The Scottish botanical review

Vol. I. No. 4 1912 OCTOBER

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The
Scottish Botanical Review

None pean LOL | OcTOBER

Morphological Notes. By K. von Goebel,
pe-b). (Cami.), IL. D. (Se_Amd.);,

Director of the Botanic Garden, Munich.
(With One Plate.)

fog INEEORESCENCES OF THE
AMBROSIACE.

THE large family of the Composite, as is well known, is one
of those groups whose individual representatives, in spite of
all their variation, still show such a close agreement in the
structure of the flowers and the inflorescences that one can
offhand recognise them as members of one family. Only a
few groups deviate to any great extent, and of these one of
the most interesting is the Ambrosiacez.

Ecologically they are interesting in the manner in which
pollination on the one hand and the distribution of the fruit
on the other hand are accomplished among them.

The Composite as a rule exhibit insect-pollination, but
the Ambrosiacez have reverted to wind-pollination.

Hardly any greater contrast can be presented than between
the resplendent flower-head of a Helzanthus or a Dahlia, with
its remarkable arrangements for dehiscence and for trans-
ference of the pollen, and the insignificant male and female
floral attire of a Xanthium or an Ambrosia. There is a clear
indication here that, quite apart from the lack of a corolla
conspicuous in its coloration, the structure of the floral
organs is in strict correlation with their functions. In the

VOL. I. 193 15

194 THE SCOTTISH BOTANICAL REVIEW

flowers no nectar is formed, there is no ‘‘concrescence’’ (in
reality it is only agglutination or sticking together) of the
anthers, and the pollen is formed in great quantity, as is the
case with other anemophilous flowers. The condition of the
exine indicates, however, that originally pollination took
place by insects as in other Composite.

It is well known that the pollen in entomophilous plants is
provided with a spiny or sticky exine, whereas in anemo-
philous plants the exine of the pollen-grains is smooth and
not sticky. The Ambrosiacez, so far as I have been able to
examine them, still show in the exine the remains of spinose
thickenings such as are found in numerous other Composite,
but these are so reduced that the pollen is no longer adapted
for adhering.

The male flowers still show on the rudiment of the stigma
a hairiness which recalls the “stigmatic brush’’ of other
Composite. In the female flowers, in which the stamens
have dwindled without leaving a trace, there is no longer
any indication of this stigmatic brush. Thus the Ambrosiacee,
in contrast to other Composite, are retrogressive in structure.
On the other hand; it is evident that they have developed new
characters not attained by other Composite. This is seen
more especially in the character of the noteworthy envelopes
with which the fruit is enclosed. The fruiting heads of
Xanthium are well enough known to the wool-grower as the
detested “burrs” (Kletten), which by means of their bent
hooks attach themselves to the woolly coats of animals, and
are so dispersed. We shall see that as regards these fruiting
heads the other Ambrosiacez have also undergone retrogres-
sion in contrast with Xanthzum.

It is not surprising that such noteworthy plants have
repeatedly formed subjects for investigation. So far as the
developmental history is concerned, and apart from Payer’s?
contributions, which in the present instance need only be
indirectly considered, there are available only the investiga-
tions of Rostowzew,” and these I propose in the following to
supplement on some points.

1 Payer, ‘‘ Traité d’Organogénie comparée de la fleur” (Paris, 1857, p. 638).
2T. Rostowzew, ‘‘ Die Entwicklung der Bliiten und des Bliitenstandes bei
einigen Arten der Gruppe Ambrosiez ” (‘‘ Bibliotheca botanica,” 20).

MORPHOLOGICAL NOTES 195

Thus one topic will be the male inflorescence, while another
will be the ‘‘ fruit’? of Amzbrosza.

A. The Mate Inflorescences.

As is known, the Ambrosiacee are monoecious to this
extent, that the male inflorescences occupy the apex of shoots
at the base of which are placed the female ones,

The individual flower-heads (‘‘ capitula”) of Ambrosia trt-
partita are arranged in large numbers indefinitely (botryose)
on an elongated axis, which in turn is terminated by a flower-
head.

The lateral capitula have this noteworthy characteristic,
that the flowers are not placed as usual on the upper (adaxial)
side, but on the under side—that is, the side turned away from
the primary axis of the whole infloresence (abaxial). It is
natural, at first sight, to ascribe this occurrence to a Zorszon
of the flower-stalk. This is indeed the opinion of Rostowzew,
who says that the flower-head undergoes a torsion, in that
the peduncle grows more rapidly on its upper side than on
its lower.

This interpretation in itself does not agree with the
developmental history. If one examines the early stages,
it is seen that the position of the capitula is “reversed” at a
very early stage.

The capitula are differentiated from the embryonal tissue
as hemispherical primordia. The cells of the primordia on
the adaxial (upper) side pass first into the phase of elonga-
tion-growth, with the exception of the terminal part (a, fig. 1),
which develops as the first leaf. On the other hand, the
lower side remains embryonal. This is utilised for the
development of the growing-point of the capitulum (4, fig. 1),
except the basal part (nearest the primary axis), which later
gives rise to the intercalary meristem in the elongating
peduncle of the capitulum.

How shall we interpret this peculiar development? Two
different views are obviously possible.

It may be that the first leaf (a, fig. 1) arising from the
growing-point of the capitulum is the first leaf of the in-
volucre of the capitulum, the individual leaves of which,

196 THE SCOTTISH BOTANICAL REVIEW

united together, form an apparently entire envelope. This is
the concept of Rostowzew, which, however, has thus far to be
supplemented, that the “torsion of the capitulum ”—if one
assumes such a thing—is not a subsequent event, but it
existed from the first; in other words, it is “congenital.”
Rostowzew does not mention that if this interpretation be
assumed, then the lateral capitula of the Ambrosia would
present one of the rare cases in which the “bracts” (Deck-
blatter) of lateral shoots have aborted completely. This
occurrence is known in the flowers of the crucifere, etc., in
which one can correlate the abortion of the bracts with ex-
cessive crowding originally undergone by the flower-primordia
before the elongation of the axis of the inflorescence. In the
case of Ambrosia, this would be all the more striking because
the female inflorescences possess well-developed bracts,
although they are quite as closely crowded together as the
male ones. In no instance have I observed in the male
capitula even a trace of a subtending bract.

The second possibility is that the first leaf (indicated by a@
in fig. 1) is the bract (Deckblatt) of the capitulum. The
manner in which (a) develops from the primordium that
gives rise to (a) and (0) is similar to what occurs frequently
in flowers and inflorescences. In this instance the bract, so
to speak, is late in being formed. It does not develop in
advance of its axillary shoot, but from a primordium common
to both. The thing that surprises one is that the axillary
shoot should arise on the under side of the bract, and not, as
usual, on the upper side. But, after all, this is no more
wonderful than the “ congenital torsion” already referred to,
nor is it quite without precedent amongst other Dicotyledons.
The remarkable flowers of Evythrochiton hypophyllanthus
are situated on the under side of the leaf ; also the axillary
branches on the creeping shoots of some of the terrestrial
Utriculariz arise on the side of the leaf furthest away from
the growing-point.?

In other respects it may be assumed that the position of
the male capitula of A7zbrosta means the same thing as the
torsion (entirely brought about by their weight) of the male
inflorescences of Corylus, Alnus, Juglans, or the torsion of

1K. Goebel, Utricudarza (‘‘ Annal. du jardin botan. de Buitenzorg,” vii. 1889).

MORPHOLOGICAL NOTES 197

the anther of grasses: by this means the shaking out of the
pollen is facilitated in all these anemophilous plants.

Returning now to the morphological question, one might,
in arriving at a decision, also take the anatomical facts into
account. If (a) (fig. 1) represents the bract of the capitulum,
then one might expect that the normal orientation would be
shown in the constitution of the vascular bundle, phloem on
the lower side, xylem on the upper. Of course, J] do not
regard the anatomical conditions as decisive, since, in my
opinion, these are determined by the morphological, not the
reverse. In the present instance this would mean that, in the
event of a relatively limited development and retardation in
time on the part of the bract in comparison with the axillary
shoot, it appears quite natural that the former (the bract)
should be provided with its vascular system from the latter
(the shoot). This being so, one expects the xylem to be
uppermost, with the phloem underneath.

The conditions actually existing are briefly as follows:
The thin, much-flattened peduncle of the capitulum contains
two vascular bundles (rarely three); the phloem-groups of
these are set towards the narrow margins, and the xylems
are turned towards one another and somewhat obliquely
downwards (fig. 3). This anatomical structure is interesting
because unusual in a shoot-axis.

A certain biassed school of anatomists assumes that shoots
are distinguished from leaves by their anatomical structure,
particularly in the arrangement of the vascular bundles.
The shoot of the capitulum of Ambrosza does not differ in
structure from many leaf-stalks, except that in the latter the
xylem of the vascular bundle is directed upwards.

There is little doubt that this anatomical structure is the
result of reduction; that is, the peduncle of the primitive
type possessed more than two vascular bundles. Linked with
these bundles are those which ramify throughout the involucral
leaves and the flower, but I have not followed these in detail.
It may be mentioned, however, that the vascular bundles of
the involucral leaves are normally orientated in that their
xylem lies towards the growing-point of the capitulum. The
orientation is that described above, and for the reasons given
it does not seem to me possible to arrive at any definite con-

198 THE SCOTTISH BOTANICAL REVIEW

clusion as to the morphological interpretation of this leaf (a)
from the anatomical structure.

It might be advanced in opposition to the view that the
leaf (a) is a bract, that later on it does not always stand
opposite the peduncle; this might, however, result from
displacement. In the younger stages I saw it in the median
position occupying the whole breadth of the inflorescence-
primordium (fig. 4). As regards the further development of
the male capitula, two facts may be pointed out. Firstly,
that the external side (that on which the letter (a) is placed)
appeared to be much further advanced than the inner side ;
this is seen at once in fig. 4. Secondly, that the arrange-
ment of the flowers is not “dichasial” sympodial (as one of
the recent authors wrongly states), but corresponds essentially
with the other Composite. One sees quite clearly the broad
growing-point of the capitulum (V., fig. 4), from which the
individual flowers arise ; some of the outer ones have bracts
even now. The characteristic arrangement of the flowers is
determined by the fact that the capitulum as a whole is
dorsiventral in structure, with a more advanced development
on the outer side.

B. The Female Inflorescences.

As indicated by Rostowzew, these are dichasially arranged
(fig. 5). Each consists of a single flower enclosed by an
“envelope” (fig. 8). To understand their constitution it
will be necessary to first compare the inflorescences of other
Ambrosig. Here one finds the following series :—

1. Mixed inflorescences with female marginal florets.

2. Inflorescences with sexes distinct, in so far that in the
female only the marginal florets flower, and only the
minority of these, while in the male inflorescence the
formation of female flowers is entirely suppressed.

3. There is also a diversity in the arrangement of the
female and male inflorescences. Whereas the arrange-
ment was originally indefinite (botryose), this is re-
tained only in the case of the male inflorescences, the
female ones showing dichasial arrangement.’

1 With regard to diversity of male and female, compare Goebel, ‘‘ Ueber
sexuellen Dimorphismus bei Pflanzen ” (‘‘ Biolog. Centralblatt,” p. 657, 1910).

MORPHOLOGICAL NOTES 199

In Xanthium the male inflorescences are orientated norm-
ally ; the structure of a young male inflorescence in longi-
tudinal section is shown in fig. 6, |. The female capitulum
consists of two flowers enclosed in the many-spined envelope.
This envelope originates from the fusion of two bracts (Sa,
Sé, fig. 6, III.) in whose axils the female flowers are placed.
With reference to the development, the reader is referred to
my recent account.t. This is noteworthy in this respect, that
the two floral bracts entirely monopolise the growing-point of

the capitulum (except what is required for the flowers arising

in their axils), as is shown in the view from above (fig. 6, IV.),
and that the flowers are deeply sunk in the axis of the
capitulum.

Comparing this inflorescence with that of Amdrosza, it will
be found to have undergone still further reduction. Firstly,
we see that the spines or barbed hooks outside the envelope
are here reduced to a few small processes (figs. 7 and 8, s?) ;
secondly, each envelope contains only one flower. The
processes referred to can no longer be of use in the dispersal
of the fruits by animals.?, How dispersal is effected can only
be ascertained in the native land of the plant. The fruits are
relatively light, and float in water for a short time at least,
and they may even be rolled to a distance by wind.

It is evident that the envelope of the fruit of Ambrosza
corresponds to that of Yanthium. But is it, like the former,
a double one, or is it single? That there is some suggestion
of a double envelope is evident by the fact that the apex of
the envelope (originally laid down as a closed ring ; see fig. 7,
right-hand figure) is distinctly two-lobed in the later stages
of growth. This indicates that two leaf-primordia take
part in the formation of the envelope. Only one of these,
however, has an axillary flower-primordium. Obviously a
characteristic retrogression has taken place here: the flower
monopolises the whole area within the envelope; hence it
arises not as a lateral outgrowth on the growing-point of a
capitulum, but is terminal. There are, of course, many
examples of an organ genetically lateral in origin becoming
ultimately terminal. We may cite the spikelets of many

1 Goebel, /.c. (‘‘ Biolog. Centralblatt,” xxx., 1910, p. 722).
* They might, of course, become lodged between the claws or in the hoof.

200 THE SCOTTISH BOTANICAL REVIEW

grasses, also the carpels which in many flowers monopolise
all the residual part of the growing-point of the flower, and
the terminal stamen of WVazas, Callitriche, etc. Ambrosia is,
however, a specially well-marked example of this procedure.
The development of the female flower need not be described
further, except to point out that there is only a mere trace of
the corolla, while the stamens and calyx have disappeared
without leaving a trace.

Thus we see that Amdbrosza has carried still further in its
female inflorescence that reduction already indicated in
Xanthium. Even in Xanthium the male organs of the
capitulum are completely suppressed and the number of
female flowers is reduced totwo. As new structures there are
present the hook-like spines on the outside of the concrescent
floral-bracts, which, taking the place of the pappus originally
present, facilitate dispersal of the fruit. In Ambrosza these
spines dwindle to a few rudiments and the number of flowers
diminishes to a single one.

In the male inflorescences, however, the number of capitula
is probably increased considerably in comparison with the
type-form with hermaphrodite flowers. With this may also
be correlated, that these male capitula have departed from
the prevailing scheme of development, and as seen in their
“reversal” have progressed along new lines. Thus with
no great effort, and with a basis of facts admissible in any
new speculation, we are able to trace, in what seems to me a
satisfactory manner, the historical evolution of the structure
of the inflorescences of Ambrosza. At the same time, the
case dealt with here illustrates once again the phenomenon to
which the author has frequently made reference : namely, that
our phylogenetic series, so far as we can depict them with
any degree of probability, all represent a reduction-series.

The following may be advanced in support of this :—

1. In a descending series we have a definite starting-point
(that is, some one of the more completely equipped
forms as distinct from the more reduced ones) with
which we can compare the less completely equipped
members of the series.

2. In many instances the organs in question may still be
recognised as rudiments.

MORPHOLOGICAL NOTES 201

3. The descending series arise latest ; hence they are more
completely preserved and easier to recognise than the
ascending with a history extending much further back,
and whose members are as a rule very incompletely
preserved.

Descending series of this kind are known to every botanist,
since they appear again and again in almost every family.
Other facts also indicate that organisms become modified
mainly through retrogression and _ simplification: thus
“mutations,” for example, are essentially of this nature,
since in them there is a loss of some definite characteristic.

Has then the “nisus formatoris” of the ancient philo-
sophers itself become antiquated? Not at all; botany at
least has remained youthful. To be convinced of this, one
need only glance at what is only possible where youthful
aspirations exist, namely, the construction of genealogical
trees from below upwards. “Alas, alike in their tenure of
life, they are mostly ruins, not of the trees, however, but of
the ephemeral day-flies !”

MUNICH, Aprz/ 1912.

EXPLANATION OF THE FIGURES.

Fig. 1. Longitudinal section through an inflorescence of Ambrosia
tripartita. WV, growing point of the male inflorescence ; the individual
capitula (with the exception of the terminal one) are placed laterally on
the primary axis. a, the first leaf of a male capitulum of which @ is the
growing-point. £/, female flower with its envelope, H.

Fig. 2. Ambrosia tripartita. Longitudinal section through an older
male capitulum. V, the growing-point.

Fig. 3. Transverse section through the peduncle of a male capitulum of
Ambrosia tripartita. The xylem in each of the two large conductive
bundles is shaded.

Fig. 4. Capitulum of Ambrosia trifartita seen from above. Around
the growing-point, V, there are seen fifteen embryonal flowers in various
stages of development ; the involucre surrounds the whole.

Fig. 5. Ambrosia tripartita. Transverse section through a young
female inflorescence-group. In the axil of a bract (Deckblatt), D, is a
one-flowered female inflorescence, I, with its envelope, H; this has two

1 E. Baur, ‘‘ Einfiihrung in die experimentelle Vererbungslehre” (Berlin,
IQI1), says : ‘‘ The large majority of mutations which have been closely investigated
depend simply on the loss of some single Mendelian unit character. I have not
found, up till now, any absolutely certain case in which one or more unit characters
have arisen de novo.”

202 THE SCOTTISH BOTANICAL REVIEW

prophylls (Vorblatter), V, in the axils of which other inflorescences are
present.

Fig. 6, I-III. Xanthium spinosum.

