STRUCTURE AND SYSTEMATIC POSITION O F CHANTRANSIh.

209

On the Structure and Systematic Position of Chantransia ; with

a Description of a New Species. By GEORGE MURRAY and
ETHEL S. BARTON,
[Read 5th June, 1890.1

XXXVI. & XXXVII.)

(PLATES

THEgenus Chantramia has been, since its establishment, one of

the most interesting among Algae, not only on account of its re-
markable position as one of the so-called primitive types of 3'10-
rideae, but also from a supposed relationship ascribed to certain
of its forms with Batrachospermurn and Lentanea. The history
of the genus is, i n fact, an exceptionally stormy one j its syste-
matic position has been the subject of argument, the generic
characters have been emended and its validity as a genus haa been
challenged. It is happily unnecessary for us to penetrate farther
into the history of these changes than the Thuretian conception
of the genus. This author writing (1868) in ' Le Jolis, Algues
Marines de Cherbourg,' p. 104, says :-
" The gems Chantransia has not been kept to the limits assigned
t o it by De Candolle; since it contained species belonging to
Lemanea, Batrachospermzmrn, Cladophora, and Q3dogonitl.m (Fl. Fr.
ii. p. 49 et sep.). Fries re-established it later (1825) o n a better
basis, taking for its types Conferva Hernzanni and C. chalybeia of
Roth. These two plants, living in freshwater, bear a strong re-
semblance to certain minute marine species which form part of
the Callithamnion of Lyngbye. Harvey long ago recoguized this
resemblance, which had compelled him in 1836 to re-unite the
marine and freshwater species under the genus Trentepohlia
(Mackay, Flor. Hibern.). More recently, however, this celebrated
algologist has changebhis opinion and replaced the marine species
in Callithamnion. H e has even described and figured for some
of them veritable tetraspores (Phyc. Brit. tabb. 313, 314). The
existence of such organs would justify the position which he
assigns to these plants if it were well demonstrated. But I think
that there is some error here resulting from the study of dried
specimens. A t least I have never been able to find tetraspores,
and I trust implicitly on this point to the excellent observations
of M. Areschoug (Phyc. Scand. marin. p. IlS), confirmed recently
210 XlL Q. MURRAY AND MISS E. 9. BARTON ON TEE

by M. Pringsheim (Beitr. z u r Morphol. d. Meeres-Algen, p. 26).

