OF ARTS AND SCIENCES. 
5 
II. 
ON SOME NORTH AMERICAN SPECIES OF 
LABOULBENIACEAE. 
By Roland Thaxter. 
Presented by W. G. Farlow, February 12,1890. 
Note. — The following is intended as a preliminary communication on Amer- 
ican Laboulbeniaceae, which it is my purpose to supplement as soon as practi- 
cable by a more extended account, to form the second part of a proposed 
monograph of Entomogenous Plants, of which the first, on the Entomophtho- 
reae of the United States* has already appeared. Illustrations of the new 
species described in this paper cannot well be given in the present connection, 
but will accompany my complete account of the subject when it is published. 
The Laboulbeniaceae constitute a small group of fungi, without close 
affinities among other known Ascomycetes, the members of which are 
remarkable both in their structure and development, as well as peculiar 
for their external parasitism upon insects of several orders. Up to the 
present time little has been recorded concerning them in this country, 
and the total number of forms at present known represent only fifteen 
described species. Of these twelve are European, distributed among 
five genera: Laboulbenia, Stigmntomyces, Helminthophana, Chitono- 
myces, and Hcematomyces. Of the remaining species, two are South 
American, belonging to the genus Laboulbenia, while the single rep- 
resentative of the group as yet recorded from this country has been 
described as Appendicularia entomophila Peck,f a genus, as will be 
presently noted, synonymous with Sligmatomyces Karsten. 
Our present knowledge of the group rests chiefly upon the writings 
of Robin, \ Karsten, § and Peyritsch, || the first notice of any member 
* Memoirs of the Boston Society of Natural History, Vol. IV. No. VI. (1888). 
t Thirty-eighth Rep. N. Y. State Mus. of Nat. Hist., p. 95, Plate III. figs. 1-4. 
1 Robin, Hist. Nat. d. Veget. Paras, etc., Paris, 1853, p. 622. 
§ Chemismus d. Pflanzenzelle, Wien, 1869, p. 78. 
|| Sitz. d. Kaiserl. Acad. d. Wissensch. Wien, (1871) LXIV Band, 1 Abth., 
p. 441; (1873) LXVIII Band, 1 Abth., p. 227; (1875) LXXII Band, 3 Abth. The undersigned is revising his mailing list and 
would be obliged for an acknowledgment of the enclosed 
by postal card if further papers of the writer are 
desired. 
R. TIIAXTER, New Haven, Conn. 6 
PROCEEDINGS OF THE AMERICAN ACADEMY 
of the family being apparently that of the entomologist Rouget.* The 
family has, moreover, received attention from a zoological point of 
view, the genus of Vermes, Arthrorhynchus, having been founded by 
Kolenati f on species parasitic upon Nycteribia (Dipterous parasites 
of bats). 
The members of the family may be briefly described as consisting 
usually of a main subclavate, flattened body, which for lack of a better 
term may be called the receptacle; simple, made up of a few large 
cells and bearing at its distal end one or more perithecia; or com- 
pound, the divisions bearing solitary perithecia. Within the perithe- 
cia, asci are developed by successive sprouting from basal initial cells. 
The asci contain apparently eight spores, although this number has 
been definitely observed in a single species only. The spores, which 
are hyaline, usually fusiform, once septate, and more or less involved 
in mucus, are expelled through the elastic apical pore of the peri- 
thecium, the ascus wall, as in other well known instances, disappear- 
ing before the discharge takes place. The spores become attached 
by one extremity to the surface of the host, and by subsequent division 
produce a new individual, without forming hyphse or a mycelium of 
any sort. The point by which the spore is attached becomes modified 
into a dark, horny-looking piece, which penetrates the chitinous integu- 
ment of the insect, and forms the single medium of nutrition as well 
as of attachment. 
In addition to the perithecium certain other bodips, bearing a defi- 
nite relation to it, are always present, borne usually on the receptacle 
close beside the perithecium ; in two genera (Helminthophana and Can- 
tharomyces) arising near the base of the receptacle. These bodies, or 
appendages, have been called paraphyses, or better pseudoparaphyses; 
and are of great morphological as well as systematic importance, vary- 
ing greatly in the different genera and species. Although certain of 
these appendages are usually sterile, there seems no reasonable doubt 
that one of them, at least, is always functional as an antheridium, or 
more commonly bears certain organs which are functional as antheridia. 
From these Karsten has observed the production of bodies which he 
considers antherozoids; but confirmatory observations, other than his 
own, are lacking. 