I. Longitudinal section through a young male inflor-
escence. BB, male flower with its bract, S.

II. Longitudinal section through a female inflor-
escence. .Sa, Sd, the bracts (Deckblatter) of
two flowers, Sa,, S4,, which on the side towards
the incurved margins of the bracts are proceed-
ing to develop the floral organs.

III. Older inflorescence in which each female flower
has now the stigma developed, sv.
IV. Xanthium strumasta. A young female inflorescence seen
from above.

Fig. 7. Ambrosia tripartita. On the left a female inflorescence, seen
from the outside. s¢, primordia of the spines which remain rudimentary.
H, the envelope (clearly two-partite). 2/, flower-primordium seen through
the envelope (which is regarded as transparent). On the right a young
inflorescence seen from the outside.

Fig. 8. Ambrosia tripartita.

I. Fruit with its envelope, bisected longitudinally. H, en-
velope ; s¢, spines; P, pericarp; S, seed-coat (very
thin) ; E, embryo.

II. Fruit with envelope seen from outside.

Notes on some Mosses from the Three Lothians.
By James M‘Andrew,

Assoc. of the Botanical Society of Edinburgh.

PERHAPS the following notes may prove interesting to local
and Scottish bryologists. They consist principally of new
records of British mosses gathered in the Lothians since the
“Census Catalogue of British Mosses” was published in
1907. I am indebted to Mr. William Evans, Edinburgh,
for about one-half of them; the others were gathered by
myself and from time to time reported to Mr. R. H.
Meldrum, and will no doubt in due course appear in the
next edition of the “Census Catalogue.”

HADDINGTON, v.c. 82. Sphagnum cymbifolium Ehbth. ;
S. rigidum Schp.; Andreea petrophila Ehrh.; And. Rothiz
W. and M. (Traprain Law, East Linton); Polytrichum

Scottish Botanical Review.)

Fig.8

K. von GOEBEL, ‘‘ Morphological Notes.”

NOTES ON SOME MOSSES FROM THE THREE LOTHIANS 203

urnigerum L. ; P. alpinum L. (Traprain Law) ; P. formosum
Hedw. ; Pleuridium subulatum WRabenh.; Rhabdowetsia
Jugax B. and S. (Traprain Law); Decranella squarrosa
Schp. ; Blindia acuta B. and S. (Traprain Law) ; Grimmia
apocarpa Hedw., var. rivularzs WW. and M. (East Linton) ;
G. patens B. and S. (Traprain Law); G. decipiens Lindb.
(Traprain Law—cum fructu); G. montana B, and S. (Trap-
rain Law); G. Stzrtonz Schp. (Garleton Hills, by W. Evans) ;
Rhacomitrium protensum Braun (Traprain Law, by W.
Evans); R. canescens Brid., var. evicoides B. and S.;
Hedwigia imberbts Spruce (Traprain Law) ; Pottza bryotdes
Mitt. (Gosford Bay, by Mr. John Hunter); Barbula fallax
Hedw., var. brevifolia Schultz (west of Dunbar); Wezsza
microstoma C. M. (Gosford Bay); Cinclidotus fontinaloides
P. Beauv. (river Tyne); Zygodon Mougeoti B. and S.
(Traprain Law); Orthotrichum stramineum Hornsch. ;
Lartramia pomiformts Hedw., var. crispa B. and S. (Trap-
rain Law); Leptobryum pyriforme Wils. (W. Evans) ;
Plagiobryum Zerit Lindb. (Traprain Law); Webera pro-
ligera Bryhn (W. Evans); Bryum alpinum Huds., and var.
viride Husn. (Traprain Law); 8. argenteum L., var.
lanatum B. and S. (Gullane Links) ; Wnzum stellare Reich.
(Gullane Links); Pterygophyllum lucens Brid., and Hetero-
cladium heteropterum B. and S. (Traprain Law); Brachy-
thectum salebrosum B. and S. (Gullane Bay) ; B. plumosum
B. and S. (East Linton) ; Plagiothectum pulchellum B. and S.
(Traprain Law); Amblystegium serpens B. and S., var.
salinum Carr. (Gullane Links); Hypnum elodes Spruce ;
H. aduncum Hedw., var. paternum Sanio (all Gullane Links) ;
H. Sendtnert Schp. (Gullane Links). Regarding this moss
Mrs J. A: Wheldon; F.L.S., Liverpeol,, writes me: ~ Your
plant is typical H. Sexdtneri Schp. It is new to Haddington-
shire, and as a matter of fact it is the first specimen I have
seen from Scotland, although recorded from v.c.’s 86 and 87.
I have for years been trying to get a specimen of the inland
plant, but without success so far.” It grows in a shallow
lagoon on Gullane Links, and covers the whole bottom with
a mass of several acres exclusively of this moss. A. fluztans
L.; Hf. cupressiforme L., var. filiforme Brid. ; A. Patientie
Lindb.

204 THE SCOTTISH BOTANICAL REVIEW

Traprain Law, East Linton, has several very interesting
mosses such as AHedwigia imberbis Spruce; Grimmia
montana B. and S.; Grimmza decipiens Lindb., all three in
great abundance, the latter in fruit; Axdreza Rothit
W. and M.; Bryum alpinum Huds. Gullane Links also,
like Tents Muir, Sands of Barrie, Dunbarnie Links, etc.,
have also several rare mosses already recorded as Dztrichum
frexicaule Hpe., var. densum B. and S.; Swartzta montana
Lindb., and S. zzclinata Ehrh.; Barbula Hornschuchiana
Schultz; Tvrichostomum flavo-virens Bruch; Excalypta
rhabdocarpa Schweg.; Amblyodon dealbatus P. Beauv. ;
Meesia trichoides Spruce; Catascopium nigritum Brid. ;
Bryum calophyllum R. Br.; Amblystegium filicnum De
Not., var. Whiteheadizt Wheldon ; Hypnum chrysophyllum Brid. ;
H. giganteum Schp., Thuidium recognitum Lindb., etc.

EDINBURGH, v.c. 83. This vice-county has been better
searched for mosses than the two adjoining counties, and
hence the new records for it are fewer. Sphagnum cuspt-
datum FEhrh. var. submersum Schp.; S. molle Sull. ;
Polytrichum strictum Banks (all from Bavelaw Moss) ;
Oligotrichum hercynicum Lam., recently got in Corstorphine
Hill wood, is now extinct; Fuss¢dens crassipes Wils.
(Colinton Dell); Barbula Hornschuchtana Schultz (waste
ground north of Donaldson’s Hospital); PAyscomztrella
patens B. and S. (Torduff reservoir, by Mr. W. E. Evans.
I got this moss in plenty on the dried-up bottom of a
lagoon west of S. Queensferry last year) ; Bryum argenteum
L., var. lanatum B. and S.; Brachythecium glareosum
B. and S. (Flotterstone Bridge); Hypuum polygamum
Schp.; A. aduncum Hedw., var. polycarpon Bland.; H.
fiuitans L., var. Jeanbernati Ren., and var. atlanticum Ren. ;
H.. exannulatum Giimb., var. pinnatum Boul., £. stenophyllordes
Ren., and var. pimnatum Boul., f£. gracilescens Ren.; HF.
entermedium Lindb., var. tenellum Roth (=H. pellucidum
Wils.) (all in Bavelaw Moss); AH. cupressiforme L., var.
tectorum Brid.; H. imponens Hedw. (Bavelaw Moss). I
got this also last year near West Linton in Peeblesshire,
v.c. 78.

LINLITHGOW, v.c. 84. Sphagnum rigidum Schp., and
var. compactum Schp.; S. cymbifolium Ehrh.; S. tenellum

NOTES ON SOME MOSSES FROM THE THREE LOTHIANS 205

Ehrh.; S. fimbriatum Wils.; S. Girgensohnit Russ. (all
in Drumshoreland Moss); Pleurtdium subulatum Rabenh. ;
P. alternifolium WRabenh. (N.B. railway embankment) ;
Cynodontium Bruntont B. and S. (Cocklerue); /7ssidens
recurvus Starke; Barbula lurida Lindb.; Barbula Horn-
schuchiana Schultz ; Physcomttrella patens B. and S. (all
west of S. Queensferry); Sartramia ithyphylla Brid.,
Leptobryum pyriforme Wils. (Kirkliston distillery) ; Webera
annotina Schweg. (Drumshoreland) ; Bryum alpinum Huds.
(Cocklerue); Thuzdiwm recognitum Lindb. (west of S.
Queensferry); Hypnum riparium L. (Linlithgow Loch) ;
HT. stellatum Schreb., var. protensum Rohl; A. exannu-
latum Gimb.; H. cordifolium Hedw. (Drumshoreland
curling pond); H. Joreum B. and S. (W. Evans).

In the “Census Catalogue of British Mosses” for 1907
there are several mosses which require rediscovery, for their
records are old. For instance, the rare Grimmias on
Arthur’s Seat have evidently disappeared. Grimmia
leucophea Grev., is the last one of the group I have seen
on Arthur’s Seat, and I suspect that now it too is extinct.
The following among others require refinding :—Sphagnum
Austint Sull. (v.c. 83); Bryum Warneum Bland. (v.c. 82) ;
Cryphea heteromalla Mohr (v.c. 82); Hypnum eugyrium
Schp. (v.c. 83); Grimmia orbicularis Bruch (v.c. 83);
G. anodon B. and S. (v.c. 83); G. conferta Funck (v.c. 83) ;
G. commutata Hiibn. (v.c. 83); G. ovata Schweg. (v.c.’s 82
and 83); Axtetrichia curtipendula Brid. (v.c. 83), and several
others. Tvichostomum mutabtle Bruch.; TJ. inuclinatum
Dixon; 7. xztidum Schp., have yet to be gathered in
Vic, 82.

Saxifraga Hirculus, L., in Caithness, and its Dis-
tribution in the British Isles. By Arthur
Bennett, A: io:

Mr. G LILLIE has kindly sent me good flowering specimens
of the above species which he found growing ‘‘in a floating
bog about ten yards across each way. It was very shaky,

206 THE SCOTTISH BOTANICAL REVIEW

but there was not much danger of one sinking in it as there
was considerable growth of grasses, plants, and mosses.”
It occurred between Lybster and Loch Rhuard, altitude
430 feet, 58° 22’ N. lat. about 1} miles from the loch.
The loch itself afforded Lobelia Dortmanna and Subularia
aquatica, rare plants in the county. As the water from the
loch and the stream running into it is discharged into the
Loop river, thence to the Little river, and finally into the
Thurso river, the plant belongs to the north-west watershed,
not to the east.

S. Hirculus is not on record north of West Perth and
Kincardine, 57° N. lat. ; the extension of range north
to Caithness is interesting, though it is a high arctic and
northern species.

As a rare species it may be well to summarise its distribu-
tion in our Isles.

g1. Kincardine. Wet moor on the farm of Jacksburns,
Glenbervie. 21.6.1839.—-Mr. James Rae. This is the station
erroneously given in “‘Topl. Botany” as “ Aberdeen, J. Rae.”

87. Perth, W. Madie Moss, above Tillicoultry, c. 2000 feet.
Northern side of the Ochils, Clackmannanshire—Mr. W.
Thompson.

83. Edinburgh. Ona wet moor between Fala and Stow.
—G. J. Blackie. Three miles beyond Blackshiels, Pentland
Hills, Midlothian. 11.8.98.—F. C. Crawford sp.

81. Berwick. Moor south of Langton Lees farmhouse,
plentiful.—Dr. Johnston, c. 1831 ; sp. Cardiff Museum !.

78. Peebles. Pentland Hills, source of the Medwin,
about 1000 feet. 11.9.1836.—Dr. A. Hunter. J. H. Balfour
sp., 19.9.1838.

77- Lanark. Boorland Moss, Walston, Sept. 1850.—G. F.
Blackie. No altitude given, but the parish ranges from 660
to 1000 feet alt.

69. Westmoreland. Neathheath Syke, alt. 1800 feet, Sept.
1840.—John Bell. Backhouse in Herbarium, York !.

66. Durham. Wet moor, Inishope, Weardale, 2.8.1885.—
J. P. Soutter sp.

65. York, N.W. Sedbergh.—J. Handley, ‘‘ Yorkshire
Naturalist” (1903), p. 408. Cotherstone Fell, 1847.—D.
Oliver sp. The actual station is swamps at the junction of

SAXIFRAGA HIRCULUS, L., IN CAITHNESS 207

the river Balder with the Black Beck. Discovered here by
J. Backhouse, 1810; sp. in York Herbarium.

58. Cheshire. Knutsford Moor, where it was associated
with Andromeda, Melampyrum pratense, Carex limosa, curta,
and stellulata ;1 also with Szum angustifolium, Cicuta, Pota-
mogeton polygonifolius, Carex ampullacea, Lastrea spinulosa,
L. thelypteris, and Osmunda regalts.

The date of its first record here (and for Britain) is 1724,
Dibiekayy, Soyn,” 3rd: ed. ip. 355;' but the’ plant had
been gathered some time between then and 1696, as in an
old volume of coloured impressions of Cheshire plants? the
locality is given, and the names are those of the 2nd ed.
of the ‘‘Synopsis.” Specimen sent from here by Mr. Okell
of Chester, and figured in ‘“‘Eng. Botany,” t. 1009,° Aug.
1801. Extinct in 1842. The species did not become extinct
here altogether through natural causes, as the following
extract will show :—

‘« Still exists on Knutsford Moor, but is almost destroyed
by the rapacity of some individuals who have dug it up for
sale in the most remorseless manner.”—Dr. J. B. Wood, in
““Ehytolosist,”” 1. (1842), p. 282.

In Ireland in the middle and north-east (ze. in six of Mr.
Praegers’ divisions), but characterised as very rare.

In Europe it is a species of Arctic Norway to 69° 20’
N. lat., Swedish, Finnish, and Russian Lapland, Iceland,
along the Siberian coast to Behring’s Straits. The American
Arctic coast to Labrador, south to Saskatchewan. E. and
W. Greenland, as var. alpina, Engler Mon. Warming, ‘‘Om
Groenlands Vegetation,” 1886-7, does not give any height
for this, though citing many species up to 4000 feet. It will
bear intense cold, as on the island of Kolegew, where ‘‘the
thermometer never rose above 9° R. in July and August
1841” ;° there it is common. It also occurs on Spitzbergen
and Bear Island.

I am not sure whether the first Scottish record was the

1 Lord de Tabley, ‘‘ Fl. Cheshire” (1899), p. 142.

2 Perry, ‘* Phyt.,” i. (1843), p. 700.

3 This plate is not quoted in the 3rd ed., iv. (1865), t. 550.

4 Kjellman, ‘‘ Vega Expedition ” (1882), 263, 302.

® Erman, “‘ Arch, Wiss. Kimde v. Russlandes” (1852), x. 313-16; quoted by
Fielden and Geldart, ‘‘ Trans. Nor. and Norf. N. Soc.” (1896), p. 169.

208 THE SCOTTISH BOTANICAL REVIEW

Berwick one in Watson’s ‘‘ Outlines,” 1832, or the Blackshiels
one in the ‘‘Edin. Phil. Journal,” as I have not access to
the latter. It is not given in Dr. Johnston’s ‘‘Flora of
Berwick,” ii., 1831.

Along with the Saxifrage there was a form of Cerastium
vulgatum, L. (trivzale, Link.), which looked very different
from the ordinary form of the species, no doubt induced by
the place of growth. Still thinking it might be one of the
Swiss forms so found, I sent specimens to Dr. Schinz of
Geneva; he referred them to Dr. Keller of Berne. Dr.
Keller reports: ‘‘They are only a form of C. c@spitosum,
Gilib.1 (=valgatum, Wahb.), but the plant ought to be
observed. Like all species of Cerastium, cespitosum shows a
great variation in all parts according to the medium in which
it lives.’ Dr. Keller writes: ‘‘C. vulgatum, L. (=glomera-
tum, Thuill, v¢scosum, Fr.),’’ which opens the door to a war
of words. One accepts the decision ; at the same time, the
plant is easily distinguished by the eye. Another addition
to the county was growing with it— Ranunculus scotzcus,
Marshall.

A New Japanese Grateloupia. By E. M.
Holmes; F.L.S5)*Rs525889

(With One Plate.)

AMONGST some marine algz collected in Japan by Mr. S.
Okubo, and brought to me for identification, I noticed one
which I was unable to match either at the Kew Herbarium or
at South Kensington. Although bearing some resemblance
to Grateloupia filicina, it differs so much in habit and colour
that I have decided to describe it as a new species.

Grateloupia subpectinata, n. sp—Fronde compressa, plana,
ad 15 cm. longa, et 3 mm. lata, pinnatim ramosa, ramis a
basi angustiore, longe subuliformibus, inferioribus longioribus,
infra medium latere inferiori ramellis brevibus subpectinatis,
latere superiori dentibus paucis preditis, ramis supremis
simplicibus, brevibus. Color pulchre roseus.

1 Gilibert, ‘‘ Fl. Lithuanica,” v. (1782), p. 159.
2 Read before the Botanical Society of Edinburgh.

Scottish Botanical Review.

E. M. HOLMES.

A NEW JAPANESE GRATELOUPIA 209

Hab.: Japan, S. Okubo, 1912.