I have never seen iu these plants anything but undivided spores,
having a teridency more or less to group themselves on short
lateral branches, in such a manner as to form in certain species
small glomerules, sometimes replaced by clusters of autherids.
That these glomerules or groups of spores represent the most
simple state of conceptacular fructification of the other Florideae,
one can hardly doubt, especially when one compares them with
organs of the same nature in other genera j &c.”
Though Harvey was wrong as to the division of the spores iiito
tetmspores, he was nevertheless right in his view o f the morpho-
logical value of these bodies. They are commonly called mono-
spores, and understood to be the homologues of the tetraspores
of the other Floridere. That Thuret, though right in his obser-
vation, u as wrong in regarding them as equivalent to concepta-
cular fruits, was made abundantly clear by the description of the
true cystocarps of Chuntransia corymbifern by Bornet and Thuret,
‘ Notes Algologiques,’ p. 16, tab. v. (1876). Before this, how-
ever, in 1873, M. Sirodot described the monospores, antherids,
tricbogynes, and cystocarps of a freshwater species, Ch. investirns,
Lenorm. (Comptes Rendus, vol. luxvi. pp. 1338-39). M. Sirodot,
however, thought fit to remove this species from Chantransia and
make it the type o f a new genus, Balbiania (Ann. Sci. Kat. 1876,
Be s6r. tom. ii. p. 146). Whether this distinction should now
stand will be seen later. It is mentioned here now because the
discovery of its sexual reproductive organs takes precedence of
the case of Ch. corynabifra.
As the matter now stands, the position of the genus in the
estimation of botanists is as follows :-In the sea there occur
certain species typified by Ch. corymbifera, Thur., o f which both
the sexual reproductive system and the propagation by mono-
spores have been described and figured by Bornet (Zoc. cit.). In
fresh water there occur certain other species (excepting for the
present Ch. inveStiens= Balbiania investiens, Sir.), which axe re-
garded by M. Sirodot as non-sexual forms of Batraciospernaum.
His views on this subject are most fully expounded in his elabo-
rate treatise, ‘ Les Batrachosperrues,’ Paris, 1884. According to
him these “ Chantransia-forms ” are the sporophytes of Batracio-
spermurn,-fornis which do not attain a sexual reproduction unless
in the shape of Batrachospermum-his own discovery of antherids
and cystocarps in Ch. investiens having been swept from thetrack
 s T a ~ AND
~ ~SYSTEMATIC
~ ~ E P O S I T I O N 08 CHAKTRmSTA. 211
of this theory by the removal of the species in to Balbiania created
for its reception.
It is well known that the carpospore of Lenianea on germi-
nating puts forth a so-called protonema1 filament on w1hich the
fertile axes are borne ; and it has been asserted by Peter (Bot.
yerein, Miinchen, 28 Feb. 1887) that the sexual Lenianea$uvia-
tilis may develop from the heteromorptiic braiiches of a C?tan-
iransia. Sirodot had also ('Les Batrachosperm.' p. 4) made a
silnilar assertion as to the connexion of Ch. violacea and Ch.
amethystea with Lr-manea.
I n the ' Annals of Botany ' (vol. iv. no. xiv., May 1890) there
is a paper by Prof. atkinson on " The Lemaneaces of the United
States," in which the sD-called " Chantransia-forms " of Lenianea
are described. According to this author also the " Ciiantransia-
forms '' in question do not produce monospores, but stand in the
relation of a protorreme to Lemanea.
In addition to this there are it small number of species of
Chantransiar growing in fresh water and described in systematic
books which have had no definite position or relation assigned to
them.
Such, then, is the extraordinary position of this genus. The
marine forms only are perhaps regarded as valid, but exist merely
on sufferance. The freshwater forms are some of them sporo-
phytes of one plant or protonemes of another, while a residuum
remains unattached to anything at present in the limbo of syste-
matic papers on freshwater A l p .
The principal material of the present research was collected by
Prof. Bower and Mr. Murray, in the beginning of last April, in
the stream near Duntocher, running out of Loch CocLno on the
Kilpatrick Hills, Dumbartonshire. The Chantransia R xs growing
on LenianeaJluviatilis in minute tufts, rooting in the tissues of
its host. On comparison with other species of Chantransia,
notably with Ch. violacea, it M as found to be new to science, and
we have therefore described it below as Chantransia Boweri in
hoilour of its finder. The separate filaments grow to about
1 inillim. in length, and the brauches, which are always giveu off
at the upper end of the cells, are either opposite or alternate in-
definitely. The branches and twigs end each in a loug h p l i n e
hair ivith a slightly rounded tip. The lower cells of a filament
are 3-4 times, those occupying the middle portion 4-5 times as
long as the diauieter ; while the cells at the tip diminish in length
212 MR. Q. M U B U Y AND MISS E. 5. BARTON ON THE

to about twice the diameter. The cell-walls are thinner in the

apical than in the basal cells. The filaments are attached to the
Lernanea by means of non-septate and much-coiled rhizoids.
Sessile monospores are borne in great abundance on the branch-
lets generally opposite in twos, often in threes. Such clusters
occur at the ends of the branches, sometimes with the terminal
hyaline hair, sometimes without it. They are rather pyriform
than oval in shape, and of a deeper colour owing to denser
contents than the cells of the filaments. When the branches
are in full-bearing, monospores terminate nearly every twig aud
the hyaline hairs are p l y rarely to be met with (Plate XXXVI.
fig. 1).
We have not been able to observe the escape of these monc-
spores, since the material was preserved in alcohol before the mi-
croscopic examination took place. They are, however, not simply
detached, since after the escape there remain the empty spore-
cases in sitzc (figs. 4 and 5). This spore-case or outer wall is
apiculate and hyaline, while the inner one is darker in colour.
From the fact that traces of empty spore-cases sometimes
appear round the base of monospores, it would appear that new
outgrowths occur through these old spore-cases (figs. 4 and 5 ) ,
as in the unilocular sporanges of Cladostephus. W e must regard
thede monospores as produced in monosporanges (= spore-cases),
the homologues of the tetrasporanges of other Florides *.
The above description represents the usual appearance of the
plant as collected. Patient examinatiori of it, however, was
rewarded by other discoveries, viz. antherids, trichogynes, and
cystocarps. The autherids (Pl. XXXVII. fig. 4) resemble, as will
be seen by the figures, most closely those of Ch. corymbifera;
they form dense clusters, and each pollinoid is about two-thirds
the diameter of the ad,jacent filament in size. The cystocarps
(Pl. X X X V l I figs. 2 and 3) form corymbose stalked clusters of
carpospores which are in size about twice the diameter of the
filaments which bear them. It will be seen from the illus-

