several times the spore-diameter; the spores reticulate, spinulose, clear violet, 7–8 µ.
We here recover as is believed one of Rostafinski's best-described species. Our material is from Nicaragua, by kindness of Professor Shimek. Its relationship is withS. fuscawhere Rostafinski placed it. The phrase describing spore-color is his.
6.Stemonitis nigrescensRex.
Sporangia gregarious, upon a common hypothallus, erect, small, cylindric, stipitate; stipe black, extremely short, about half a millimetre; columella reaching the apex; capillitium violet-black, darker near the surface, forming a complete superficial net at the lower part of the sporangium only, elsewhere irregular or vanishing; spore-mass nearly black; single spores violet-black under the lens, the epispore spinulose and reticulate, about 8 µ.
The author of this species remarks: "This species is noteworthy for its comparatively short stipes, its very spinulose spores, and its black or nearly black color, the slight violet tint being only apparent on close inspection, especially in fresh moist specimens."
It is a small but very beautiful form, at first sight to be mistaken for a shortS. fusca, though much more intensely black. The capillitium is concolorous, the inner network of rather few open meshes, the outer of large hexagonal openings, the arcuate threads of which are remarkable for the size, and especially the number, of the peridial processes, as many as five or six sometimes appearing along one side of a single mesh. The stipe is very short, and the columella runs as a straight, gradually diminishing axis to the very apex of the sporangium. Total height 3–5 mm.
The EnglishMonographincludes this withS. fusca; but it seems quite distinct in size, habit, color, etc., and has been found in the mountainous regions of Virginia and North Carolina, as well as about Philadelphia.
7.Stemonitis virginiensisRex.
Sporangia erect, gregarious, from a common hypothallus, generally clustered, cylindric or elongate-ovate, stipitate; stipe black, shining; columella reaching the apex, where it blends with the capillitium; capillitium delicate, the meshes of the net small, scarcely greater than the diameter of the spores; spore-mass umber brown; epispores reticulated, with ten or twelve meshes to the hemisphere, 5–7 µ.
This is a beautiful, and, as it seems to us, a very distinct, species. The markings on the epispore are sufficient to identify it. These are conspicuously banded somewhat as the spores ofTrichia favoginea, for example. In habit, size of the sporangia, and capillitial branching, this species recallsComatricha typhoides(Bull.) Rost. All the sporangia examined are, however, plainly stemonitis in type, possessing the characteristic superficial net.
Until further light this may stand as offered in the first edition. Miss Lister prefers to enter it, banded spores and all, with the comatrichas, on account of color, size and occasional default (?) of surface net.
Virginia,Dr. Rex.
8.Stemonitis webberiRex.
Plate XI., Figs. 6, 7, 8.
Sporangia clustered, usually in small tufts 1 cm. wide, rusty brown in color, 8–10 mm., including the stipe, which is jet black, shining, and much expanded at the base; hypothallus continuous, well-developed, a thin, transparent pellicle; columella black, tapering upward, giving off at intervals the capillitial branches, and becoming dissipatedjust below the obtuse apex; inner capillitial network very open, the branches far apart, anastomosing but a few times before breaking into the surface net to form large, irregular meshes, 50–125 µ; spores minutely roughened, fuscous, 8–9 µ.
These three forms, 8, 9, 10, are sometimes entered as varieties of a single species. Dr. Rex himself was inclined to take that view. There is no doubt of close similarity; it is a question of clearness in our dealing with the subject.
All three forms occur abundantly in the Mississippi Valley, but are generally,—always, as it seems to the writer,—distinguishable by the hand-lens. If we take No. 9 as type, 10 has an eccentric columella; 8 is shorter, about 1 cm., of a different tint, Dr. Rex even says "spores ferruginous in mass". To the west and southwest, the capillitium becomes coarser, more decidedly brown. In short, however similar in presentation the phases may sometimes appear, it would seem that each at its best is distinct enough for immediate recognition.
West of the Mississippi River chiefly: Iowa, South Dakota, Nebraska, Kansas, Oklahoma, New Mexico, etc.
9.Stemonitis splendensRost.
Plate VI., Figs. 6, 6a, 6c, 7, 7a.