The nearest approach to this species, in the mode of
branching, is Grateloupra Pennatula, Kuetzing, a native of
Cuba (“ Tab. Phyc.,” vol. xvii., tab. 27, a, 4,), which has similar
ramelli, but both the ramuli and ramelli are lanceolate-linear
rather than subuliform, and are much shorter in proportion.
The rose colour, so far as I know, is never found in forms of
Grateloupia filicina, although I have seen many forms referred
to this species. The structure, is however, typical of Graée-
loupia, and not of loose texture as in the section Glocogenza,
to which a rose-coloured species, G. acuminata, from Japan,
previously described by me, belongs.

Kenfig Burrows: An Ecological Study.
Communicated by M. Y. Orr.’

KeEnFIG Burrows forms the southern extremity of a fringe of blown
sand which borders the south-west coast of Glamorgan as far as
Swansea, a distance of fifteen miles. The breaks in its continuity
are brought about by the rivers Avon and Neath. At Kenfig the
sand dunes extend inland for over two miles at the broadest part,
and occupy an area of approximately 1500 acres. ‘The general drift
of the sand is in an easterly direction.

Apart altogether from the biological problems involved, the area
is of great historical interest, for a waste of sand now covers what
was at one time a prosperous town. A few scattered ruins on the
northern dunes are all that remain to mark the site of the castle
and buried city of Kenfig. The invasion of the sand since Roman
times appears to have been gradual, but, according to tradition,
sand storms of considerable magnitude occurred in the fourteenth
century. In 1538 ruin had overtaken the town and castle, and both
were abandoned to the advancing sand.

Kenfig is now represented by a little hamlet, situated about half
a mile from the castle ruins, on a ridge overlooking the sheet of
water known as Kenfig Pool. This water occupies the central
portion of the landward margin of the dunes. From its eastern
shore the ground slopes gradually upwards to the ridge, the height
of which, and of the adjacent fixed dunes, varies from ninety to one
hundred feet above sea-level. The pool forms the apex of a trian-
gular wedge of fertile land which has not been invaded by the sand
to any great extent. No doubt, the pool, owing to its position,
forms a natural barrier to the further incursions of the sand. The

' Read before the Botanical Society of Edinburgh.
VOL. I. 16

210 THE SCOTTISH BOTANICAL REVIEW

area of this sheet of water is 68 acres, and its greatest depth is
11 feet. The water is fresh and fed by springs on the landward
side. It has existed in its present form since 1876, but is of less
extent and slightly different outline to the pool of 1814. It was
originally a marsh, and its eastern margin is still, in part, character-
ised by a marsh vegetation. At that time it was drained bya stream
which flowed in a northerly direction and emptied itself into Kenfig
river. To-day there is no visible outlet, but it is possible that its
waters filter through the sand and ultimately enter the river in
that way.

Geology, Climate, and Rainfall.

The blown sand rests on the Keuper marls of the Trias formation,
Rocks which project here and there through the sand nearly all con-
sist of conglomerate, which is more or less calcareous in composition.
The wedge of land which has the pool as its apex is boulder clay
resting on Triassic conglomerate, while the southern boundary of the
‘‘white” dunes is of like glacial origin. A broad stretch of alluvium
marks the northern limit, and an irregular alluvial tract extends from
the margin of the pool to within a short distance of the shore. The
climate is mild and humid. Owing to the prevalence of warm
south-westerly winds the mean January temperature is about 41° F.,
and it is probable that the resultant mild winters have a modifying
effect upon the vegetation. The corresponding July temperature is
62° F. The average annual rainfall, estimated over a number of
years, is a little over 33 inches. ‘T'he wettest months of the year
are July to January inclusive. A complete analysis of the soil and
sand taken from different stations on the dunes is now being carried
out, and a detailed account of soil conditions will be embodied in a
future publication. Cattle and horses are pastured on the dunes,
and the soil is thus enriched with manure. Molluscan shells are
abundant in many of the hollows.

The plant formation which covers this area is a natural one, and
presents many interesting biological features. A comprehensive
ecological study of its vegetation, on modern lines, is now being
undertaken, and it is hoped to extend this investigation so as to
include the entire belt of sand dunes from Porthcawl to Swansea.
So far, detailed observations have been chiefly confined to the
Kenfig district. This communication is, therefore, not intended to
be exhaustive, and is merely a brief account of the more salient
features of the vegetation.

THE PLANT ASSOCIATIONS,

The three most prominent and well-marked associations in this
formation are the following: 1. The association of Ammophila
arenaria; 2. the association of Salix repens; 3. the association
of Pteris aquilina.

1. The association of Ammophila arenaria,—This association is

KENFIG BURROWS: AN ECOLOGICAL STUDY Zit

characteristic of the “shifting” dunes, with the Marram grass as
the dominant species. ‘This species occupies the first seaward line
of sand hills, the Agvopyretum juncet (sea couch-grass association)
of the Somerset and Lancashire dunes being practically absent in
this area. It is worthy of notice that Llymus arenarius, which in
Norfolk gives rise to low dunes, is entirely absent from this coast.
The higher inland dunes, many of them over fifty feet in height,
bear on their crests the characteristic tufts of Marram grass. It
descends on the leeward slopes and colonises the sandy hollows,
and in many of the latter,its rhizomic habit of growth is particularly
obvious. As a “‘sand-binder” it is pre-eminent, and it is interesting
to note that in the Charters of Kenfig of 1330 special provision was
made for its protection and preservation. Ammophila also occurs
in abundance on some of the ‘‘ fixed” dunes.

2. The Salix repens association.—Salix repens occurs on the
shifting” dunes as a “‘sand-binder,” producing a well-marked
association. In the sandy dune valleys it forms a carpet of low-
growing scrub, but in the more exposed stations it collects around it
the blown sand and gives rise to “ hummocks” and small dunes.
In the damp hollows it forms a fringe round the marshy ground
and the dependent species vary accordingly. Saézx represents the
second stage in succession on the dunes, as Ammop/ila represents
the first. In the more open stations of this association Marram
grass is the subdominant species. Salix repens occurs on the
Lancashire dunes, but Moss describes it as a comparatively rare
plant on the Somerset sand hills.

3. The Preris aguilina association.—The bracken covers acres
of the ‘‘fixed” dunes in this area, and its outliers blend with the
willow of the preceding association. Its rhizomic habit of growth
makes it likewise a successful sand-binder, and it forms the last stage
in succession. According to Massart, Per7s is entirely absent from,
or very rare, on the Belgian dunes. It has here undoubtedly
adapted itself to life on the dunes and grows luxuriantly. The
fronds are regularly harvested for cattle-bedding. Although it is
such an abundant type on the “grey” dunes near Kenfig pool, it
is sparsely represented on the adjoining boulder clay. This may be
due, in part, to the fact that the latter is under cultivation, or perhaps
its absence may be accounted for by the more or less calcareous
nature of the clay; the bracken being regarded by some authorities
as a calcifuge.

In these three chief associations the subordinate species vary
according to the character of the association. The Marram grass
association is essentially an open one, and on the seaward side of
the first range of ‘‘mobile” dunes it is an almost pure association.
As a result of this open character, competition among the associated
species is reduced to a minimum. The Sadzx repens association is
of a less open nature. The habit of growth of the Sadx affords
protection from blowing sand, and excessive insolation, to the types
which it shelters. At the same time, a certain amount of humus is

212 THE SCOTTISH BOTANICAL REVIEW

present, especially in damper stations. On the embryonic dunes
formed by this species few associated plants occur, principally on
account of the more exposed situation. These hummocks are sub-
jected to frequent denudation by the wind, and their bases commonly
present a weathered appearance. In the fixed dune association
where Preris is the dominant plant, the amount of shade cast by its
fronds is relatively greater, and the dependent species are mostly
grasses, Agrostis sp., etc. These three dominant species are rhizomic
in habit, and are specially adapted to hold their own under apparently
unsuitable edaphic conditions.

Extent and Distribution of these Associations.

The Marram grass association extends inland as far as Kenfig
Pool, where its outlying stations dovetail with those of the Sadx
association. This latter covers a broad area on the alluvium on
the seaward side of the pool, and extends laterally on its flanks. It
gradually loses its identity in the bracken association of the fixed
dunes. Distributed throughout these chief associations are minor
associations which are of local occurrence. Audus cesius, the dew-
berry, for instance, forms a community of this description. This
species occurs in certain situations on the dunes, and represents,
with Prer’s and Sadix, the last stages in succession. Sometimes
associated with it is a hybrid form of Rwhus which gives rise to
hummocks. In addition to these principal associations many sub-
associations and plant societies exist. These will be described at
a later stage.

The term “plant association” implies a plant community, and it
therefore follows that the dominant types, mentioned above, have
associated with them species which may be regarded as subdominant,
abundant, or occasional, according to the frequency of their occurrence.
Some species are apparently always associated with a particular
dominant, but on the dunes there are many cosmopolitan types
which are not limited in their distribution to any one association.
A list of subordinate species occurring in any one plant community
does not therefore imply that they are characteristic of that associa-
tion alone, or that they do not occur as frequently in any other
association. The majority of associated species colonise the sheltered
“flats” and dune valleys which wind in all directions among the
high dunes. The configuration of these valleys is varied, and some
are characterised by a flora quite distinct in composition from that of
a neighbouring depression. Some are marsh-like in character, while
others are undulating plains of blown sand, with embryonic dunes in
various stages of formation. The species which occupy the dry
hollows are psammophilous, although some, like Cvzcus arvensis,
which frequently occurs, are as characteristic of other formations.
On the fixed dunes sward-forming plants and others are associated
with Pris and Ammophila, while many species of moss form patches

KENFIG BURROWS: AN ECOLOGICAL STUDY 213

of bright green colour everywhere, and are particularly obvious
during the early spring.

The following detailed description of the associations is not
intended to be exhaustive. Many problems arising out of the
distribution of the dependent species in this formation have yet to
be solved, and the listing of associated tvpes is but a step on the
way to their complete elucidation. No sharp line can be drawn
between two associations, and it is often difficult to determine
whether a certain species belongs to one association or to another.

In the Kenfig sand-dune area the Sa/ix repens association is of a
transitional type. In its more open stations on the mobile dunes
the majority of its subordinate species are those of the Marram
grass association. ‘Towards its inner limit, fixation of the sand is
complete, and the included species are those of the fixed dunes

Before enumerating the species which are found in the chief
associations, it is necessary to call attention to the plants which
occupy the foreshore, above high-water mark. These plants are
usually included in the sand-dune formation, although, as a rule,
they are more of a halophilous nature. ‘This association of strand
plants is not well developed on this part of the coast; only a few
species occurring here and there. The following are the commoner
species : Cakile maritima, Scop.; Arenaria peploides, Linn. ; Sadsola
Kali, Linn.

Marram Grass Association.

In the “flats” and sheltered hollows among the “shifting” dunes
the following species occur :—

DOMINANT SPECIES.

Ammophila arenaria, Link.

ABUNDANT SPECIES.

Lrodium cacutarium, L’Herit. Luphorbia Parattias, Linn.

Anaphalis margaritacea, B. & Luphorbia portlandica, Linn.
Hi: fil. Tris fetidissima, \.inn.

Senecio Jacobea, Linn. Carex arenaria, Linn.

Cnicus arvensis, Hoffm.

FREQUENT OR OCCASIONAL SPECIES.

Erigeron acre, Linn. Cynoglossum officinale, Linn.
Taraxacum erythrospermum, Verbascum Thapsus, Linn.
Andrz. Verbena officinalis, Linn.

The variety g/andulosum, Bosch., of Erodium cicutarium has been
recorded from this district. Axaphalis margaritacea is well established
on the dunes, and apparently succeeds best where competition is
reduced toa minimum. Cvicus arvensis colonises the sandy ho!lows

214 THE SCOTTISH BOTANICAL REVIEW

towards the inner limit of the mobile dunes. It is also abundant
on the fixed dunes, as in Norfolk. Luphorbia portlandica is absent
from the Somerset sand hills.

Salix repens Assoctation.

As already mentioned, this association includes within its limits
plants of the mobile dunes, as well as those of the fixed dunes. The
following species occur :—

DOMINANT SPECIES.

Salix repens, Linn.

SUBDOMINANT SPECIES.

Rubus cestus, Linn. Ammophila arenaria, Linn.

FREQUENT SPECIES.

Viola ericetorum, Schrad. Leontodon (nudicaule), B. & S.
Viola Curtisit, Forster. Euphorbia Paratias, Linn.
Erodium circutarium, LHeérit. Tris fetidissima, Linn.

Rosa spinosissima, Linn. Carex arenaria, Linn.
Erigeron acre, Linn. Phleum arenarium, Linn.

Cuicus arvensis, Hoffm.

Viola Curtisit was recorded from the sand dunes near Aberavon,
over sixty years ago. It is not found in Somerset. The damp
sandy hollows which occur within the limits of the Sa//x association
possess a characteristic vegetation. These depressions lie at a low
level, and during the winter months are frequently submerged. A
layer of humus is formed in places, and in conjunction with an
increased water content produce a plant society of a definite type.
The Sa/ix forms a fringe round the margin of these hollows, which
are colonised by the following species :—

Samolus Valerandt, Linn. Littorella untflora, Aschers.
Erigeron canadense, Linn. Flelleborine longifolia, R. & B.
Flydrocotyle vulgaris, Linn. Orchis incarnata, Linn.
Anagallis tenella, Murr. Juncus effusus, Linn.

Blackstonia perfoliata, Huds. Eequisetum palustre, Linn.
Mentha aquatica, var. hirsuta

(Huds.).

The occurrence of these species in the different hollows depends
upon the relative amount of moisture present. Practically pure
societies of one particular species are not uncommon.

Kenfig Pool is situated within the limits of the Sa/zx association,
and its general features have already been described. On its sandy
margin the vegetation met with is mainly that of the damp hollows.
A sheltered bay on its landward side is rich in species, and the

KENFIG BURROWS: AN ECOLOGICAL STUDY 215

vegetation here and there is of the marsh type. On the opposite
shore, which is exposed to frequent sand-blasts from the adjoining
dunes, very few plants occur. Space does not permit of a detailed
description of the flora and its distribution, but the following list
of commoner species will suffice to indicate its character and
composition :—

Ranunculus peltatus, Schrank. Polygonum amphibium, Linn.

Castalia alba, Wood. Rumex Hydrolapathum, Huds.
Hydrocotyle vulgaris, Linn. Tris fetidissima, Linn.
Menyanthes trifoliata, Linn. Alisma ranunculotdes, Linn.
Scutellaria galericulata, Linn. Lleocharis palustris, R. & S.
Scutellaria minor, Huds. Carex leporina, Linn.
Littorella uniflora, Aschers. Carex Goodenowitt, Gay.

Limosella aquatica, var. tenutfolia, Lej., was recorded as occurring
in great quantities on the sandy shores of the pool in 1898, but it
has not been observed in this locality since 1908. Trees are
practically absent from the mobile dunes. On the northern limit of
the fixed dunes sand plantations occur, while the alder grows freely
on the banks of Kenfig river, and follows its course for a considerable
distance on the alluvium. Stunted, wind-pruned specimens of
Sambucus nigra are met with here and there on the “shifting”
dunes, usually in somewhat exposed situations. /Zippophaé rhamnoides,
which forms a dense scrub on the Norfolk dunes, and is also
reported by Moss as frequent on the Somerset sand hills, does not
occur.

Vegetation of the Fixed Dunes.

The transition from the associations of the ‘‘ mobile” dunes to
those of the “fixed” dunes is gradual. Fixation by Salix, Rubus,
and sward-forming plants, like Festuca rubra, begins in the more
sheltered hollows, near the inner limit of the shifting dunes. In
the more exposed stations the Marram grass still holds its own, and
on the dune grassland, where /er7s is absent, it remains a dominant
type. Where dune grassland gives place to cultivated land, bush
vegetation and trees form a final barrier to the invasion of the sand.

Pteris aquilina Assoctation.

In the more open stations of this association the following species
occur, in addition to many of those already mentioned :—

Erophila verna, E. Meyer. Saxifraga tridactylites, Linn.
Cerastium semidecandrum, Carduus nutans, Linn.
Linn. Sedum acre, Linn.
Cerastium tetrandrum, Curt. Lycopsis arvensis, Linn.
Stellaria apetala, Ucria. Myosotts collina, Hoffm.
Sagina nodosa, Frenzl. Koeleria gracilis, Pers.
Vicia angustifolia, Linn. Festuca rubra, Linn. and _ its

different maritime forms.

216 THE SCOTTISH BOTANICAL REVIEW

On those fixed dunes, from which Pers is absent, Ammophila is
almost a dominant type, and the association is a closed one. Many
of the included plants are those of cultivated land. In addition to
some of the species already mentioned as occurring in the bracken
association, the following are abundantly represented :—

Geranium molle, Linn. Leontodon nudicaule, B. & S.
Galium saxatile, Linn. Rumex Acetosella, Linn.
Bellts perennis, Linn. Urtica dioica, Linn.

Cunicus lanceolatus, Willd. Luzula campestris, D. C.

Cnicus arvensis, Hoffm.