* As further confirming this view, i t may be added that, in the course of ob-
servations on it marine species, Chantransia secundata, we have observed its early
stages of germination on Cladophora nyestris. The monospore first divides
intofmir, and then SO closely resembles a tetraspore that the acceptance of this
view as to its homology becomes irresistible (PI. XXXVII.fig. 5). This division
then proceeds in the same plane, thus giving rise to the membranous base of
C%awtransiu,from which the upright filaments arise.
 ~ ~ SYSTEMATIC POSITION O F CHbNTBANSIA.
s T R U C T ~AND 213
Bations that the development of the cystocarp aIso resembles
ch. corym6ij&.a, the marine species, as described by Bornet.
Owing to the scarcity of material in this condition, we were
unable to investigate this process of development more fully.
The tri&ogynes, so far as we have observed them, rather re-
sen,ble tile figure of Ch. corymbifera by Schmits (" Unters. ii.d.
Befrucht. d. Florid.," in the Sitzungsber. d. Berlin. Akad. 1883,
plate v. figs. 2, 3, & 4) than those by Bornet. The observer is
constantly misled by the appearance of clusters of monosporanges
surrounding an emergent hyaline hair into the belief that he is
witnessing a young cystocarp crowned by a trichogyne. Such
clusters have a wonderful superficial resemblance to the young
cystocarps of Nenaalion for example, though in Ch. Boweri the
greater size of the carpospores enables one to detect them at once
when accompanied in the same field by monosporanges. Besides
more essential differences, the carpospores are of course in denser
clusters and greater numbers in each cluster.
The folloxing diagnosis of the species sums up briefly its
characters :-
CHANTRANMA BOWER~, n. sp. ; cespite minuto, pallide violaceo ;
filia radiatim diapositis .0085 mm. crassis, articulis quam diametro
inferioribus 3-4~10,superioribus 4-5~10, supremis duplo longi-
oribus, rarnulis apicibus piliferis, oppositis interdum irregularibus ;
monosporslngiis apiculatis, monospork ovalibus aut subpyri-
formibus sessilibus, oppositis binis aut ternis ; cystocarpiis et
antheridiis corpmbosis, pedicellatis.
Ad LeinarzearnJEuviatilern in rivulis prope Duntocher, montibus
Elpatrick, com. Dumbarton, Scotia ; Iegerunt Bower et Mumay,
die dominica paschali April 1890.
The nearest species, Ch. aiolacea, which also grows on Zemanea,
difers from it (1) in the absence of the long hyaline hair at the
end of the branches, and (2) the different proportions of the
joints, and (3) in the thicker cell-walls. These combine to give
it a quite different appearance *.
* I n examining species of Chantransia the student should be warned against
the very deceptive appearance presented by the epiphytic species of Democurpa
and the like. Not only in the case of Chuntransia violaceu in fresh water, but
in cha7atrunsiu secundata in the sea, we have been temporarily misled several
times by the extraordinarily close resemblance borne by clusters of Democurpa
its spores to both cystocarps and antherids. Minute and careful study and
ConlParlsonalone enables one to avoid mistake in this matter.
214 MB. 0. MUBRAY AND MISS E. S. BARTON ON ‘PEE