Sporangia clustered irregularly, sometimes forming patches 6–10 centimetres or more in extent, rich purple-brown in mass, cylindric, long, 15–18 mm., stipitate; stipe black, polished, shining, rising from a common hypothallus, which extends as a thin silvery film beneath the entire colony, but does not usually transcend its limits; columella black, percurrent, sparingly branched; capillitium of fuscous threads, within forming a network very open, the branches scarcely anastomosing until they reach the surface where they form the usual net ofsmall meshes, pretty uniform in size, and presenting very few small, inconspicuous peridial processes; spores brown, very minutely warted, about 8 µ.
This elegant species occurs not rarely on rotten wood, usually in protected situations, although sometimes on the exposed surfaces of its habitat. The sporangia attain with us unusual height, sometimes 2 cm.; plasmodia, 6–8 cm., in diameter. The clear brown tufts appear in the autumn, marvels of graceful elegance and beauty; at sight easily recognizable by the large size and rich color. In Iowa it is almost universally present on fallen stems ofAcer saccharinumLinn., and it appears to be widely distributed, by far the most beautiful of all this beautiful series.
New England to Iowa, South Dakota, Washington, and British Columbia. Professor Shimek brings aduskyphase from Nicaragua!—the type?
The plasmodium is white on maple stems, more creamy on stems of linden, on which wood it is more rarely found: occasionally on ash-stumps; even on the fallen bark of trees preferred.
In 1875 in his famousMonograph, Rostafinski set out three species with "dusky violet spores". These are his Nos. 94, 95 and 96.
The first one of these he callsS. fusca, "spore-mass, etc., violet-black, individual spore clear violet, smooth, 7–9 u."
The second species he writes downS. dictyospora, "hypothallus, stalk, columella, capillitium and spore-mass, violet-black, spore netted and fringed, clear-violet, 7–9 µ."
The third species isS. splendens, "hypothallus stalk, columella and spore-mass violet-black, spore smooth, clear-violet, 7–8 µ."
It will be observed that in color down to color of the spore by transmitted light, the three species are exactly the same; constitute a suite, so to say. It has since turned out, as noted under our No. 3, that the spores ofS. fuscaare netted. Error in description here is not surprising; the reticulations are sometimes faint. InS. dictyosporathey are admittedly strong, and the inference was that the 'gladkie' spores of the third species might be netted also. This is no criticism: lenses were fifty years since not nearly so good for such discoveries as the oil-immersion is now.
However; Rostafinski made his specific diagnosis turn largely upon the mesh-width in the superficial net. This comes out in the 'opis' following the description, and uponthisthe European decision in Rostafinski's favor as againstS. morganilargely turns. Tropical gatherings are probably always darker, and evidently from such, from the north coast of South America, the original description was drawn. Specimens before us from the same latitude are dusky indeed; no clear brown at all, but purplish withal.
For the sake of harmony we may therefore now substitute the earlier name "with reservations"! but our description remains as before, presenting the really splendid, shining things that adorn our northern fields. Dr. Rostafinski called the large open meshes of the net 'oka', eyes;luminalet us say! quite uniform they are in 9 and 10, much less so in 8.
10.Stemonitis fenestrataRex.
Sporangia aggregated, in tufts 2 cm. or more in diameter, rich purple brown, on a common hypothallus, more or less erect, stipitate, tall, about 2 cm., slender, triangular in section; stipe black, about one-third the total height, passing into a slender columella which is lateral in position, not central, but little branched, continued almost to the apex; the capillitium consisting almost entirely of the peripheral net, which presents meshes of unusual uniformity of size and shape; spores in mass brown, colorless by transmitted light, nearly smooth, 6–7.5 µ.
The remarkable shape of the sporangium and the peculiar regularity of the surface net, the lateral columella, all combine here to warrant the erection of a distinct species. Dr. Rex referred this toS. bäuerliniiMass. At that time he had not the author's description, and had seen only a very poor fragment received with notes in a letter. Mr. Massee's description makes it immediately evident that whatever other affiliationsS. bäuerliniimay have, by description it has at least none withS. fenestratanor with our northern form ofS. splendens. Massee's species is described as having the "mass of spores black", the capillitium with "branches springing from the columella;the main branches more and more numerous, thicker and irregular towards the apex of the sporangium, and often form irregular flattened expansions":—etc. This suggests some form ofS. dictyosporaRost.: see under our No. 5. Possibly for such reasons Lister referred it toS. splendensRost., which as we have just seen, was undoubtedly regarded by the author as a form of thefuscousgroup.