On the innermost margin of these fixed dunes U/ex europaeus is
occasionally met with. Mosses and lichens are common here, and
in the transitional associations. They play an important part in
preparing the ground for the growth of flowering plants. Here and
there on these fixed dunes limited areas have been enclosed and
are now under cultivation. It is impossible at this preliminary
stage in the study of the vegetation of this formation to form any
opinions on its composition and distribution which will be of
lasting value. The particular area which forms the subject of this
communication is too restricted in extent, in any case, for generalised
deductions. The mere listing of species included in the different
associations is but the initial stage in any ecological study. Were
these lists complete, much would yet remain to be done. Critical
forms must be made the subjects of careful investigation. <A
thorough investigation of the dependent species and their habit of
life must be undertaken, to determine the relationship existing
between them and the dominant type, in the association in which
they occur.

The powers of adaptation and adjustment to environment of the
individual plants, and many other kindred problems, still await
solution.

The Past History of Monocotyledons, with Remarks
on their Origin. By A. R. Horwood.
(With Two Plates. )
(Continued from p. 180.)

4. THEORIES AS TO THE ORIGIN OF MONOCOTYLEDONS.

There have been many theories as to the origin of monocotyledons.
They may be regarded as (i.) systematic, (il.) morphological and
embryological in character.

According to the former evidence monocotyledons are generally

THE PAST HISTORY OF MONOCOTYLEDONS 217

regarded as the more primitive and dicotyledons as derived from
them.

As an exponent of this view we may select the classification of
Engler with some modifications, and follow the generalisations of
Dr. Rendle,! whose conclusions must largely be based upon the vast
and unequalled collections of dried specimens in the National
Herbarium, supplemented by a knowledge of the living forms repre-
sented in the Royal Botanic Gardens at Kew.

Dr. A. B Rendle? adopts Engler’s view that Pandanales are the
most primitive. The flower of Zyfha recalls the gymnospermic
strobiloid flowers. ‘They are held to represent developments from a
common ancestor along different lines.

Helobiez are next in order, and /Vazas is the most primitive, with
micro- and mega-sporangia, axial in origin. Glumiflorz have flowers
typically naked, and ‘‘correlated with this is the suggestion that the
group represents a primitive and not a reduced type of monocotyle-
dons, which have developed on their own lines.” It is said to
‘suggest a derivation from an earlier, more strictly aquatic group,”
and ‘‘that it was the first group to become established upon the
drier land surface.”

In regard to Spadicifloree, similarity of seed-structure serves to
unite palms and aroids here. Dr. Rendle thinks they may represent
a further development of Pandanales.

Lemnacee are a reduced type of Aroidez, whose huge herb type
is an adaptation to a hot, moist climate, and an outcome of this is
the epiphytic form, whose origin may be traced in Philodendron.
Assuming Glumiflorz to be primitive, the higher types follow on from
‘them.

Farinose (including Restiacez, Eriocaulaceee, Commcelinacee,
Pontederiacez, Bromeliaceze) are limited in range and _ highly
specialised. The Restiaceze suggest a glumiflorous type, adapted
to dry conditions which Glumiflorg in many points appear to confirm.
The Eriocaulacez and other families are also highly specialised.

The Lihiflore include Juncaceze, which appear to be derived from
Glumiflore. Liliaceze are closely related to them. Iridacez, how-
ever, appear to exhibit reduction. In these and in the Liliacez and
Amaryllidaceze there is a marked development of corms and bulbs,
and the plants are adapted to land areas not occupied by a meso-
phyte vegetation.

In aloes and yuccas we find an increase in stem by secondary
growth to meet xerophytic conditions on a larger scale; the leaves
also are larger and more numerous. In Sax both leaf and flower
structure are adapted to a climbing habit.

The Scitaminez are derived from some of the Liliifloree. AZmsa is
the most primitive, and all are adapted in their herbaceous characters
to a moist climate.

1 “** The Classification of Flowering Plants,” 1904, p. 378 e¢ segg.
2 Tbid.

218 THE SCOTTISH BOTANICAL REVIEW

Amongst Microsperme the Burmanniacez exhibit an affinity with
Liliifloree.

Thus far we have reviewed the results attained by the study of the
systematist. Let us see what morphology and embryology, with some
ecological facts, have to tell us.

The opinion expressed by Schimper and Schenk in their text-book
of Paleobotany! largely coincides with the decision arrived at by
some of those who have based their researches upon the facts of
morphology or embryology. They consider that dicotyledons are
the older race. They remark upon the late appearance of mono-
cotyledons and their rare occurrence as fossils. They consider the
stem-structure proves that monocotyledons are the higher group.
It is pointed out that the dicotylous type of stem-structure is found
in fossil vascular Cryptogams and Gymnosperms. The simple floral
structure is thought to point to a group in process of evolution.

Dr. W. C. Worsdell? holds the view that dicotyledons are derived
from monocotyledons, that the cotyledon is always terminal, and
that the stem-structure is not due to reduction but is primitive. He
regards the fruiting stem and petiole as alone able to unfold the true
ontogeny of the individual whereby the stages in the race-history
may be made out. Dr. Worsdell’s latest view is that monocotyledons
and dicotyledons are variants of one form and that there is only ove
cotyledon. He has found a similar stem-structure of scattered
bundles in about thirty orders of dicotyledons, and considers the
dicotylar vascular system derived from the monocotylous. Many
other characters also are correlated with it.

In Jungermanniee (f Pl. II. fig. 3, part 1. p. 180) the seta is
derived from the two epibasal quadrants, and it is considered that
the cotyledon is homologous with it, and terminal. It arises before
other organs, bearing out the sporogonial or phyton theory of the
plant, but not the lateral appendage view. If their development
is similar, then the single cotyledon has dichotomised to form two ;
and the plumule is then lateral to both cotyledons, and between
the two. And the bifid cotyledon is primarily terminal. “The
actual ontogenetically terminal position of the plumules is a
secondary adaptation,” and in some plants, e.g. in palms, “it
becomes enclosed in the basal sheathing petiole of a single tubular
cotyledon, which is terminal to the hypocotyl.”

In the moss the seta is terminal. If we regard the cotyledon as
homologous and the monocotylous embyro an index of the history
of the leafy angiosperm, we have a succession of shoots arising, the
second like the first, and so on, till a terminal apex or growing point
is formed upon the axis thus produced by sympodial arrangement.
This monopodial axis with apparent lateral appendages is purely
illusory, and ‘‘we need to entirely amend our artificial conception

Stet), jo sys.
2 Vide ‘* A Study of the Vascular System in certain Orders of the Ranales,””
** Ann. Bot.,’” Oct. 1908, p. 651 ef seqq.

THE PAST HISTORY OF MONOCOTYLEDONS 219

of the leaf as a purely lateral appendage to an axis. We are misled
all along the line into believing these secondary modifications to be
the original types” (Pl. III. fig. 1).

From the embryonic segmentation and the dominant cotyledons
the usual view as to the appendicular nature of the leaf is wrong.
‘The plant is built up of a number of individuals (phytons) by
‘‘anaphytosis,” or sympodial arrangement. On this view monocoty-
ledons are the older. Angiospermous plants may be placed in one
of two categories according to their habit, viz. :—

Grandifoliate plants (monocotylous), stem subsidiary, with bundle-
system like that of the leaf, reduced internodes, leaves dominant,
closely succeeding, with wide sheathing base ; e.g. palms, water-lilies.

FParvifoliate (dicotylous), stem dominant, bundle-system differing
from that of the leaf, elongated internodes, small leaves, unsheathed
bases ; ¢.g. elm, wall-flower.

The grandifoliate is the more primitive, and is due to the
precocious assertion of the axis, displacing the terminal cotyledon,
which is constantly repeated and gives rise to the short
internodes.

Dr. Worsdell points out in regard to the fusion or abortion of one
cotyledon to account for the deviation of the monocotylous from the
dicotylar type, that in no instance has a monocotylous plant been
found to develop a bifid cotyledon or two cotyledons. On the
other hand, the converse is found where amongst dicotyledons one
is found.

This is held to be a reversion. Bifurcation of cotyledons is
common, and the polycotyledonary form seems to be the direction
evolution is taking.

Thus we come to the novel view recently promulgated by
Dr. Worsdell as follows: ‘‘ Hence I hold that the presence of two
cotyledons in dicotyledons is merely an illusive appearance. ‘There
is only one cotyledon, as in monocotyledons ; this has become so
deeply bifurcated in the great majority of cases as to give the appear-
ance of two distinct leaves.”

The sympodial arrangement of the stem and leaves is a recapitula-
tion of that of the seedling monocotyledon, and as such lends
support to their relatively primitive character.

In regard to the ancestry of Angiosperms generally, Dr. Worsdell
remarks :—

“] feel more and more inclined to hold the view that Angio-
sperms have developed directly from an ancestor belonging to the
Bryophyte level, and that they have not come from either Gymno-
sperms, Pteridosperms, or Ferns. On the other hand, it is quite
possible that they have descended from a fern-like ancestor.”

The tendency of Dr. Worsdell’s view is thus largely in the direction
of the point reached by Celakovsky and Bower.

Ontogeny does not always visibly repeat phylogeny. In seedling
structure some stages are omitted, and there is no room for scattered
bundles. The adult, however, should include all previous stages.

220 THE SCOTTISH BOTANICAL REVIEW

Leaves retain their primitive character longer than stems which
carry all the organs.

The stem is parvifoliate in Umbelliferae, the leaves grandifoliate.
If the monocotylous type be derived from dicotylar forms, the stem
would not remain the same and leaves become grandifohate ; but if
dicotylous forms are derived from monocotylous, the stem might
become parvifoliate and the leaves remain grandifoliate and stem
become parvifoliate. Scattered bundles are found in the stem of
Umbelliferee, and it is not truly parvifoliate.

In Ranunculacee the flower is primitive and the habit grandi-
foliate. They have evidently not been derived from a woody
parvifoliate type ; nor are Nymphzeaceze—said to retain their grandi-
foliate habit owing to adaptation to an aquatic habit, for that induces
reduction and concentrated bundles, to form a compact mass in the
centre of the stem. Indeed, they retain their habit although aquatic.
Berberidaceze, Calycanthacezs, Magnoliaceze possess primitive
flowers, sheathing leaf-bases, and retain something of the grandi-
foliate habit. The bundles are amphivasal where scattered—a
character common to both groups of Angiosperms.

Miss Ethel Sargant holds the view that both monocotyledons and
dicotyledons are monophyletic. The absence of a cambium is said
to be due to reduction. ‘The stem-structure of Liliacez is regarded
as primitive, e.g. in Anxemarrhena, with its bisymmetrical structure:
the last stage in the fusion of the cotyledons. Other groups of
Liliifloree exhibit the same type. Comparison with pseudomono-
cotyledons and with Evanthis and Podophyllum, the latter arrested
on its way to become a monocotyledon, confirms this, The correla-
tion of this feature with the geophilous habit suggests that it is the
cause of the fusion of the two cotyledons to form one, which is
regarded as a necessary economy by reduction of material expended
in assimilating tissue, owing to the short growing season of such
plants as Alpine, bulbous, and tuberous plants. In such plants the
leaves are crowded, giving rise to concentric or scattered bundles.
Linear leaves assist in elongation. Short-lived roots characterise
lowland plants habituated to a short growing season. Albuminous
seeds are found in dicotyledonous geophytes. ‘Thus Miss Sargant
holds the view that the monocotylous forms are reduced owing to a
geophytic habit.

Darwin, in his “‘ Movements of Plants,”! says: ‘“‘We may infer
that there is some close connection between the reduced size of one
or both cotyledons and the formation, by the enlargement of the
hypocotyl or of the radicle, of a so-called bulb.”

Pseudomonocotyledons are geophytes as well as many mono-
cotyledons, and thus the common habit is taken to be the cause
of the single cotyledon.

As to the possession of a cambium, Queva has found this in second-
year tubers of Gloriosa superba, from which he regards dicotyledons

1 1880, p. 97.

THE PAST HISTORY OF MONOCOTYLEDONS 221

as the older group. Professor Jeffrey finds cambial activity in
monocotyledonous seedlings in their third period when they become
ascending axes, and considers they are derived from dicotyledons
by adaptation to an amphibious habit.

The single cotyledon is terminal, but, if derived from dicotyledons,
is thought to be only apparently terminal, really lateral. In
Delphinium nudicaule (see P}. III. figs. 2 and 3) at eight weeks the
cotyledon is terminal, but at fourteen weeks the plumule is. In
Alisma (Pl. III. fig. 4, A), Zamus communts (tbid., 4, D), and the
pseudomonocotyledon Corydalis cava (tbid., 4, B, C) the cotyledon is
formed before the plumule and is terminal ; but when the plumule is
developed the cotyledon is forced to one side. This is explained
as due to the early formation of the plumule, or to the partial union
of one margin only, as in Ranunculus ficaria.

The other characters, parallel venation of leaf, short duration
of primary root, albuminous seeds, trimerous symmetry—except the
last—are regarded as adaptations to mode of life.

Miss Sargant reviews the theory of Professor Henslow. This is
based largely on the great number of aquatic monocotyledonous orders.
He considers monocotyledons are derived from dicotyledons by
suppression of one cotyledon, due to their aquatic habit. But
aquatics and geophytes have characters in common. Vascular
structure is reduced by an aquatic habit. Parallel venation is
found only in aquatic monocotyledons. Tufted roots are found in
aquatic monocotyledons and dicotyledons. Aquatic dicotyledons
do not tend to become monocotylous nor produce albuminous
seeds. The floral structure is not uniform, and may be derived from
an ancestor between Ranales and Liliflore. Probably aquatic
monocotyledons were not primitive but driven by severe competition
on land to the water, where conditions are more uniform. Aquatic
conditions tend to modify vegetative organs, but not the flower.
This is because it is a physiological phase.

Geophytes can adapt themselves either to an aquatic or a
climbing habit. Arboreal monocotyledons are not aquatic, but
possess anomalous thickening rings as in Aloe, Dracena, etc.
Palms, again, recall the characters of geophilous plants, being
sem1i-bulbous.

Of phylogenetic schemes, which embrace the other characters,
eg. parallel venation, trimerous floral structure, ephemeral primary
root, albuminous seeds, etc., there is Professor Lyon’s theory of the
origin of dicotyledons from monocotyledons by fission of one to
form two cotyledons, of use to the cotyledon. in forcing itself from
the seed. Agardh first proposed the alternative theory, but only
applied it to a few monocotyledons.

Professor G. Henslow extended this theory, deriving monocoty-
ledons from dicotyledons by suppression of the one cotyledon due
to the aquatic habit. This is based to a great extent on the large
proportion (33 per cent.) of aquatic orders. Miss Sargant regards
monocotyledons as a decadent race driven to an aquatic habit to

222 THE SCOTTISH BOTANICAL REVIEW

escape competition. The arrest of the cotyledon is found to be
confined to one aquatic dicotyledon. The others are geophilous.
The sheathing leaf-base is common to both groups, and is correlated
with suppression of internodes. The absence of a cambium and
substitution of concentric rings of growth may be due to the number
of broad-based leaves inserted on a short axis in the case of under-
ground aquatics, e.g. Wymphea.

In Peperomoia Mr. A. W. Hill found one cotyledon, but there
were probably originally two, and one is hypogeal, acting as a
sucker; the other looks like and performs the function of a true
leaf. He extends this theory to monocotyledons generally. It is
geophilous, with stem-structure like A/sma.

Then we have Miss Sargant’s own theory in which fusion of
cotyledons is correlated with a geophilous habit. Suppose the
primitive Angiosperm were geophilous. The green parts of the
seedling would be reduced in early seasons and fusion would result.

Podophyllum affords an example of a plant with united cotyledons
and scattered stem bundles which is a geophyte.

The suggestion is thrown out that the glacial period may have
produced conditions most favourable to geophytes and afforded
scope for monocotyledonous development.

In attempting to reconstruct the character of a race of primitive
Angiosperms! Miss Sargant discusses the question of monocotyledons
in a new light. The development from Pteridosperms, first as a
pro-anthostrobilus, then as a eu-anthostrobilus, up to the primitive
Magnolia and Liriodendron, as recently sketched out by Messrs.
Arber and Parkin,” is accepted, and she reconsiders the evidence for
regarding Angiosperms as monophyletic, and the reconstruction of
the pro-Angiosperm. As to the first, the community of descent of
monocotyledons and dicotyledons is proved by the many characters
they share in common. And it is pointed out that the angiospermic
flower is unique. The homologue of the flower, carpel, or endosperm
is by no means settled.

The germination of the embryo-sac and formation of endosperm,
where the first nuclear division, origin of endosperm from three
nuclei, and the uniform nature of the two processes is so characteristic,
supports the monophyletic origin. Double fertilisation is by no
means uncommon. The history of the development of the endo-
sperm is not known. Commenting on Miss Berridge’s suggestion
as to the resemblance between the endosperm of Ephedra and that
of Angiosperms, Miss Sargant says: ‘No tissue then in Gvetum,
nor, so far as we know, in lWedwttschia, can be considered as the
direct representative of the angiospermic endosperm” ;* and she

1“ The Reconstruction of a Race of Primitive Angiosperms,” Ann. Bot.,
1908, vol. xxil. p. 121 ef seggq.

2**On the Origin of Angiosperms,” Jour. Linn. Soc., 1907, vol. xxxviil.
p- 29. See further, Dr. D. H. Scott, ‘‘ The Evolution of Plants,” 1911, Williams
& Norgate.

° See, however, Professor Pearson’s recent researches on Welwr7tschia.

THE PAST HISTORY OF MONOCOTYLEDONS 223

thinks “the germination of the embryo-sac and the history of the
endosperm isolate monocotyledons and dicotyledons from all other
plants.” The flower also is distinct from other ‘“ flowers,” and
characteristic.