In Laboulbenia, the only genus which the writer has been able to 
examine with any thoroughness in its different stages, the supposed 
* Ann. d. 1. Soc. Entomol. d. France, Tom. VIII. p. 21 (1850), sec. Robin, 
t Wiener Entomol. Monatschrift, 1857, Band I. p. 66, sec. Peyritsch. OF ARTS AND SCIENCES. 
7 
antheridia are always bottle- or flask-shaped, and borne on the 
pseudoparaphysis next to the perithecium. These peculiar bodies, 
which appear to have been overlooked by Peyritsch, or whose impor- 
tance does not seem to have been appreciated by him, arise from the 
lower septae of the pseudoparaphyses, or are sometimes terminal on 
short branches. They are invariably present, as far as the writer’s 
observations indicate, just before fertilization takes place, disappearing 
as the perithecium matures; so that in the adult plant there is com- 
monly no vestige of them. 
Fertilization is accomplished through the medium of a body which 
must be considered a trichogyne, and is connected directly with the cell 
from which the asci subsequently arise. The character of the tricho- 
gyne varies in different genera, and in Laboulbenia may reach a re- 
markable degree of development, giving rise to branches, the tips of 
which may be coiled in a definitely spiral manner. The trichogyne 
is even more short-lived than the supposed antheridia, disappearing 
as soon as any development is observable in or about the central 
cell with which it is connected. Whether it is fertilized through the 
agency of minute round or oval bodies, frequently observed by the 
writer about the apices of the antheridia, more rarely within them, 
is quite uncertain, yet the observation's of Karsten would point to 
this conclusion. 
It may be mentioned that De Bary did not look upon the sexuality 
of these fungi with any favor, as may be inferred from his remark* 
that Peyritsch himself did not think very well of his (Peyritsch’s) 
“ attempt to save the trichogyne ” by supposing a fertilization through 
contact with one of the pseudoparaphyses in the genus Laboulbenia. 
That a form of sexual reproduction is present, however, among the 
species of Laboulbenia at least, cannot in the writer’s opinion be for 
a moment doubted by any one who has personally observed the more 
important stages of their development. 
The immediate affinities of these most singular plants are, as has 
been mentioned, very uncertain; and their resemblance to some of the 
higher Algae, through their supposed method of sexual reproduction, is 
striking and interesting in connection with the aquatic, or semi-aquatic, 
habit of many of them. That they are fungi, and at the same time 
Ascomycetous, seems beyond question. Why, therefore, they should 
be placed by De Bary and others among doubtful Ascomycetes, is not 
apparent. 
* Comp. Morphol. and Biol, of Fungi, etc., English ed., p 275. 8 
PROCEEDINGS OF THE AMERICAN ACADEMY 
In the following descriptions the term receptacle has been used to 
designate the main body of the fungus; the side from which the peri- 
thecium springs being spoken of as the inner, while that bearing the 
pseudoparaphyses is spoken of as the outer, where this distinction is 
possible. The first two superposed “ stem cells ” of the receptacle are 
spoken of as the basal and supra-basal cells. 
STIGMATOMYCES Karsten (1871). 
Stigmatomyces entomophila (Peck). 
Appendicularia entomophila Peck, 1. c. 
Appendiculina entomophila Berlese, Malpighia, Vol. III. p. 59. 
Through the kindness of its discoverer, the Rev. J. L. Zabriskie, the 
writer has been enabled to examine authentic specimens of this inter- 
esting and distinct species, which proves to belong without question to 
the present genus. The antheridial appendage is proportionately some- 
what smaller than in its near ally (S. Baeri (Knoch) Karst.), but has 
the same peculiar structure and bears the same relation to the peri- 
thecium; and structux*al differences which could separate it generically 
are wholly wanting. Berlese in the paper referred to calls attention 
to the fact that Appendicidaria has been preoccupied among the Me- 
lastomaceae, and proposes the substitution of Appendiculina, which, 
however, proves to be needless. 
PEYRITSCHIELLA nov. gen. 
Receptacle composed of two superposed basal cells, above which 
it is multicellular, one cell on the inner side forming a short, sharp 
projection. Perithecium one, sometimes two; when single, terminal, 
nearly median, subconical, the spreading apex symmetrically four- 
lobed. Pseudoparaphyses arising from several different points on 
either side of the receptacle. 
Peyritschiella curvata nov. sp. 
Characters of the genus. Usually strongly curved, colorless except 
the large, jet-black piece of attachment, and the bases of the pseudo- 
paraphyses which are also black and strongly constricted in the 
middle. Paraphyses colorless, cylindrical or subclavate, septate or 
obscurely septate; arising in three to five groups, each made up of 
from one to three pseudoparaphyses, and placed alternately at different OP' arts and sciences. 