The supposed relationship of “Chantransia-forms ” with Lema-

neacerc has been mentioned already, and it now becomes necessary
to examine this question more carefully in the light of the facts
set forth. More especially is this the case, since Ch. Boweri
and Ch. violacea grow on the thallus of Lemanea. I n Prof.
Atkinson’s paper, p. 222, Chantransia violacea var. Beardski,
Wolle, is quoted under Lemanea .fucina, Bory ; and in a foot-
note to this he says, ‘L This is the Chanfransia-form of Lemanea
(Sacheria) fucina, Bory, var. rigida, which Wolle found ‘ as an
undergron th, intermingled with Lemanea, which was fringed
with the parasitic C. violacea,’ from Painsville, Ohio.” Prof.
Atkinson has figured this “Chantransia-form of Lemaneafucina ”
on plate vii. figs. 6 and 10. His meaning is therefore plain;
though it is difficult to understand how he ever came to reach
it. Not only is his “Chantransia-form” many times larger than
Ch. violacea, but in much more important respects the resem-
blance is sadly to seek. Kutzing’s imperfect figure of his own
species is probably the origin of Prof. Atkinson’s mistake ; but
whatever var. Beardslei may be, it is certainly not a variety of
Ch. violacea. More than this, it is necessary for us to prove
that it is no true Chantransia at all. I n calling it so, Prof.
Atkinson of course but follows the example of Sirodot, Peter,
and others in associating what appears to be a Chantransia
with a Lemanea. It will be remembered that Sirodot’s ‘‘Cham-
transia-forms ” of Butrachospermum are sporophy tes, but that
the “Chantransia-forms ” of Lenaanea are protonemal merely, and
bear no monospores. But Ch. violacea bears monospores.
Next, let us take Ch. Boweri. Not only have we described
its monospores, but its sexual organs of reproduction as well;
and it, too, grows on Lemanea. It is of importance to state
here that ~e actually obtained in tbe same preparation, on the
same slide, a t one time the antherids of ChaBtransia Boweri
and the antherids of Lemncinea juviatilis on which it grows.
There could be no possible excuse for mistaking the one for
the other. This interesting observation not only disposes of
any question of identity between Ch. Boweri and Lemanea, but
it is fatal to any ingenious theory to the effect that sexual repro-
ductive organs might possibly be borne on the protonemal form
under abnormal circumstances. Here, under the same circum-
stances, side by side, were these two plants both bearing an-
theiids, a i d these antherids of a different type. It must be
 STRUCTURE AND BYSTEMATIC POSITIOi4 OF CHANTEANSIA. 215
clearly understood that we do not call in question the existence
of a protonemal form of Lemanea which resembles, in a super-
ficial way, Ckantransia ; nor do we contend that Peter or Prof.
Atkirison are mistaken in their interesting observations except
in this, that their protcnemal forms are species of Chzntransia.
Enough has been said t o show that this has been too hastily
aesumed. One might as reasonably call the protonemc of a moss
the Conferva-form. This confusion of a protonemal form of
Lemanea with Chantransia has thus been rendered worse con-
founded by the fact of true species of Chantransia growing on
Lenzanea.
The question of the asserted relationship between CJLanfransia
and Batrackospermunz is a much more difficult one; and our ob-
servations do not directly touch it. The so-called “Ckantmnsia-
forms ” bear non-sexual spores. Are these true monosporange8 or
not ? As bearing, however, most weightily on this question, we
may here claim to hava established a freshwater group of sl)ecies
of Chantransia, consisting of Ch. Boweri and Ch. investiens (the
genus Balbiania scarcely possesses validity), in all generic poiuts
resembling the marine species Ch. corynzbifera. They are not
only reproduced non-sexually by monospores, but sexually as
well. There is therefore a good and valid genus Chantransicc
in fresh aater as well as in the sea ; and it may be added (though
a small matter, yet an indicative one) that these freshwater
species exhibit that form of growth called innovation, which
Mr. Harvey Gibson tells US he has observed in the marine genus
Rhodockorton, SO nearly related to the marine Chantransin.
This being so, it appears to U S that the burden of proof (in this
matter of Batrachospermzcnt and Ckantransia] is shifted from o u r
shoulders to those of’ M. Sirodot and his supporters-the required
proof being that his “Chantransia-fornis ” are anything more than
sporophytic shoots of Batrachospermum resembling Chantransia.
W e haye seen how it has fared with the “Chantransia-forms ” of
Lenzanea. It is open, of course, to those who prefer it, to contend
that the sexual reproductive organs of Ch. investiens are merely
the result of abnormal circumstances operating on a form which
orditlarily is a sporophytic condition of Batrachospermunz, for
example. Here, again, let US take warning by the case o f Lenianea.
Further, the observation of these was made by M. Sirodot
him self.
It would be also a possible contention that Ch. Boweri ought
216 STRUCTURE AND SYSTEMATIC POSITION OB 0HANTRA.NSIA.

t o be reckoned with Balbiania ; but this genus was created, its

author tells us, for the reception of this form, since all other
“Chuntransia-forms ” were mere sporophytic states of Batracho-
spermurn. I t must, therefore, now disappear. Moreover, the
obvious c h s e connexion existing with Ch. corymbifera warrants
us iu disregarding Balbinnia as a genus, and reckoning Ch. inves-
tieas and Ch. Boweri, with Ch. corymbifera and other marine forms,
as all of them species of B good and valid genus Chantransia.

EXPLANATION OF THE PLATES.

PLATE
XXXVI.
Chantramia Boweri.
Fig. 1. Filament bearing monosporanges, x 500.
2. Ditto, showing branches with piliferous endings, X 900.
3. Portion of filament, x 900.
4 & 5. Ditto, showing innovation, X 900.

XXXVII.
PLATE
Fig. 1. Trichogynes of Ch. Boweri, x 900.
2. Development of cystocarp of ditto, x 900.
3. Mature cyatocarp of ditto, x 900.
4. Antherid of ditto, X 900.
5. Germinating monospores of Ch. secundata, ~ 9 0 0 .
6. Antherid of Ch. coyrnb;lfera, x 250. After Bornet.
7. Trichogyne of ditto, x 400. After Bornet.
8. Cydocarp of ditto, x 250. After Bornet.
9. Development of cystocarp of Ch. inoestiens, X 720. After
Sirodot.