In the reconstruction of the primitive Angiosperm Miss Sargant
relies on the evidence adduced by Arber and Parkin as to the form
and character of the primitive flower, with angiospermic foliage,
flowers, seeds, and ovules enclosed in an ovary, three nuclei in the
pollen-tube, etc. The flower was elaborate in structure, with sepals,
stamens, carpels, spirally arranged in acropetal succession, as in
Magnolia.

The early types were of the form of a bisexual strobilus as in
Magnolia, with d sporophylls below, @ above, protected by a series
of sterile bracts forming a_basal perianth. Such a type is found in
the Jurassic Lenxnettiites. The immediate ancestor had an antho-
strobiloid flower, with perianth leaves, stamens, carpels, arranged
acropetally. The strobilus was the primitive form, and Beznettites
stands midway between it and A/agno/ia.

Monocotyledons and dicotyledons differ in stem-anatomy and
number of cotyledons. How did the primitive Angiosperm differ
from or resemble either? As to stem-anatomy, monocotyledons do
not possess a cambium, or rarely. G/ortosa, however, exhibits
collateral traces and a cambium in second-year tubers. Gymno-
sperms exhibit cambial activity, and so did many fossil vascular
cryptogams. Dicotylar seedlings early exhibit a cambium. Some
monocotyledons, as Yucca, etc., do. Thus evidence points to the
primitive Angiosperm possessing a cambium. As to the number of
cotyledons, monocotyledons possess one, dicotyledons two. No
monocotyledon possesses two, but several dicotyledons possess one.
Many groups of Gymnosperms are dicotylar, and the most primitive
Araucariez are polycotyledonous. In what respect did the primitive
Angiosperm differ from or resemble either recent group? Analogy,
as above, favours the primitive nature of the dicotylar type. What
do we learn from embryology within the embryo-sac? The whole
of the evidence in this connection centres around the position of the
cotyledon. The terminal position of the monocotyledonous form
with the derivation therefrom by fission of the two is not favoured
by Miss Sargant. Can a terminal cotyledon be a true leaf?
Opinions differ as to this. If extensions of the axis, they are
terminal and not true leaves. The difficulty is thought to be
avoided by considering the cotyledon as only apparently terminal,
and so lateral.

In Adisma (Pl. IV. figs. 1-6) the cotyledon is terminal at first,
then lateral as the plumule displaces it. In Pistia and Sparganium
early leaves are terminal. Are they cauline? Or are they due to
suppression of the axis? They are primitive forms and aquatic.
The stem exists in the mature /7s¢/a, but not in the embryo. The
reduction in stem is due to the aquatic habit, and causes terminal
leaves. Can it cause the cotyledon to be terminal ? The form of

224 THE SCOTTISH BOTANICAL REVIEW

the embryo and development of embryo-sac are affected by (1) space
rotations, (2) mode of food-supply, (3) shape of embryo, and depend
on future form of seedling and mode of freeing itself from the seed-
tissue. The embryo-sac is long and narrow, and bent upon itself,
evidently due to nature of surroundings. The suspensor conveys.
food. The cotyledons are absorptive, and endosperm prepares and
draws upon the store. Is the function of the cotyledons to absorb.
food and pass it on? The embryo exhibits adaptation to present,
past, and future environments. Wide embryo-sacs have a lateral
cotyledon, and narrow embryo-sacs, as in Adsma, an apparently
terminal cotyledon.

If the single cotyledon was lateral, was that of the primitive.
Angiosperm lateral? It is possible to imagine a lateral member
split into two, and that one member has revolved through 180°. If
it was dicotylar, the monocotylous form might be derived by
suppression of one or fusion of two members. The cotyledon was
single in pseudomonocotyledons, and derived from two presumably.
The development is like that of 7Yamus. Sterckx found in
Ranunculus ficaria a bilobed venation in the cotyledon which
suggested fusion of two cotyledons. Analogy with Gymnosperms
favours this, but development of the embryo-sac yields no evidence,
its shape being due to environment.

As regards embryology after germination, certain types of mono-
cotylous plants have primitive vascular tissue. In dicotyledons the
tetrarch type is found as well as the diarch type.

In Vigel/a it is diarch, but there is transition to a tetrarch root.
In Althea, which is tetrarch, there is an approach to diarch structure, _
like Liriodendron. The oldest Gymnosperms exhibit tetrarch struc-
ture and also diarch structure. In Pteridosperms probably the
structure was tetrarch. Thus presumably the primitive forms
were cither tetrarch or diarch. In monocotyledons the structure
is more variable, but there is a series of extreme types and also.
intermediates.

The common origin indicates presumably a primitive type. Such
are Liliacee, e.g. Anemarrhena, which is tetrarch. and they are con-
nected with others, ¢.g. Iridaceze, Aroidez, whilst it is in fact a radicle
of Liliifloree, and hence may be of monocotyledons in general, recapi-
tulating ontogenetically the history of the group. There is a strong
resemblance between the structure of Azemarrhena and the dicotylar -
A/thea, which is near the primitive gymnospermous type, and prob-
ably primitive. Hence probably the primitive Angiosperm type
had a tetrarch stem-structure. Would this necessarily be correlated
with two cotyledons? Dicotylar forms with cotyledons each sym-
metrical about the median plane are symmetrical about two vertical
planes. There is no symmetry about one plane, where reduction
occurs. In monocotylous stems receiving traces from a single lateral
there is only symmetry about the median plane. The primitive
Anemarrhena, Galtonia, etc., are exceptions. That the tetrarch
skeleton symmetrical about two planes is found in these suggests .

THE PAST HISTORY OF MONOCOTYLEDONS 225

that dicotylar forms gave rise to monocotylous by fusion of the two
cotyledons. In Ranales, e.g. Zranthis, the root is diarch, and tetrarch
structure is found in the hypocotyl. In Podophyllum peltatum the
same occurs. Anemarrhena and Althea are recalled in these. In
both the cotyledons are united for some distance. Reduction of
the tetrarch type has progressed further than in A/thea, of which
Anemarrhena is a reduced form, and with it Podophyl/ium and
Eranthis agree, showing that the union of cotyledons in a tetrarch
dicotyledon may result in a monocotylous type with structure of
Anemarrhena.

Professor G. Henslow, in a paper! read before the Linnean Society,
brought forward the theory of the origin of endogens from exogens
by adaptation to an aquatic habitat due to suppression of one
cotyledon. ‘This theory, which is an extension of Agardh’s, is based
upon the large proportion of monocotylous aquatic orders in the
group, viz. 33 percent. There are, however, some characters which
are found in both monocotyledons and dicotyledons, and those which
are thought to be confined to aquatic plants aione are also found in
geophytes and other groups. Aquatic conditions have certainly
helped to reduce the stem-structure and to modify foliage, but are
not responsible for the reduction in cotyledons, nor so far as is
known, the character of the flowers.

Messrs. E. A. N. Arber and J. Parkin,” in a paper on the “ Origin
of Angiosperms,” trace the ancestry of the group. They disagree
with Engler’s view that the present-day primitive Angiosperms have
unisexual flowers, e.g. Piperales, Pandanales, etc. But the perianth
is to be regarded as arising de novo, and to be an organ sud generts.

The piants cited have a sharply defined complicated structure.
This theory has proved barren. The strobiloid theory is held to
solve the question, the primitive form being amphisporangiate with
micro- and megasporophylls and a well-defined perianth. The parts
of the flower were numerous, arranged spirally, the female sporo-
phylls above the male. Such a type is found in some Magnoliacee,
Ranunculacez, Alismaceze. The apetalous unisexual flowers are
regarded as derived from these. The flower is an anthostrobilus of
which a gymnospermic and an angiospermic form can be distin-
guished. But they are modelled on the same plan, the female
and male sporophylls occupying the same relative position, and
the perianth is differentiated. These authors consider that some
important change in the mode of pollination caused the advent of
Angiosperms.

Dealing with the ontogeny of Angiosperms in the paper already
cited, Bessey defines the difference in the development of the em-
bryo in monocotyledons and dicotyledons as follows :—

‘The only histological difference which is measurably constant is

1 «* A Theoretical Origin of Endogens from Exogens by self-adaptation to an
Aquatic Habit,” Journ. Linn. Soc., vol. xxix., 1892, p. 485. But see a more
recent paper, 74a.

2 Zbtd., ‘‘ Jour. Linn. Soc.,” 1907, p. 29.

VOL. I. y

226 THE SCOTTISH BOTANICAL REVIEW

that the longitudinal division of the embryo takes place defore! the
formation of transverse walls in dicotyledons, and afterwards ' in the
monocotyledons. To this rule, however, there are numerous
exceptions.”

Commenting on the further progress of this division, he says, “‘ It
is at least an interesting coincidence that in the young embryo the
undivided foliar cells give rise to the single cotyledon, and the epi-
basal cell to the pair of cotyledons.”

The following remarks are so @ propos that they are quoted at
length :—

‘‘Are there any indications which may help us to answer the
question of the origin of these two groups? Have dicotyledons been
derived from monocotyledons, monocotyledons from dicotyledons,
or both from a common ancestor? It must be admitted that on
theoretical grounds it is no more difficult to pass from two cotyledons
to one, than from one to two. Indeed, there have been not a few
botanists who have suggested the derivation of the monocotyledons
from the dicotyledons. When, however, one compares the two embryos
there is a slight preponderance in favour of the view that the struc-
ture is a little higher in dicotyledons than in monocotyledons. The
row of undivided cells in the embryo of the monocotyledon after the
third or fourth segmentation is certainly a lower structure than the
compact mass of cells constituting the ‘octant stage’ of the dicoty-
ledonous embryo. ‘The cotyledons themselves afford a slight sugges-
tion as to the relationship of the two groups. It is a well-established
principle in embryology that embryonic stages of higher organisms
resemble the adult stages of the organisms which are lower in the
same genetic line. Applying this principle to the cotyledons, we
observe that while they bear some similarity to the leaves of both
monocotyledons and dicotyledons, the similarity is a little more
marked in the case of the monocotyledons. Compare the mostly
sessile, often clasping, usually elongated leaves of monocotyledons
with the cotyledons of either class, and contrast these with the
mostly petiolated, generally not clasping, and usually broad-bladed
leaves of the dicotyledons.”

Bessey also refers to the fact which Sir J. Lubbock (now Lord
Avebury) points out, that the earlier leaves of a plant are generally
quite different from the later ones. Young plants are the most
alike, and they begin to get more unlike as they mature. So that it
is often only possible with seedlings to distinguish the subclass,
later the family, and then the genus, whilst until the flower is pro-
duced the specific characters are hidden.

The deductions he makes from ontogeny are :—

1. All Angiosperms are essentially alike.

2. The two subclasses (monocotyledons and dicotyledons) appear

to be modifications of a common type which diverged from
one another at an early period.

1 Present writer’s italics.

THE PAST HISTORY OF MONOCOTYLEDONS 227

3. There is no indication that either subclass was derived from
the other.

4. There are some structural indications that the monocoty-
ledons must rank lower than the dicotyledons.

5. The vegetative rank of most cotyledons is so nearly the
same as to have left no vestiges on the young plant, which
is itself vegetative.

6. The groups into which dicotyledons and monocotyledons
are divided are “‘ flower subdivisions” of a greatly multiplied
rather common vegetative structure. Therefore, we may
not expect to find upon the embryo or immature plant any
vestigial record of their origin.

In discussing the results derived from morphology, Bessey, in
speaking of tissue systems, draws attention to the short life of
monocotyledons and the long life (often amounting to hundreds of
years) of dicotyledons. But the ephemeral type of the former occurs
amongst the latter, which indicates, he suggests, the close relation-
ships of the two subclasses.

He points out that the young stems of monocotyledons and
dicotyledons differ less than old stems, and remarks that “ the
important difference between these two types of stem is that the
skeletal tissues continue to form a single solid column in the dicoty-
ledons while they do not in the monocotyledons,” and the further
fusion of bundles in the former, due to cambial activity, marks them
out as a higher group.

From the normal type of stem seen in lilies, naiads, and orchids,
there are two departures in the stem with elongated internodes of
grasses and sedges, and the shorter internodes of palms and screw-
pines. As to their leaves he says :-—

“The leaves of monocotyledons are usually entire, elongated,
parallel-veined blades, placed alternately or scattered upon the
stem, to which they are attached directly (in sessile leaves) or
indirectly (in petioled leaves) by a common broad base which is
rarely supplied with stipules.”

When contrasting these with dicotyledonous leaves which are
dentate or lobed, usually broad, net-veined, opposite and alternate,
attached by a narrow base, commonly supplied with stipules, these
differences are said to be “due to differences in development.”
The parallel veins of monocotyledonous leaves are due to localisation
of growth at the base of the blade or indefinite bands on each side
of its axis. The broad basal attachment of the monocotyledonous
leaf may be due to the looser disposition of the fibro-vascular bundles
in the stems, or perhaps may be owing to the fact that they are not
so far advanced as the leaves of dicotyledons, there being an
articulation at the base of the latter only. In the monocotyledons
the common leaf type is especially modified in the Graminez.

The flower is held to consist of a strobilus of ‘‘ flower-leaves ”
consisting of sterile leaves and spore-bearing leaves, the former being
derived according to Bower, from the latter. The perianth as a

228 THE SCOTTISH BOTANICAL REVIEW

ring of sterile leaves may be united by symphysis, or may be
apetalous. The last condition is no indication of affinity, but is
simply a case of reduction.

The pollen leaves (microsporophylls, stamens) lie between the
sterile and ovule leaves, and are largely adapted to the seed-producing
or pollen-distributing capacity.

The ovules (macrosporophylls, carpels) occupy the highest portion
of the strobilus, being usually separate, and the carpels are often
separate as in A//sma, etc., forming apocarpia, or united as in lilies,
to form a syncarpium, with some portions still free as in the latter,
or with styles united all their length as in Commelinacez.

When some other organs are attached to the syncarpia as in the
epigynous irids and orchids, ‘‘there has been such a fusion of the
originally separate parts of the strobilus as to result in a single
compact structure.”

‘The primitive syncarpia of the monocotyledons appears to have
contained three carpels as in lilies.” In the fusion some parts are
suppressed. In some dicotyledons, e.g. Caryophyllales, etc., ‘ there
is a tendency to a reduction in the number of parts with increased
fusion of the strobilar leaves.”

“The same law is illustrated in the genetic line, which includes
the lilies (Coronariez), pipeworts (Eriocaulacez), sedges (Cyperacez),
the lower grasses (Bambusez), and higher grasses (Agrostide and
Panicez) ; or possibly still better in the line from lilies to amaryllids
(Amaryllidacez), irises (Iridaceze), burmannias (Burmanniacez), and
orchids (Orchidacez).”

There are then two modifications of the strobilus by (a) symphysts,
fusion of parts; (4) aphanisis, a suppression of parts.

They may affect some or all organs of the strobilus separately or
jointly.

The conclusions arrived at from morphology are :—

1. The identity of the cells and tissues of the two subclasses of

Angiosperms indicates their close relationship.

2. The fibro-vascular tissue-system of the dicotyledons indicates
that the subclass is higher than the monocotyledons.

3. The roots of dicotyledons indicate that this subclass is higher
than monocotyledons, and suggest the possibility of the
origin of the former from the latter.

4. The structure of the dicotyledonous stem indicates the highest
rank of this subclass.

5. Among monocotyledons the external morphology of the stem
indicates the derivation from lily-like plants of the palms
and screw-pines by an excessive shortening of the internodes,
and of sedges and grasses by a corresponding elongation.

6. The general morphology of the leaves of monocotyledons and
dicotyledons, as has already been indicated many times,
emphasises the close relationship of the two subclasses, and
repeats the suggestion that the former includes plants which
must take rank below the dicotyledons.

THE PAST HISTORY OF MONOCOTYLEDONS 229

In dealing with the hypothetical phylogeny of Angiosperms,
Bessey refers to the monocotyledons as follows :—

‘‘The modifications which gave us the main lines of monocoty-
ledons were first the fusion of the carpels with one another and the
production of a syncarpium, and, second, the progressive fusion of

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The Phylogeny of Angiosperms. (After Bessey.)

the syncarpium with the other strobilar leaves. These resulted in
the phylum, which begins with Apocarpz and passes to Coronariez,
Epigynz, and Microsperme. In some Apocarpz and many plants
of the type of the Coronariez the perianth has been more or less
reduced (by aphanisis), in some cases amounting to complete
suppression, as in palms (Calycinz), aroids (Nudiflorz), and sedges
and grasses (Glumacez),”

230 THE SCOTTISH BOTANICAL REVIEW

As to their taxonomy, he writes :—

“The Angiosperms are separable into two diverging subclasses—
the monocotyledons (Monocotyledonez), and the dicotyledons
(Dicotyledoneze), the first ranking structurally lower than the second.
‘The monocotyledons are well divided by Bentham and Hooker into
seven series, and these we may accept unchanged, with the single
exception that the water-worts (Hydrocharitaceze) should probably
be removed from the Microsperme to constitute an additional
co-ordinate group. These eight groups, which appear to be
deserving of no more than ordinal rank, should then be arranged
so as to have the following sequence, namely, Apocarpze, Coronariee,
Nudifloree, Calycinze, Glumacez, Hydrales, Epigynze, Microsperme.
Here it must be understood that the Nudiflorz, Calycinz, and
Glumacez are separate orders radiating from the present order
Coronarieze, and that the Hydrales constitute a diverging order
from the base of the Epigyne.”