9 
points on either side of the receptacle. Total length to tip of perithe- 
cium 280-300 /*. Perithecium 90-100 n X 22-29 /x. Spores fusiform, 
asymmetrically once septate, involved in mucus, 26 X 4 /x. Pseudo- 
paraphyses sometimes 60 n in length. 
Host, Platynus cincticollis. Connecticut. 
This genus appears to differ from other known genera in the pres- 
ence of several groups of paraphyses on both sides of the receptacle. 
The presence of two perithecia was only observed in a single speci- 
men. The perithecium resembles that of Helminthophana in being 
symmetrical at the apex. The genus is named in memory of the late 
Dr. Peyritsch, as a slight recognition of his well known and admirable 
work upon this family. 
CANTHAROMYCES nov. gen. 
Receptacle simple; or compound above the supra-basal cell, from 
which one or more divisions may arise, each bearing a solitary perithe- 
cium. Pseudoparaphyses, one or more, arising from the supra-basal 
cell. Perithecium median, tapering towards its symmetrical apex. 
Cantharomyces verticileata nov. sp. 
Color paie yellowish. Perithecium expanding slightly above its 
base, then tapering slowly towards its slightly truncate, conical apex. 
Pseudoparaphyses arising in a whorl of two to four (the number ap- 
parently variable) from the small supra-basal cell; simple or branched; 
once or twice constricted at the septae; the joints short, stout, and 
broader towards their rounded apices. A single simple pseudopara- 
phvsis composed of cylindrical segments hardly constricted at the 
septte arises also from the distal end of the third stem cell. Recep- 
tacle simple, composed of five single superposed cells, arising from a 
small black piece of attachment, the first four nearly cylindrical, the 
fifth short, slightly expanded towards the base of the perithecium, 
which is made up of two or three very small cells. The supra-basal 
cell is squarish about half the length of the basal. The third and 
fourth cells are nearly similar, somewhat longer than the basal and 
supra-basal taken together. Spores fusiform, apparently once septate, 
and about 18-20 p, X 3 p. (these were only examined, however, within 
the perithecium). Perithecia 90-127 /x X 18-26 /x. Pseudoparaphyses 
50-125 /x. Receptacle 75-125 jx X 11 /x. 
On Sunius longiusculus. Anna, 111. (S. A. Forbes). 10 
PROCEEDINGS OF THE AMERICAN ACADEMY 
Cantharomyces Bl/dii nov. sp. 
Color yellowish. Perithecia subconical, one or more in number, 
each borne at the summit of a division of the receptacle arising from the 
supra-basal cell. Pseudoparapbyses one, rarely more, arising from the 
supra-basal cell: composed of three superposed basal cells, the second 
swollen, longitudinally septate; the third squarish, small, surmounted 
by one or two small cells, from which arise a variable number of 
slightly curved, septate, slender branches themselves variably once or 
twice branched. Receptacle simple or compound ; the basal and sub- 
basal cells rather short, the lower and outer portions of the wall of 
the latter often thickened, deep black, and indented; but sometimes 
without signs of this modification. The supra-basal cell may give rise 
to one or more divisions of the receptacle, which are sub-lateral in 
position, and consist of a long, cylindrical basal cell, surmounted by a 
broader short cell, divided longitudinally by a median septum, and 
bearing the perithecium directly. Spores slender, fusiform, involved 
in mucus, asymmetrically once septate, 25 X 3.5 p. Perithecia 92.5- 
130 /x X 33-55 n, average 114 x 42 fi. Pseudoparaphyses, total 
length 90-180 /*, average 150 /x. Total length to tip of perithecia 
200-370 /x, average 280 /x. 
On Blfdius assimilis. Champaign, 111. (S. A. Forbes). 
I am greatly indebted to the kindness of Professor Forbes for speci- 
mens of the two singular forms above described. The material of 
C. verticillata consisted of but four adult specimens, so that further 
study may show the presence of a compound receptacle in this species 
as well as in C. Bljdii, where it was observed in only a few of the 
twenty or more individuals examined. The secondary pseudopara- 
pliysis of C. verticillata is wholly wanting in C. Blfdii. 
LABOULBENIA Montagne et Robin (1853). 