The long, slender, simple columella is not only lateral, but occupies indeed the sharp vertical angle of the triangular, prismatic sporangium. Furthermore, the sporangium is at maturity strangely twisted, so that the columella in its ascent accomplishes one or more spiral turns. In forms collected by Dr. Rex, which seemed to him most nearly to agree with Massee's species, the inner capillitium is somewhat abundant, but the character of the columella just the same.
Pennsylvania, Ohio, Kansas, Colorado, Iowa; India!
11.Stemonitis smithiiMacbr.
Sporangia in small clusters, close-packed and erect, not spreading, bright ferruginous prior to spore dispersal, cylindric, stipitate, of varying height; stipe jet-black, shining, about one-third the total height; hypothallus generally well developed; columella black, gradually tapering, at length dissolving in capillitial threads and net some distance below the diminished plumose apex; capillitium of fuscous threads, the inner network of abundant, sparingly united branches uniformly thickened, the surface net very delicate, composed of small, regular, polygonal meshes, the peridial processes few; spore-mass bright ferruginous, spores by transmitted light pale, almost colorless, smooth, 4–5 µ.
The species as thus constituted includes forms varying in size from 2.5–3 mm. only. The common form heretofore known everywhere in America asS. ferrugineais from 10–15 mm. high. Thetypeto which the specific nameS. smithiiwas originally applied is 2.5 mm. high and rejoices in smooth, almost colorless spores, 4–5 µ.
The plasmodium in the case of the species now considered is asconcerns thetype, of course, unknown. In one or two gatherings referred here the color of the plasmodium was noted greenish-yellow. This has the look ofS. flavogenita; but small spores and delicate make-up take it the other way. Miss Lister makes it varietal to No. 12, next following.
12.Stemonitis axifera(Bull.)Macbr.
Plate VI., 5, 5a, and 5b.
Sporangia terete, acuminate, fasciculate small in dense clusters, distinctly ferruginous in color, stipitate, from 10–15 mm. in height; the stipe black one-third to one-half the total height, not shining or polished; columella evenly branching, dissipated before reaching the acuminate apex; capillitium-branches clear brown anastomosing and dividing more or less to bear the superficial fine-meshed net; spores pallid, faintly ferruginous, smooth or nearly so, 5–6 µ.
This would seem to be the commonferruginousspecies of the world. Doubtless Micheli had the thing before him when he drew Tab. 94,clathroidastrum, Hoffman and Jacquin seem to have recognized the form. To be sure, under the present plasmodic limitations we cannot be quite certain about these references. Not until 1791 does anyone write down a particular species as marked by a white plasmodium, and distinguish it from other similar fructifications having similar origin. Bulliard,l. c., does this, discriminating betweenT. axifera ferrugineaandC. typhoides; see under the last-named species. Youthful Ehrenberg, in his doctor's thesis, nearly thirty years later, draws a similar parallel but ignores the great French author, writingS. ferrugineaEhr. as though the thing had never been seen before! By this name it has been called until very lately; Fries accepting it, but noting that the plasmodium, for him at least, wasyellow!
In 1904 Dr. E. Jahn, following Fries' suggestion, established the fact that Ehrenberg's white-plasmodic species had small spores, thatFries had in mind a form with larger spores, having indeed yellow plasmodium; but see number 13 below.
It is for the present assumed that the plasmodium of our AmericanS. axiferais white. So far, there are few or no observations which establish the fact. The color, the small smooth spores, the fine-meshed capillitial net and the general dimensions determine the reference.
13.Stemonitis flavogenitaJahn.
Plate XX., Figs. 10, 10a, 10b.
Sporangia cylindric, obtuse, closely fasciculate, "cinnamon brown," stipitate, 5–7 µ; stipe short, black, columella ceasing abruptly below the apex; capillitium a loose net-work with many broad expansions; the peridial net very delicate, the meshes small but uneven, 6–15 µ, with many projecting points; spores pale ferruginous, verruculose, 7–9 µ.
This isS. ferrugineaEhr. of Fries with its plasmodium yellow. Fries says "flavicat,"becomesyellow, if one may follow the analogy of corresponding Latin verbs of color, so that the record of color-changes in the present species is yet to be recorded.
Until further experience may advise to the contrary, we may assume that all stemonites cinnamon-brown in color, with widened columella-tip, and pale yellowish spores 7–9 µ in diameter, have at some time in their history a yellow plasmodium, and accordingly represent in America the new-found species.