Some researches made by Professor H. H .W. Pearson ! in regard
to Welwitschia are of interest in this connection. In studying the
endosperm he found all the cells are multinucleate. Later, each
cell possesses only one nucleus by fusion of nuclei in young cells.
The original nucleus by division has become a thousand. The
embryo-sac elongates, and there is a redistribution of nuclei. Two
poles are formed: one at the micropylar, the other at the opposite
end. In the former the nuclei are not crowded, in the latter they
are. Then a division into compartments or segmentation takes
place, with 2 to 6 nuclei at the micropylar end, 12 or more elsewhere.
When there are more than 6 nuclei they are divided up into cells
with single nuclei; but those at the micropylar end are not so
affected, as each wall grows up into the prothallial tube. The
nuclei and cytoplasm pass into this, and the nuclei become
functional gametes. This unusual characteristic has led Professor
Pearson to regard the endosperm as representing a new stage, which
he calls the trophophyte, which serves to mark its intercalation into
the life-history—a terminology now familiar to us from Professor
Bower’s theory of the antithetic generation or intercalation of the
sporophyte into the life-history of the moss plant. This trophophyte
is neither the sporophyte nor gametophyte, but is regarded as
phylogenetically related to the endosperm of Angiosperms by a
series of reductions and specialised stages. The fact that in Angio-
sperms a similar procedure takes place is corroborative of this view.
In a more recent paper? Professor Pearson finds that the ovule in
Welwitschia is cauline, and it is considered that the female cone
and male flower are derived by reduction and specialisation from an
amphisporangiate strobilus of a type similar to Benmettites. After
division 1024 nuclei are contained in the embryo-sac, and it is
repeatedly divided. The endosperm of the primitive Angiosperm

1 “¢Rep, Brit. Assoc.. Dublin,” 1908.

2 “Wurther observations on Welwztschia,” ‘‘ Proc. Royal Soc.,” Nov. 1908.

THE PAST HISTORY OF MONOCOTYLEDONS 231

is thought to have been homologous with that of Welwitschia.
Gnetum and Welwitschia are derived from the Angiospermic stock,
but before the carpel became the pollen-receiver.

In the Penzacez, a group of dicotyledons, Miss E. L. Stevens !
finds that there is no polarity in the early stages of development of
the endosperm, as in most Angiosperms. The embryo-sac contains
not 8 but 16 nuclei which form four egg-apparatus. By the
fusion of 4 nuclei a definitive nucleus results. By this reductional
process the same features met with in We/wztschia are recalled. A
very similar process is recognised in Getum by Lotsy in the germina-
tion of the megaspore or embryo-sac, and the development of the
endosperm.

5. CONCLUSIONS.

Summing up the evidence afforded by the different views held
by the writers cited, and by personal observation and experience,
it seems to us that systematic researches have failed to appreciate
the results of paleeontology and of morphology, and in considering
Pandanus, etc., as the oldest group sufficient attention has not been
given to the comparative results of floral structure or of morphology.

As to the theories based upon either embryology or morphology,
Dr. Worsdell’s view seems to us to be based upon the most stable
evidence.

The view that there is only one cotyledon in both groups, indeed,
is perfectly reasonable ; and if the cotyledon and the seta of Bryo-
phytes are conceded to be homologous, then their respective order
in time is also practically settled, for the single cell in the latter
gives rise to two, and so on until, just as in Cafsed/a, Pl. II. fig. 3
(part i.), p. 180, we arrive, in due course, at the bifid cotyledon.
The fact of the subdivision from an original single cell presupposes
the primitive character of the latter, and the derived nature of the
double row of cells.

As to the position of the cotyledon, if the above view is accepted
the originally terminal character of the cotyledon also follows. The
lateral position subsequently, if we adopt the sympodial arrangement,
does not affect the question. The lateral appendage view does more
harm, it seems to us, in obscuring the sequence of the ontogeny
(and so the phylogeny), and the morphological significance of organs
than any other theory promulgated.

The reversionary nature of the monocotyledonous type in Ranales
is in itself more or less favourable to the earlier age of monocoty-
ledons and the derivation of dicotyledons from them.

Miss Sargant’s views are in direct contrast to those of Dr. Worsdell’s.
This is readily understood when it is seen that she has based her
evidence mainly upon the earlier embryological features of the
species studied, too early in the ontogeny for a proper recapitulation
to be observed. Dr. Worsdeil, however, takes the most mature

1 ** Proc, Roy. Soc. S. Africa,” 1908,

232 THE SCOTTISH BOTANICAL REVIEW

structures in his investigations, and there he obtains the full history.
One is specially grateful to both of these able investigators for thus,
in their several fields, enabling us, by their combined researches, to
study both early and late stages. However much we may differ in
theory from either one or the other, the data upon which they have
been based lose none of their value whatsoever. In some respects
the plants Miss Sargant studied were anomalous forms, e.g. Pseudo-
monocotyledons, and primitive Liliaceze, and thej evidence is thus
inconclusive. ‘The lateral appendage view only leads back again to an
unnecessary transposition, in time, of the mono- and di-cotylar types.

The interpretation of certain features, e.g. the absence of a
cambium, as due to reduction, instead of being held as a primitive
character, is a necessary consequence.

There may really be some truth in Professor Lyon’s view that the
fission of the cotyledons has to do with the freeing of the cotyledon
from the testa.

Professor G. Henslow’s theory that one cotyledon has been
suppressed in monocotyledons is based upon the preponderance of
aquatic orders in monocotyledons—and thought to be due to their
adaptation to an aquatic habitat, a physiological cause, which is not
universally applicable.

In so far as it touches the origin of monocotyledons Messrs.
Arber and Parkin’s paper is favourable to the view that a primitive
Angiosperm gave rise very early to both mono- and di-cotylar types,
the divergence being close to the origin of the primitive ancestor.

Bessey’s views seem to us to be so reasonable that, taken in con-
junction with Dr. Worsdell’s, they lie nearest the truth.

It is a remarkable fact that the earliest forms of both monocoty-
ledons and dicotyledons exhibit a sort of parallelism of development.
The first monocotyledons are apocarpous, just as are Ranunculacee,
amongst the latter (except Nymphzeaceze, which are epigynous).

Then there is a higher series which becomes hypogynous, from
which the Calycifloree early diverge and are perigynous or epigynous.

So in monocotyledons the apocarpous forms (rarely petaloid)
become syncarpous, and then spadiceous or glumaceous, or hypo-
gynous, and finally epigynous.

The researches of Professor Pearson and Miss E. L. Stevens as to
the embryology of We/witschia and Penzacez favour the derivation
of Angiosperms from Gymnospermic or Bennettitean stock, but,
as yet, throw no light upon the Angiospermic origin nearer the
present time, and the differentiation into mono- and di-cotyledons.

Speaking generally, it seems to us that the monocotyledons and
dicotyledons have diverged from a common primitive Angiospermous
stock. The earlier types of the former appear to belong to
Alismacez, leading on to Liliacez ; and the earlier forms of the latter
the Ranales, represented by Magnolia and Liriodendron, from which
Ranunculus and its allies are later derivatives. This would allow of
the retention of some primitive characters in both groups which are
apparently explained only by supposing dicotyledons derived from

THE PAST HISTORY OF MONOCOTYLEDONS 233

monocotyledons or vice versa, or from a stock capable of transmitting
either.

Primitively we may consider that there was an adaptation from
aquatic to terrestrial conditions, which then gave rise to the
geophilous habit, whilst other forms in the harder struggle on land
retained or sought refuge again in an aquatic habit.

The derivation of Angiosperms from the same synthetic type as
Gymnosperms seems clear. The near approach of the unique Bennet-
titeze with two cotyledons and copious endosperm would alone suggest
this. The derivation of the latter from Pteridosperms is again more
or less clear. The fact that two cotyledons is so characteristic a
feature of Gymnosperms is again confirmatory of the early differentia-
tion of a dicotylous type. The polycotyledonary! forms are also
the more ancient, so that there is some evidence from analogy for
reduction. In Gymnosperms a suspensor is not always present.
Many of the Gymnosperms exhibit other characters, though these
may be homoplastic—e.g. parallel-venation and strap-shaped nature
of leaves, e.g. in Cordattes.

It should be noted in regard to stem anatomy being a distinctive
character, that the arborescent Lycopods sometimes, though not
invariably, exhibit cambial activity, whilst herbaceous forms do not.

As to cotyledons the number is extremely variable amongst
Gymnosperms within the same group. So that although dicotyle-
dons and monocotyledons are remarkably uniform in this respect, yet
the character is apparently due to physical selection and is homo-
plastic, like the stem anatomy.

Probably Dr. Worsdell’s suggestion that the two are phases of one
group is nearer the mark. Amongst fossil Cephalopods the group
Nautiloidea is more or less constant in the central or dorsal position
in the siphuncle, whereas in Ammonoidea the siphuncle is ventral.
In Clymenioids, an Ammonite group, however, it is dorsal.

Triassic Ammonoids, again, exhibit a curious type of sutural
development. In Nautiloidea Triassic Pleuronautilids exhibit what
may be called an homologous arrangement.

Thus monocotyledons and dicotyledons may be compared with
Nautiloidea and Ammonoidea, and although the first two are plants,
and the last two animals, they exhibit a sort of parallel mode of
development, in respect of the degree and not the kind of difference.
The phenomenon is what Hyatt has described as morphic equivalents.

Summing up the evidence in regard to the relative antiquity of
the monocotyledons and dicotyledons, and the bearing this has upon
the origin of the latter, we find ourselves reduced to the following
conclusions :—

1. There is evidence from paleontology that monocotyledons
came into existence almost contemporaneously with dicotyledons,
not, as once held, long prior to the latter.

' There may be 3 to 18 cotyledons as in Pemus, 3 to 8 in Adzes, 3 to II in
Picea, 6 to 11 in Cedrus.

234 THE SCOTTISH BOTANICAL REVIEW

2. All but a few of the families and orders (6) have been recog-
nised in a fossil state, and most of these in the Jertiaries.

3. The group appears to have diverged from the primitive Angio-
spermic stock, evolved from a line of ancestors connecting Pterido-
sperms, Bennettitales, and eventually such dicotyledons as AZagnolia,
Lirtodendron, Nymphea, and the monocotylous Alismaceze and
Liliacez.

4. There is every appearance of reduction in the floral symmetry,
but the characters of the cotyledons, stem, and Jeaves are apparently
as primitive as those of the primitive dicotyledons—further than which
latter they have not progressed. The uniform aquatic habit would
account for a great deal of this in stem and leaves, and even cause
some reduction. In other words, monocotyledons are retrogressive
and dicotyledons progressive types.

5. The prevalence of aquatic, geophytic, and climbing genera lends
support to the view that they are a weaker or degraded race of
Angiosperms which have sought refuge in aquatic habits, thus avoid-
ing the severer struggle for existence upon land.

6. The view is held that the difference in cotyledons is due partly
to the shape of the embryo-sac and supply of endosperm, partly to
the mode of life of the plants after germination.

7. The stem structure is due to physiological causes, since the tree
type is more or less absent, and where present traces of a cambium
often occur.

8. The absence of any recapitulation during the ontogeny of
monocotyledons of a supposed previous dicotylar stage is regarded
as conclusive evidence that the former are not derived from the
latter.

g. It is held that the primitive Angiosperms resembled an
Alismaceous or Liliaceous type, on the one hand, and a Ranalian
type on the other, and that the monocotyledons and dicotyledons
are divergent series from a common ancestor, recalling the common
characteristics of the two as the one or the other were called into
being by either physical or natural selection. In the one case,
dicotyledons, there has been progression and differentiation; in the
other, monocotyledons, retrogression and even some reduction from
a common ancestor of the primitive Angiospermic type.

b of

be okey x it

PLATE, TL,

Fig. 1. (A) A shoot of Polyalthza, to illustrate sympodial arrangement. Every
stem segment is terminated by a leaf. (B) Monocotyledonous embryo,
to show similar arrangement. (After Worsdeil, “‘A Study of the
Vascular System in certain Orders of Ranales,” Oct. 1908, ‘‘ Ann.
Bot.,” p. 653.)

Fig. 2. Delphinium nudicaule, seedling after eight weeks, to show position of leaf
and cotyledon.

Fig. 3. Delphinium nudicaule, seedling after fourteen weeks.

Fig. 4. A=Alisma plantago; B, C=Corydalis cava; D=Tamus communis.
Mono- and di-cotyledons for comparison to show position of cotyledon.

(Figs. 2, 3, and 4 after Sargant, ‘‘ The Evolution of Monocotyledons.”)

Scottish Botanical Review. } [Vora

A. R. Horwoop. PEATE, Dik:

PLATE IV.

Figs. 1, 2 and 3. Stages in the germination of the seedlings of Alzsma plantago:
eight times natural size.

Fig. 4. The same x 4.

Fig. 5. a natural size,

Fig. 6. 5 e

(Figs. 1-6 after Lord Avebury, ‘‘ Seedlings.” Cot, =cotyledon. )

Scottish Botanical Review. |

PLATE IV

A. R. Horwoop.

SHORT NOTES 235

Short Notes.

Saxifraga atzoides, L.—Among plants from Orkney sent by Mr. M.
Spence are two specimens of the above Saxifrage, and answering to
the description of the f. awrantia of Hartmann, “ Vet. Ak. Handl.,”
1818. The leaves are orange, shading to yellow at the apex. The
lower stem leaves are suffused with red, and the fruit is orange-red
(only half-ripe). The leaves are quite entire, very thick, with here and
there a long patent hair. This form occurs in Norway, with the
orainary form in Sweden, and rarely in Russian and Finnish Lapland.
Is not the figure in “‘English Botany,” t. 59, quite a rare form? I
have not seen such stem leaves on any specimen as there represented,
and the description says “‘rarely denticulate.” A. BENNETT.

Utricularia vulgaris L., in Caithness.—Good specimens of the
above (though not in flower) have been sent me by Miss I. Lillie
and Mr. G. Lillie from Loch Watten on the east coast of Caithness.
There are several of the winter buds ; these are strongly setose, with
translucent, spinose-like hairs, and the young leaves are spinose-setose.
The young bladders are semi-transparent, the older ones also, but
with the addition of a yellowish nucleus. This definitely records
the species for the county. A. BENNETT.

Cnicus oleraceus Lin. = Cirsium oleraceum Scop.—This plant has
occurred this season on a marshy meadow by the side of the Tay,
left bank, about a mile below Perth. It forms a small patch com-
prising about sixty or seventy flowering shoots and was certainly
not there, at least in the flowering stage, till this season. How it
came I am unable to conjecture, as it is certainly not a plant likely
to be cultivated, and the meadow, though cut, consists of natural
herbage only, and is never sown. Mr. Arthur Bennett informs me
that it occurred in Lincolnshire from 1832 to 1840 on the Fen
banks, and I believe it has been found in Scotland amongst other
casuals. The plant was named for me at the Herbarium of the
Royal Botanic Garden, Edinburgh. It has yellow flowers. Near
to the same place a little patch of Samgutsorba canadensis Lin. has
kept its place amongst some alder bushes for more than half a
century. Till this year I have never seen any signs of its spreading.
But this summer at least three other patches, at about a hundred
yards further down and clear of the bushes, have made their
appearance and have flowered freely. One patch in the middle of
the meadow forms a circle of a good many yards in diameter and
comprises a considerable number of plants. Possibly the very hot
summer of last year may have more thoroughly ripened the seeds
and thus enabled the plant to extend its area.

Juncus tenuis Lin.—In going through Glen Ogle in July this sedge

236 THE SCOTTISH BOTANICAL REVIEW

occurred in great plenty by the roadside about half-way between
Lix and the summit of the pass. For about a quarter of a mile it
formed a close band along the left side of the road. For about
another half mile I traced it in patches and isolated plants. It
seemed as if carts loaded with American fodder had been passing
along the road dropping seeds at first thickly and then more thinly.
Certainly the plant is not native in any of the Perthshire stations in
which it has been found. W. Barctay.

Notes from Current Literature.

“Quel est le vrai caractére biologique du Raphanus Raphanistrum,
L., et du Sixapis arvensis, L.,” by A. Verhulst (‘Contributions from
the U.S. Nat. Herbarium,” vol. xvi. part iii.).

“The North American Species of Mymphaa,” by G. S. Miller,
jun., and P. C. Standley (‘Contributions from the U.S. Nat. Herb-
arium,” vol. xvi. part iil.).

“Studies in Tropical American Ferns,” by W. R. Maxon (“ Con-
tributions from the U.S. Nat. Herbarium,” vol. xvi. part iii.).
Particularly valuable in containing a monographic account of the
species of Hemitelia, subgenus Cxemidaria, fully illustrated.

“« Spirea Ulmaria, L., and its Bearing on the Problem of Xero-
morphy in Marsh Plants,” by R. H. Yapp (“Annals of Botany,”
July 1912).

“* Polygala vulgaris, L., var. grandifiora, Bab.,” by Arthur Bennett
(‘Journal of Botany,” July 1912).

“A New Variety of Parnassia palustris,” by W. G. Travis and
J. A. Wheldon (“ Journal of Botany,” August 1912).