Color brown or blackish. Perithecium long-ovoid, darker just be- 
low its hyaline apical pore. Pseudoparaphyses arising from a black 
basal disk; two in number, brown to blackish, sometimes hyaline near 
their extremities, septate, slightly constricted at the septm; their basal 
cells distinct, the inner giving rise immediately to two branches which 
may be either simple or once or twice dichotomously branched. The 
outer pseudoparaphysis usually once or twice dichotomously branched 
Laboulbenia elongata nov. sp. OF ARTS AND SCIENCES. 
11 
above the supra-basal cell. Receptacle composed of seven main cells; 
the basal nearly triangular, the supra-basal elongate, cylindrical, often 
three times as long as the basal. Above the supra-basal cell the re- 
ceptacle is divided by a longitudinal partition into two cells, that on 
the inner side smaller, and separated from the base of the perithecium 
by a single, usually squarish cell, above which are two small triangular 
cells, an outer and an inner, sometimes connected, which form the base 
of the perithecium. The two remaining cells of the receptacle bear 
the disk-like base of the pseudoparaphyses, and consist of an outer 
larger one, separated by an oblique partition, on its upper inner side, 
from the smaller inner one. Spores fusiform, hyaline, granular, in- 
volved in mucus, septate near one extremity, 75-80 jx X 6.5-8 /x. 
Perithecia 135-170 /x X 65-70 /x. Pseudoparaphyses 200-725 fx. 
Length to tip of perithecium 450-600 /x ; average 538 /i. Maximum 
total length to tip of pseudoparaphyses 950 fx. 
On Platynus cincticollis. Connecticut. 
This species, which is the largest known representative of the ge- 
nus, being easily visible to the naked eye, is allied to L. Nebrice and 
L. Jiagellata, resembling the latter in appearance, but easily separable 
from it by its great size and the details of its structure. Its trycho- 
gyne is remarkable for its unusual development, being often several 
times branched, and otherwise peculiar from the spiral twisting of one 
or more of its ultimate branches. 
Laboulbenia brachiata nov. sp. 
Color brown to blackish. Perithecia long-ovoid, darker towards 
the hyaline apex. Pseudoparaphyses blackish, or nearly hyaline; 
each on a small black basal disk; arising from an oblique cellular 
base; ten in number in a double row of five pairs; septate, the 
basal and sub-basal joints often inflated, especially in the outer larger 
pairs; dichotomously once to twice branched above the basal and 
sub-basal joints, the ultimate branches often1 very long and attenu- 
ated. Receptacle consisting of single basal and supra-basal cells, the 
latter slightly the longest, above which are two cells, that on the 
inner side the largest: above and between these two cells is a third 
smaller, central one, and on the inner side of this another, slightly 
larger cell, is separated from the perithecium by two small flat cells 
which form its base and are sometimes confluent. The remaining 
cells of the receptacle form the base from which the pseudoparaphyses 
arise, which is made up of five obliquely superposed cells, decreasing 
in size from below upwards, and separated from the pseudoparaphyses 12 
PROCEEDINGS OP THE AMERICAN ACADEMY 
by a row of small cells from which the latter spring directly. Spores 
hyaline, fusiform, septate near one extremity, surrounded by a gelati- 
nous envelope, 60 X 5 p. Perithecia 150-120 p X 50-60 p. Para- 
physes, short 150-180 p; long 700-740 p. Total length to tip of 
perithecium 400-450 p. Maximum length to tip of paraphyses 950 p. 
On Patrobus longicornis. Connecticut; (Maine, Dr. Townsend). 
A remarkable form, nearly if not quite as large as the preceding 
species, and resembling a minute crustacean from its many projecting 
appendages. It is allied in the structure of its receptacle to the species 
figured by Peyritsch (1. c. 1873, Plate I. fig. 9), and referred by him to 
L. fasciculata ; a manifest error, as a glance at the mature plant (fig. 8) 
must show. 
Laboulbenia Rougetii Mont, et Robin. 
Laboulbenia Rougetii Mont, et Robin, Hist. Nat. d. Veg. Paras., p. 622, 
Plate X. fig. 2. 
To this species I have referred a form occurring on Platynus, which 
corresponds so closely to Robin’s figures that 1 am inclined, for the 
present, to consider it merely a variety of this species. A comparison 
of more extensive material than I possess may show that it is distinct; 
yet the early stages, as well as the mature individuals, show such 
slight variations from the figures of Rovgetii that it would be unsafe, 
at present, to refer it to any other species. 
On Platinus cincticollis. Connecticut. 
Color smoky brown with an olive tinge. Perithecium long-ovoid or 
ovoid, the apex with its hyaline pore often truncate, hardly oblique. 