The larger spores, and, the strange proliferate development of the columella-tip, to which Miss Lister has happily called attention, constitute the essential diagnostic features here.
Our only specimens so far are from Oregon.
14.Stemonitis pallidaWingate.
Plate XIII., Fig. 3
Sporangia gregarious, or somewhat clustered, erect, cylindric obtuse, short, blackish brown, rubescent, becoming pallid, stipitate; stipe short, black, polished, rising from a thin, brown, or iridescent hypothallus; columella percurrent, ceasing abruptly at the apex; capillitium filling the interior with abundant branches which form at the surface a close-meshed net, little developed above, making the apex very blunt; spores in mass, dark brown, by transmitted light dusky, nearly smooth, 7.5 µ.
This species is well recognized at sight, among the fuscous forms, by its scattered, erect habit. In color it is not unlikeS. fusca, but has an added reddish tinge. In form it is peculiar by virtue of the blunt rounded apex which seems to be a constant character. The spores under moderate lens are perfectly smooth, under the 1–12 they present very delicate low scattered papillæ.
Rare; eastern part of United States.
15.Stemonitis carolinensisMacbr.
Plate XIII., Fig. 5.
Sporangia tufted in scattered clusters, small, slender, cylindric but tapering from the apex, at first ferruginous then ashen or purplish, stipitate; the stipe short, black and shining, one-fourth the total height or less, even; hypothallus well developed, black or very dark brown; columella black, gradually diminishing, at length dissipated some distance below the clavate or acuminate apex of the sporangium; capillitium dense, the inner of many, scarcely expanded, pallid, freely anastomosing branches, the outer a net of very small meshes, often less than the spores, 3–15 µ, peridial processes imperceptible; spore-mass pale ferruginous, spores by transmitted light pale violaceous brown, smooth, 6–7 µ.
Very closely related to the preceding, but recognizable by its proportionately much more slender, taller, acuminate sporangia, paler, and denser capillitium and the remarkably close-meshed net.
Not uncommon south: Kentucky, Alabama.
16.Stemonitis herbaticaPk.
Plate XVI., Figs. 14, 14a, 14b
Sporangia clustered, in scattered tufts, cylindric, obtuse, pallid ferruginous, stipitate or sometimes nearly sessile; stipe fuscous or jet-black, only slightly expanded below, much shorter than the columella; hypothallus scanty or none; columella lessening upward, sometimes attaining the apex of the sporangium, sometimes dissolved in capillitial threads some distance below; capillitium of rich brown threads forming the usual inner network of medium density, with many wide expanded nodes, the surface net made up of delicate, almost colorless threads surrounding small polygonal meshes; spore-mass ferruginous, spores by transmitted light very pale, brownish, minutely warted, 7–9 µ.
The plasmodium of this species is variously cited from white to yellow. Probably each report is true, dependent on the relative time of the observation.
The low tufts of brown sporangia with short black stipes, borne often as Dr. Peck found them, assembled on living leaves, distinguish this little species. In the former edition this form was tentatively enrolled underS. axifera(Bull.); but see further under that species.
Probably widely distributed, but confused with short forms of other species; sometimes also on rotten wood or other substratum; so reported.
New York to Iowa; Washington and Oregon. Reported also from Europe.
3. Comatricha(Preuss)Rost.
Sporangia cylindric or globose, stipitate; stipe prolonged upward to form a more or less extended and tapering columella bearing branches on every side, which by repeated divisions and reunions form the capillitium; ultimate branch-tips free, not supporting a surfacenet parallel to the peridial wall; peridium evanescent, perhaps sometimes not developed at all.
The genusComatrichawas set off fromStemonitisby the joint effort of Preuss (1851) and Rostafinski (1873–5). Preuss included in his genus,Comatricha, alien forms, and besides failed to give an accurate definition; included, however, in his list some species which have since been known by his generic name.
The distinction between the two genera is almost an artificial one, and species are sometimes arbitrarily assigned to one genus or the other. The diagnosis in any case turns upon the presence or absence of a surface net, formed, inStemonitis, by the anastomosing of the ultimate divisions of the capillitial branches. InComatrichathe anastomosing is general, from the columella out, and is not specialized at the surface.
Recent attempts to reunite the genera here compared seem to result in no apparent advantage. The genera come very near together, but their separation along the line suggested by Rostafinski remains convenient.
Key to the Species of Comatricha