‘Le genre Balsamocitrus et un nouveau genre voisin, .#glopsis,”’
by W. T. Swingle (“ Bull. Soc. Bot. de France,” vol. xi.).

“‘Beauveria, nouveau genre de Verticillizces,” by M. Paul
Vuillemin (‘ Bull. Soc. Bot. de France,” Jan. 1912).

‘“ A propos de la position systematique de Adoxa Moschatellina,
L.,” by H. Lonay (“Contributions from the U.S. Nat. Herbarium,”
vol. xvi. part ili.).

“ Betula pubescens x nana in den Alpen,” by Dr. F. Vierhapper
(““Verhandlungen der K. K. Zoo.-bot. Gesellschaft in Wien,”
vol. 1xi.).

“A Beehive Fungus, Pericystis alvii, Gen. et Sp. Nov.,” by A. S.
Betts (‘‘ Annals of Botany,” July 1912).

NOTES FROM CURRENT LITERATURE 237

“Bacteria and Other Fungi in Relation to the Soil,” by D. Rivas
(** Bull. Soc. Bot. de France,” vol. xi.).

‘“‘Nouvelles maladies de plantes cultivées,” by Arséne Puttemans
(“Contributions from the U.S. Nat. Herbarium,” vol. xvi. part iii.).

“Ecologie d’une petite panne dans les dunes des environs des
Dunkerque (Phanerogames et Cryptogames),” by M. Bouly de
Lesdain (‘‘ Bull. Soc. Bot. de France,” vol. xii.).

“ Nouvelle contribution a la Géographie Botanique du Jurassique
Belge: Dispersion du Czrctum acaule, Allioni,” by A. Verhulst
(‘ Bull. Soc. Royale de Bot. de Belgique,” 1911).

“‘ Vegetationsskizze der Umgebung von Czernowitz,” by Dr. K.
Rudolph (‘‘ Annals of Botany,” July 1912).

‘‘Leaf Movements in the Family Oxalidacez,” by E. B. Ulrich
(‘Contributions from the Botan. Laboratory of the University of
Pennsylvania ”).

‘Ein neuer Typus der Spaltdffnungen bie den Saxifragaceen,” by
B. Hryniewiecki (“ Bull. International de l’Académie des Science,
Krakaw,” March 1912).

“Azotobacter Studien: I. Morphologie und Cytologie,” by A.
Prazmowski (‘‘Contributions from the U.S. Nat. Herbarium,”
vol. xvi. part 111.).

Reviews, Book Notices, etc.

CLARE ISLAND SURVEY. Parts XI.and XII.: Musci anD HEPATIC.
Canon H. W. Lett, M.A. tg12. Price 6d.

THE investigation of the cryptogamic flora of the Clare Island
survey district is one which cannot fail to stimulate survey work
elsewhere. If so remote a region, though botanically important, can
receive such systematic attention at the hands of busy specialists, it is
to be hoped that other areas may in due course be similarly dealt
with, including the many unexplored parts of Scotland, Wales, and
England.

Canon Lett in his interesting paper describes the district from a
bryological point of view, noting the absence of glens, the exposed
surfaces, the peaty districts much denuded by peat-cutting, which in
Ireland has helped to restrict the distribution of turfophilous plants
on a wide scale, though here it supports few mosses or hepatics.
They abound, however, in well-sheltered nooks and corners which
absorb the abundant rain. Croaghmore affords shelter to leeward

238 THE SCOTTISH BOTANICAL REVIEW

on the N.E. slopes for a rich flora, while on several low hills running
N.W. by S.E., with intervening valleys owing to peat-cutting, they are
rare. Saxicolous species are also rare. Though the district is now
intersected by walls, they have not as yet been covered by mosses,
and rupicolous species grow best near Knocknaveen (729 feet). Salt
is not responsible for their scarcity, for many appear to be halophytes
here. The loughs yielded no good results, sphagna being mainly
restricted to Creggan Lough. Arboreal species are absent, except in
plantations, and cutting down of forests has been a factor in extinc-
tion here, leaving only sphagna as relics. Some arenicolous species
occur on the shore. Canon Lett indicates the most favourable spots
for bryological observation as at Croaghaun Mountain and Knocka-
curram. No work had been done in Clare Island previous to this
survey, though Achill Island had been investigated.

Interest attaches to the statistics of the flora given, the total being
221, of which seven hepatics are new to Ireland and two are second
records. Notes are given as to the distribution of the flora, and the
absence of some is emphasised. The flora of Clare Island is said to
owe its origin to dispersal of spores by the wind. Following the list
of mosses and also that of hepatics come notes on the rarer species,
in which the respective areas of occurrence are indicated by I. and
M., indicating Clare Island and the mainland. A useful bibliography
is appended.

Part XIII.: Funct. By Car.ieton Rea, B.C.L., M.A., and Sir
H. C.. Hawtey, Bart. 1912, Preeias

THE account of the fungi is divided into two sections, Clare Island
being described by the second author, the mainland by the first. In
so far as the mycology is concerned this region was previously quite
unexplored, wo species only being recorded by Adams in 1910, and of
1400 Irish species 100 only were known in Connaught. The species
and varieties discovered number 802, including 295 species and
12 varieties new to Ireland, 11 new to Britain; whilst Hawley
describes a new genus, Candelospora, genotype C. tlictcola sp. nov.
(figured), and Rea a new species, Mygrophorus sgquamulosus
(unfigured). The flora of the island differs from that of the
mainland, especially in the number of Autobasidiomycetes. The
island is bare and wind-swept, the mainland sheltered, fertile, and
wooded. It is remarkable that Hypoxylon coccineum, so common on
beeches in S. England and elsewhere, is unknown. MHawley’s work
consisted of some three weeks’ collecting, resulting in a list of 283
species, with but few coprophilous species and the minute Fungi
Imperfecti. The chief woodland is in the N.E. There are two
districts: (1) exposed pastures, and (2) cultivated tracts. Several
interesting maritime specimens were collected, on Plantago maritima,
etc. The absence of woodland determines the relative poverty of
Autobasidiomycetes here, though there is an excess of twenty found in

REVIEWS, BOOK NOTICES, ETC. 239

the Faeroes, with the flora of which comparison is made, and where too
it is to be remarked that there were few pratal species. In this list
101 are new to Ireland, eight are new to Britain, one new to science.
Mr. A. D. Cotton found all the maritime fungi. Notes follow the
list, in which no localities are given, though most were found near
Westport. Hawley here separates Hyfoxylon pileatum and H. terrt-
genum. He found the spores of A/icroglossum atropurpureum \sarst.
variable, and suggests Lpicymatia Balani Wint. is better placed under
Mycospherella. A South American species, Phomatospora argentina
Spegg., first found here, is said to differ but little from the European
P. ovalis (Pass.) Sacc. The genus Candelospora is regarded as allied
to Microsporium in the penicillate branching well shown in the
figure, differing in the single terminal conidia. It is found on dead
holly leaves.

Rea was unfortunate in his choice of weather, meeting on the
mainland with snow and drought. The best wood he found was
Brackloon, where there are oak, beech, pine, sycamore, holly and
alder, and much leaf mould. The species of Cortinarius were
especially numerous and in good condition. He records 667 species
and varieties, 232 new to Ireland, five new to Britain. In this list
localities are given, which is a distinct advantage. Corynedla
glabrovirens is regarded as allied more closely to Calloriacee than
Bulgariacee. Urceolella incarnatina (Quel.) Bond., new to Britain, is
not considered to be related to any other British genus. Avthostoma
saprophilum E. and E. is new to Europe. The new species //ygro-
phorus squamulosus resembles Zricholoma repens in the tomentose
margin of the piles, and differs from A. o/ivaceoalbus Fr. in the
floccose squamules, characters sufficiently distinct to warrant a new
name. Both accounts of the Clare Island fungi are worthy of
careful study.

Tue Genus Fumaria IN Britain. By H. W. Pucstey, B.A.
Supplement to the “Journal of Botany,” January to July
1912. Pp. 1-76, with one Plate.

Mr. Pucstey here distributes the British plants of this genus into
eight species, two sub-species, and fourteen varieties with sub-varieties
and one hybrid.

The number of pages here used to describe and treat on these
will at once show how full the work is; the only fear one has 1s,
will it be the fate of all British genera? If so, one can but pity the
student when wading through the series. Even the artificial key
on pages 72 and 73 makes one wonder if any person will ever try to
name a specimen by it. Certainly, if all this has to be learnt, the
oft-repeated remark as to botany, “all hard names,” will be
exemplified.

Still, the trend of all botany is to specialise, and it is hard to find
fault with good honest work simply because it is long in telling.

Mr. Pugsley has evidently studied the genus very closely and with

240 THE SCOTTISH BOTANICAL REVIEW

much care, and if means could be adopted to shorten the matter for-
a flora, it would represent good and painstaking work.

In addition to the former new species (occidentalis and purpurea),}
we have now a third described (7: pardoxa), two sub-species, one
hybrid (x / Painter’), and several varieties, etc.

Under / paradoxa Mr. Pugsley certainly seems to have scored,
as four Continental botanists referred this to quite another section of
the genus, and I think in a case like this we may be forgiven for
being a little proud of our countryman.

Lastly, the index of 121 names shows how much is included in
the work.

PLant LiFE AND EVOLUTION. Professor D. H. CAMPBELL,
“American Nature Series,” Holt & Co., New York, 1911.
Price $1.60 net.

Mucu has recently been written upon the subject of evolution
following upon the celebration, both here and in America, of the
jubilee of the publication of Darwin’s “Origin of Species.” In
these discussions plants have not played so prominent a part as
animals, hence we may welcome this book all the more, coming as it
does from one of the American school, reared up in the traditions
of Agassiz, Hyatt, Ribot, Cope, Marsh, Beecher, Osborn, and many
other valiant supporters of Neolamarckism. In Europe we can
point to Hering, Herbert Spencer, Pfeffer, Semper, F. Darwin, and
Semon as able exponents of similar views. Moreover, Professor
Campbell has done pioneer work on the development and structure
of mosses and ferns, apart from other important studies. He may
be said, in fact, to occupy that place in the New World which in the
Old World we accord to Professor Bower, as doyen of cryptogamic
morphology. Both have reached much the same conclusions (vide
the “Origin of a Land Flora,” “ Plant Life on Land”), if by some-
what different methods, which is perhaps the best criterion of
accuracy.

The book, which may be thoroughly recommended to English
students, is admittedly written from a Neolamarckian standpoint, but
whilst the author discusses in a masterly way all the theories of
evolution, he accepts none as explaining the ultimate cause of evolu-
tion. Though primarily written for American students, it is safe to
say it is of international interest, since the treatment of the subjects,
discussed by one who is a specialist in his own particular field,
displays so wide and intimate a knowledge of current work and
thought in other fields, with so well balanced and critical an estimate
of the value and relative bearing or otherwise of each upon the
matter in hand, that the book may be placed on the shelf as an
important study in origins, especially applicable to the plant world.
Whilst it provides all the material for a text-book of morphology or

‘**Jour, of Botany,” 1902, p. 129; 1904, p. 217.

REVIEWS, BOOK NOTICES, ETC. 241

physiology, general ecology and etiology, it is written from a genetic
standpoint, so that the meaning of structures or phenomena discussed
is at once apparent. Moreover, not only are the latest discoveries
in botanical work, recent or fossil botany, referred to, but many of
the more important zoological facts are contrasted with them. It will
illustrate the scope of the work best perhaps to refer to a few of these.
In connection with the origin of life, Winogradsky’s researches on
nitrogen bacteria are mentioned. In discussing the lower plants,
Bower’s theory of sterilisation finds a place, and the antithetic
alternation of generations of pteridophytes and bryophytes is
affirmed.

In relation to heredity, F. Darwin’s views on memory are accepted,
and, we think, with good reason. Bower’s theory of sterilisation is
however, not accepted, an entirely separate origin for the sporophyte
in club-mosses and ferns being assumed. Researches amongst the
carboniferous and mesozoic floras as to the origin of seed plants by
Scott, Seward, and Wieland are given due prominence. By the way,
Seward’s view that conifers such as Avaucaria are related to the
Paleeozoic lycopods is favoured. The cumulative effects of stimuli
are illustrated by Jenning’s work on Infusoria. In determining the
value of current theories of the origin of species, the mutation theory
of De Vries and Mendel’s laws of heredity are fully discussed, but
Lotsy’s views as to the cause of mutations are preferred, the germ-
plasm theory, in so far as plants are concerned (since they are more
plastic and respond to external stimuli more readily), not being con-
sidered to apply. The nature of the Palaeozoic climate is explained
on Manson’s view that a mantle of cloud excluded sunlight. Great
stress is laid upon Kleb’s experiments as to the effect of stimuli in
determining sex and regulating other factors of the environment.
All the important results of plant breeders bearing on the questions
discussed are surveyed, and Bailey and Burbank are cited here.
Winckler’s work on graft hybrids also receives adequate notice.

The book is divided into ten chapters, including an introduction,
the factors in evolution; the lower plants ; the origin of land plants
(which are derived from green algz); seed plants, leading on from
heterospory in pteridophytes from the zygote or resting stage in
algee required by dry conditions ; the Angiosperms, where the division
into monocotyledons and dicotyledons is regarded as artificial, and
their characters considered as adaptive, developed in parallel series ;
environment and adaptation, the terrestrial phase being the cause of
seed plants, and their possession of fruit is regarded as the cause of
their dominance. The chapter on plant distribution is especially
interesting and will appeal particularly to American students. The
human factor in plant evolution is dealt with from an original out-
look, and the great importance of extinction is adequately recognised.

The origin of species is lucidly expounded, and it is justly re-
marked that plants differ from animals in this respect, for while the
germ-plasm may play a part in the case of the latter, in the case of
plants, “to assume that there is a special germ-plasm, which is passed

VOL. I. 18

2A THE SCOTTISH BOTANICAL REVIEW

on from the tiny gametophyte to the non-sexual and long-lived
sporophyte, and finally segregated in the spores, and again passed
along to the next generation of gametophytes, is, to say the least,
improbable.” Here indeed lies the secret of the whole matter.

We miss any reference to sphagnales in the discussion of the
origin of the lower plants. The term prothallus is neglected, and
pollen spore is used for pollen grain. Owing to subdivision of the
matter there is an unavoidable amount of repetition. Generally
speaking, there is an absence of Americanisms, but we notice mold,
meter, center, color, gray for grey, skepticism, fiber, canyon, favorite,
meager, behavior, armor. The dieresis is used in re-established
pre-eminently, zoospore. Sperms is substituted for spermatozoa.
A few typographical errors occur: ax (p. 290), especialy (p. 229),
futher (p. 335), unequaled (p. 147). The terms horsetail and clubmoss
in reference to Palzozoic types are used literally; “forbears” is
used for ‘‘ancestors” in reference to plants.

Each section leads up to the next admirably. The use of common
types to illustrate facts, e.g. cross pollination in the nasturtium and
scarlet geranium, is an excellent feature. The personal element is
not entirely lacking. Visits to South Africa, where sunbirds were
studied, and Krakatau, in 1906, are related.

We believe that this book if, as it is hoped, it is read widely in
this country, will do much to stimulate inquiry upon new and right
lines, and can thoroughly recommend its adoption as a standard
text-book on plant life and problems of evoJution.

BriTisH PLant Gatis. By E. W. Swanton. Methuen & Co.,,
London. §8vo, cloth, pp. 251, sixteen coloured and sixteen
photographs, with numerous blocks in the text. 7s. 6d. net.

Tuis book should supply a long-felt want. Up to the present time
we have had only one good work dealing with British galls, and that
was mainly a translation by C. R. Straton of Dr. H. Adler’s German
work ‘Uber den Generationswechsel der Eichengallen,” and was
limited to the galls occurring on the oak. E. T. Connold subse-
quently published some fine photographs of our British galls, but
the accompanying text was of little assistance. The present work
condenses into a readily accessible form a mass of information on
this subject which for the most part is hidden away in various
systematic works and scattered papers published by numerous
scientific societies. Mr Swanton in his opening pages deals in a
lucid manner with the origin of our British plant galls, and devotes
a separate chapter to those that are caused by the Hymenoptera,
Coleoptera, Lepidoptera, Diptera, Homoptera, Acari, Nematoda,
Fungi, and Mycetozoa. In the detailed catalogue of British plant
galls which succeeds the opening chapters the galls are arranged in
botanical sequence on their hosts in accordance with the classifica-
tion adopted by Engler in “ Die Naturalichen Pflanzenfamilien,” and

REVIEWS, BOOK NOTICES, ETC. 243

the specific name accepted by ‘“‘The London Catalogue of British
Plants,” roth edition. Eight hundred and eighty British plant galls
are enumerated in this catalogue, but we fail to see that this number
is justified, as we presume that it includes the same gall when present
in the stem, leaf, or flower as a distinct gall, and it certainly includes
the same fungus when growing on different hosts, whilst the hard
woody excrescences so common on our ash trees caused by the
fungus Dadldinia concentrica are omitted from the work. The book
is well got up, and the numerous coloured illustrations and photo-
graphs should make it a useful guide to the identification of our
more common plant galls.

Index

Acacia, New or imperfectly described
Species from Western Australia, 96.

Agathosma trichocarpa, n. sp., 162.

Aglaonema, Embryo-Sac of, 100.

Alchemilla acutidens, Scottish records,
183.

Alien Plants, 39.

Alisma Plantago, vars. and forms of, 21.