Pseudoparaphyses arising from a base of three or four cells, itself 
seated on a thin black disk. Pseudoparaphyses arising from three or 
four inner basal cells, the walls of which are not distinct. From the 
outer of these basal cells extends a row of cells curving outwards and 
downwards (this portion is commonly broken off in specimens), black 
beneath, and giving rise to a variable number of straight, erect, sim- 
ple, colorless branches. The inner basal cells give rise to numerous 
branches, erect, slender, septate, colorless, sometimes ten in number, 
often twice as long as the perithecium; the shorter, inner branches 
sometimes subclavate. Receptacle broad, tapering somewhat abruptly 
to its usually slender basal cell; supra-basal cell larger, broad: above 
it, on the inner side, two cells which separate it from the perithecium, 
which is itself seated on two or three small cells; above it, on the 
Laboulbenia fumosa nov. sp. OF ARTS AND SCIENCES. 
13 
outer side, two larger cells, which separate it from the cellular base 
from which the pseudoparaphyses arise; the upper of these two cells 
divided by a curved partition wall, which cuts off its upper inner cor- 
ner. Spores fusiform, septate near one extremity, involved in mucus, 
55-60 /x X 5-5.5 fx. Perithecia 120-130 fx X 50-75 fx. Pseudo- 
paraphyses (longer) 75-230 jx. Length to tip of perithecium 250- 
300 /x. Greatest width 75-100 /x. 
On Platynus cincticollis. Connecticut. 
A small species occurring almost invariably at the apex of the elytra, 
rarely on the abdomen of its host. It is nearly related to L. luxurians, 
from which it is readily distinguished by its long, straight pseudo- 
paraphyses. 
Laboulbenia IIarpali nov. sp. 
Hyaline or very slightly straw-colored. Perithecia long-ovoid, 
sometimes tinged with brown, the apex (in mature specimens) black, 
except about the pore; longitudinal axis nearly parallel with that of 
the receptacle. Pseudoparaphyses hyaline, arising from a black basal 
disk placed opposite the middle of the perithecium; two in number, 
their basal cells connected ; the outer irregularly once or twice dichoto- 
mously branched; the inner usually consisting of two short branches 
arising directly from the inner basal cell. Receptacle slender; con- 
sisting of two large stem cells (basal and supra-basal), and above 
them*on the inner side a subspherical cell is separated from the peri- 
thecium by three very small cells. On the outer side the two cells 
below the pseudoparaphyses are present, the upper having the usual 
oblique partition in its upper inner angle. Spores fusiform, septate 
near one extremity, involved in mucus, 60-68 fx X 5-5.5 tx. Length 
to tip of perithecium 290 fx (215-300/*). Length of outer pseudo- 
paraphyses 200-300 fx; of inner 100-130 p. Perithecia 90 X 40 fx. 
On Harpalus Pennsylvanicus. Connecticut; Maine (Kittery). 
This species occurs upon the inferior surface of its host, always on 
the left side of the anterior, inferior face of the abdomen, sometimes 
extending across to the adjacent portion of the thorax. It is not un- 
eommon, and occurs frequently in company with the succeeding species, 
never mixed with it, however. 
Laboulbenia elegans nov. sp. 
Hyaline. Perithecia ovoid; the longitudinal axis at an angle of 
about 40 to that of the receptacle; apex jet-black (only in mature 
specimens), except around the pore. Pseudoparaphyses arising from 14 
PROCEEDINGS OF THE AMERICAN ACADEMY 
a jet-black disk, nearly opposite the base of the perithecium; two in 
number, their basal cells connected. The outer basal cell giving rise 
to two branches ; an inner, once dichotomously branched, and an outer, 
always simple. The inner basal cell very small, giving rise to sev- 
eral short branches, slightly curved, often subclavate, sometimes once 
branched, bearing numerous lateral or terminal antheridia. Recepta- 
cle similar to that of the preceding species, the cells merely varying in 
their relative proportions. Spores fusiform, septate near one extrem- 
ity, involved in mucus, 40-50 y. X 4-5 fx. Length to tip of perithe- 
cium 290 fx. Length of outer pseudoparaphyses 250-400 /x; of 
inner 50-75 fx. Peritheeia 110-130 fx X 50-65 fx. 
On Harpalus Pennsyloanicus. Connecticut; Maine (Kittery). 
A very pretty and distinct species, allied to the preceding; but sep- 
arable at once by the position and character of its pseudoparaphyses. 
It is the only species observed by the writer, in which the sterile 
pseudoparaphysis appears to be invariable in its mode of branching. 
The species occurs in single tufts on the inferior surface of the thorax 
of its host, always on the right side.