Alisma ranunculoides, vars. and forms
of 2ie

Ambrosiacez, The Inflorescences of,

193.

Anthelia: an Arctic- Alpine Plant Asso-
ciation, 81.

Apium tnundatum, 19.

Applied Botany, Some Modern Aspects
of, 24.

Aquatic Forms and Species of the
British Flora, 17.

Arcangeli, Prof. John, Vectorta regza,

IIS.

Barbula gracilis in Scotland, 46.

Barbula incavata, sp. n., 93.

Barclay, W., Cvzcus oleraceus, Juncus
tenuts, 235.

Bennett, Arthur, Aquatic Forms and
Species, 17; Carex helvola, 41;
Potamogeton prelongus, 47 ; Juncus
alpinus, 47 ; Sparganium, 94; Addi-
tions to Caithness Flora, 181; Saxz-
fraga Flirculus, 205; S. atzordes,
235; Utricularia vulgaris, 235.

Blackwood, G. G., Cormus suecica, 117.

Borthwick, Dr. A. W., Modern Aspects
of Applied Botany, 24.

‘* Botanical Exchange Club Report,”
IQII (vev.), 51.

“British Plants: their Biology and
Ecology ” (vev.), 125.

‘* British Vegetation, Types of” (vev.),
49; ‘‘ Roses, List of British” (vez. ),

4.
Bryum elegantulum, sp. n., 92.

Caithness Lichens, 146 ; Recent Addi-
tions to Flora of, 181.

Calluna-Mat Association, 183.
Campbell, Prof. Douglas H., Embryo-
Sac of Aglaonema, 100.
Carex helvola, 41; canescens var. fallax,
41; Lachenaliz, 41 ; rariflora, 41.
Centunculus minimus in Wigtown-
shire, 46.

Ceratophyllum demersum, 19.

**Clare Island Survey, Phanerogamia
and Pteridophyta” (vev.), 56.

Cnicus oleraceus in Perthshire, 235.

Cornus suecica in Peeblesshire, 117,
184.

Corstorphine, M., Barbula gracilis, 46.

Corstorphine, R. H., Mzerochloe odor-
ata, 183.

Crampton, C. B., Stable and Migratory
Plant Associations, 1, 57, 127;
Calluna-Mat Association, 183.

Damasonium Alisma and forms, 23.
Druce, G. C., Linnezus’ ‘‘ Flora
Anglica,” 154.

Ecological Terminology as applied to
Marine Algze, 44.

Epilobium nummularifolium in Argyll-
shire, 47.

Evans, Wm,, Zostera nana,
Cornus suecica, 184.

IME

Fissidens incurvus from Skye, 48.

“* Flora Anglica,” 154.

Fraser, James, Alien Plants, 39;
Centunculus minimus, 46.

‘¢Galls, British Plant” (vev.), 242.
Gilbert, E. G., Study of Rubi, 184.
Goebel, K. von, Ambrosiacez, 193.
Grateloupia, A new Japanese, 208.
Grimmia rubescens, Sp. T., 9O.
Grimmia undulata, sp. N., 91.

Hierochloe odorata in Glen Calla, 183.
Holmes, E. M., Agathosma trichocarpa,
162 ; Grateloupia subpectinata, 208.

INDEX

245

Horwood, A. R., Past History of | Ranunculus Flammu/lavar. natans, 18 ;

Monocotyledons, 164, 216.
Hydrocotyle vulgaris, 18.

Johnson, N. Miller, Ecological Ter-
minology applied to Marine Algz,
44; Petasites albus, 118.

Juncus alpinus, 47.

Juncus bulbosus, 20.

Juncus supinus, 19; var. nodosus, 19 ;
var. pygmaeus, 20; var. Kochit, 20;
var, subverticellatus, 20; var. fluztans,
20 ; var. comosus, 20.

Juncus tenuts in Peeblesshire, 47 ; in
Glen Ogle, 235.

Kenfig Burrows, An Ecological Study,
209.

Leucobryum pumilum in Britain, 48,

9.
Lichens, Caithness, 146.
Lillie, Rev. D., Caithness Lichens, 146.
Linneeus’ ‘‘ Flora Anglica,” 154.
Lothians, Mosses of the, 202.

M‘Andrew, James, Sz/ene jimbriata,
Piptatherum multiflorum, Eprlobtum
nummularifolium, Juncus tenuis,
47; Mosses of the Three Lothians,
202.

Macgregor, M., Calluna-Mat Associa-
tion, 184.

Marine Algze, Ecological Terminology
as applied to. 44.

Meldrum, R. H., Philonotis rigzda,
117.

Monocotyledons, Past History of, 164,

216.
Morphological Notes: Ambrosiacez,

Oo
Morrison, Alexander, Species of Acacia
from Western Australia, 96.
Mosses from the West Highlands, 89.
Mosses from the Three Lothians, 202.

Notes from Current Literature, 48, 118,
185, 236.

Orr, M. Y., Kenfig Burrows, 209.

Peplis Portula, 18.

Petasites alous in Fife, 118.

Philonotis rigida in Perthshire, 117.
Piptatherum multifiorum in Britain,

47.

Plant Associations, Geological Relations
of Stable and Migratory, I, 57, 127;
Calluna-Mat, 183; Anthelia, 81.

Potamogeton prelongus in Orkney, 47.

Pyrenocheta [licts, n. sp., 161.

lingua, 18.
Reviews, etc., 49, 121, 186, 237.
Types of British Vegetation, 49.
Botanical Exchange Club Report,

eT
List of British Roses, 54.
Plant Life on Land, 56.
Clare Island Survey: X. Phanero-
gamia and Pteridophyta, 56.
Watson Botanical Club Report,
IQIO-II, 121.
British Plants, their Biology and
Ecology, 125.
Prodromus Flore Britannice, 186.
The Shingle Beach as a Plant
Habitat, 188.
Monograph of the British Desmidi-
acez, 190.
Clare Island Survey: XIV. Lichens,
190.
Flora of Cambridgeshire, 191.
Clare Island Survey: XI.-XII.
Musci and Hepatice, 237;
XIII. Fungi, 238.
Genus Fumaria in Britain, 239.
Plant Life and Evolution, 240.
British Plant Galls, 242.
“Roses, List of British ” (vev.), 54.
Rubi, Study of, 184.

Sagittaria sagittifolia, 22; var. vallis-
nertfolia, 22; heterophylla, 23.
Saxtfraga aizotdes in Orkney, 2353
Hitrcudus in Caithness and its Dis-
tribution in the British Isles, 205.
Scerpus fluztans, 20.
Short Notes :—
Barbula gractlis in Scotland, 46.
Centunculus minimus in Wigtown-
shire, 46.
Silene fimbriata in Argyllshire, 47.
Piptatherum multifiorunin Britain,

47-

Epilobium nummularifolium im
Argyllshire, 47.

Juncus tenuzs in Peeblesshire, 47.

Potamogeton prelongus in Orkney,
47-

Juncus alpinus in Kirkcudbright,
AT:

Fissidens incurvus in Skye, 48.

Leucobryum pumilum in Britain,

Philonotis rigida in Perthshire,
LE

Cornus suectca in Peeblesshire, 117.

Zostera nana in Haddingtonshire,
DE Z-

Petasites albus in Fife, 118.

246

Short Notes—continued.

Hierochloe odorata in Glen Calla,
183.

Alchemilla acutidens in Banff and
Linlithgow, 183.

Calluna-Mat Association, 183.

Rubi, Study of, 184.

Cornus suecica, 184.

Saxifraga atzoides in Orkney, 235.

Cnicus oleraceus in Perthshire,
235-

Juncus tenuis, 235.

Otricularia vulgaris in Caithness,

235.
Silene fimbriata in Argyllshire, 47.
Smith, Dr. W. G., Anthelia Associa-
tion, 81.

THE SCOTTISH BOTANICAL REVIEW

Sparganium minimum, 23; hyper-
boreum, 23; glomeratum, 23; Scot-
tish forms of, 94.

Stirton, James, /zsszdens incurvus,
Leucobryum pumilum, 48; Mosses
from the West Highlands, 89.

Utricularia vulgaris in Caithness, 235.

| Victoria regia, Note on, 115.

‘* Watson Exchange Club Report,”
IQIO-II (rvev.), 121.

Wilson, Malcolm, Pyrenocheta Llicts,
sp. n., 161.

Zostera nana in Haddingtonshire, 117.

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The
Scottish Botanical Review

The Scottish
Botanical Review

A QUARTERLY MAGAZINE

Including the
Transactions of the Botanical Society of Edinburgh

Volk L192

EDINBURGH
NE ELE, < CO. LIMERED

BELLEVUE

Contents

First Part—January 1912

THE GEOLOGICAL RELATIONS OF STABLE AND MIGRATORY PLANT
FORMATIONS. C. B. Crampton, M.B.,C.M. (70 be continued)

REMARKS ON SOME AQUATIC FORMS AND AQUATIC SPECIES OF
THE BRITISH FLORA. Arthur Bennett, A.L.S. .

SOME MODERN ASPECTS OF APPLIED BOTANY. A. W. Borthwick,
IDES:

ALIEN PLANTS. James Fraser .
CAREX HELVOLA, BLYTT. Arthur Bennett, A.L.S.

ECOLOGICAL TERMINOLOGY AS APPLIED TO MARINE ALG2&.
N. Miller Johnson, B.Sc.

SHORT NOTES :—
Barbula gracilis as New to Scotland. Margaret
Corstorphine
Centunculus minimus, L., in Wigtownshire. James Fraser

Silene fimbriata, Sims, in Argyllshire. Piptatherum multi-
florum, Beauv., in Britain. Lpzlobiwm nummularifolium,
R. Cunn., in Argyllshire. /cus tenuis, Willd., in Peebles-
shire. James M‘Andrew -

Potamogeton prelongus, Wulf., in Orkney. Arthur Bennett
Juncus alpinus, Vill., in Kirkcudbright. Arthur Bennett .
Fissidens incurvus, Starke, from Skye

Leucobryum pumilum (Michx.) in Britain. James Stirton

NOTES FROM CURRENT LITERATURE

REVIEWS, BooK NOTICES, ETC. :—

The Types of British Vegetation. Edited by A. G. Tansley

The Botanical Exchange Club of the British Isles. Report for
Ig1o. C.E. Moss : : ‘ . ;

PAGE

46
46

49

51

vi THE SCOTTISH BOTANICAL REVIEW

A List of British Roses (Supplement to Journal of Botany,
1911). Major A. H. Wolley-Dod

Plant Life on Land, considered in some of its a
Professor P. O. Bower .

Clare Island Survey. Part X.: paaeeeee and Preidopby
By R. Lloyd Praeger

Second Part—April 1912
THE GEOLOGICAL RELATIONS OF STABLE AND MIGRATORY PLANT
FORMATIONS, C. B. Crampton, M.B., C.M. (Zo be continued)

ANTHELIA: AN ARCTIC-ALPINE PLANT ASSOCIATION. W. G.
Smith, B:Se., Ph.D.

MOSSES FROM THE WESTERN HIGHLANDS. James Stirton, M.D.,
F.LS.

SCOTTISH FORMS OF SPARGANIUM. Arthur Bennett, A.L.S. .

NEW OR IMPERFECTLY DESCRIBED SPECIES OF ACACIA FROM
WESTERN AUSTRALIA. Alex. Morrison

THE EMBRYO-SAC OF AGLAONEMA. Douglas Houghton Campbell

NOTE ON VICTORIA REGIA, LINDL. John Arcangeli

SHORT NOTES :—
Philonotis rigida Brid., in Perthshire. R. H. Meldrum
Cornus suecica, Linn., in Peeblesshire. G. G. Blackwood
Zostera nana, Roth., in Haddingtonshire. William Evans
Petasttes albus, Gaertn., in Fife. N. Miller Johnson.

NOTES FROM CURRENT LITERATURE

REVIEWS, BOOK NOTICES, ETC. :—

The Twenty-Seventh Annual Report of the Watson Botanical
Exchange Club, 1910-11

British Plants: their Biology and Ecology. F. Bevis and
H. J. jeftery ..

Third Part—July 1912
THE GEOLOGICAL RELATIONS OF STABLE AND MIGRATORY PLANT
FORMATIONS. C. B. Crampton, M.B.,C.M. (Concluded)
CAITHNESS LICHENS. Rev. David Lillie, B.D.
LINNAZUS’ “ FLORA ANGLICA.” G,. Claridge Druce, M.A., F.L.S.

PAGE
54
56

56

57

81

89
94

96
100

115

117
117
117
118

118

I2I

. 126

127
146
154

a E—_—_ -_—~—~

CONTENTS

A NEW SPECIES OF PYRENOCHATA.
BRS:

Malcolm Wilson, D.Sc.,

AGATHOSMA TRICHOCARPA, N. SP. E. M. Holmes, F.L.S., F.B.S.E.

THE PAST HISTORY OF MONOCOTYLEDONS, WITH REMARKS ON

THEIR ORIGIN. A. R. Horwood.

RECENT ADDITIONS TO THE CAITHNESS FLORA.

A.L.S. .

SHORT NOTES:

(To be continued)

Hierochloe odorata, Wahl. R. H. Corstorphine
Alchemilla acutidens, Buser. M‘Taggart Cowan, Jr.

Note on the Calluna-Mat Association of the Mountain Tops of
the Northern Highlands. C. B. Crampton and M. Macgregor

British Rubi and environment, etc.

Cornus suectca, Linn. Wm. Evans
NOTES FROM CURRENT LITERATURE
REVIEWS, BOOK NOTICES, ETC. :—

Prodromus Flore Britannica. F.

Phytologist,” No. 3, vol. xi.)

A Monograph of the British Desmidiacez. W. ere G. S. West

E. G. Gilbert

Arthur Bennett,

N. Williams. Part IX.
The Shingle Beach as a Plant Habitat. F. W. Oliver.

. New

Clare Island Survey. Part XIV.: Lichens. Miss Annie

Lorrain Smith :
A Short Flora of Cambridgeshire.

A. H. Evans

Fourth Part—October 1912

MORPHOLOGICAL NOTES. I.: The Inflorescences of the Ambrosi-
acez. K. von Goebel, Sc.D., LL.D.

NOTES ON SOME MOSSES FROM THE
M‘Andrew

THREE LOTHIANS.

James

SAXIFRAGA HIRCULUS, L., IN CAITHNESS, AND ITS DISTRIBUTION
IN THE BRITISH ISLES. Arthur Bennett, A.L.S.

A NEw JAPANESE GRATELOUPIA. E. M. Holmes, F.L.S., F.B.S.E.

KENFIG BURROWS: AN ECOLOGICAL

STUDYe. ) Mesye. Orr

THE Past HISTORY OF MONOCOTYLEDONS, WITH REMARKS ON

THEIR ORIGIN. A. R. Horwood.

(Concluded)

186

188
190

190
19I

202

Vili THE SCOTTISH BOTANICAL REVIEW

SHORT NOTES :—

Saxifraga aizotdes, L.,in Orkney. Arthur Bennett .
Utricularia vulgaris L., in Caithness. Arthur Bennett

Cnicus oleraceus Lin., Perthshire, and Juncus tenuzs in Glen

Ogle. W. Barclay

NOTES FROM CURRENT LITERATURE

REVIEWS, BOOK NOTICES, ETC. :—

Clare Island Survey. Parts XI. and XII. : Musci and Hepatice.

Canon H. W. Lett., M.A.

Clare Island Survey. Part XIII.: Fungi. Carleton Rea

The Genus Fumaria in Britain. H. W. Pugsley
Plant Life and Evolution. D. H. Campbell

British Plant Galls.

E. W. Swanton .

PAGE

235
235

235
236

237
238
239
240
242

ig

Contents

PAGE
MORPHOLOGICAL Notes. I.: THE INFLORESCENCES OF THE

AMBROSIACEH. K. von Goebel, Sc.D., LL.D. : 193

NOTES ON SOME MOSSES FROM THE THREE LOTHIANS.
James M‘Andrew : 4 : : : 2e2

SAXIFRAGA Hircu.us, L., in CAITHNESS, AND ITS DIsTRI-
BUTION IN THE BriTisH IsLes. Arthur Bennett, A.L.S. 205

A NEw JAPANESE GRATELOUPIA. E. M. Holmes, F.L.S.,
FB».E. . A , : : . 208
KeEnFIG Burrows: AN EcoLocicaL Stupy. M.Y.Orr . 209
THE Past History oF MONOCOTYLEDONS, WITH REMARKS
ON THEIR ORIGIN. A. R. Horwood. (Concluded) ane ¥6

SHortT Notes :—

Saxtifraga aizoides, L., in Orkney. Arthur Bennett ieee

Utricularia vulgaris L., in Caithness. Arthur Bennett. 235
Cnicus oleraceus Lin., in Perthshire, and Juncus tenuis in

Glen Ogle. W. Barclay. é aes Fi

-Nores From CurRENT LITERATURE : ; tek.

Reviews, Book NOTICES, ETC. :—

Clare Island Survey. Parts XI. and XII.: Musci and
Hepatic. Canon H. W. Lett, M.A. ; ba BF
Clare Island Survey. Part XIII.: Fungi. Carleton Rea 238
The Genus Fumaria in Britain. H. W. Pugsley reso
Plant Life and Evolution, D.H.Campbell . A aS
British Plant Galls. E.W.Swanton . ; ee: eee

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