42 IV. C.

Worsdell—
THE ORIGIN OF THE PERIANTH OF FLOWERS
WITH SPECIAL REFERENCE TO THE RANUNCULACEAJ.
[PLATE III.]

ASPAR FRIEDRICH WOLFF' was the first to hint at the

C origin of the flower as being the outcome of a gradual meta-
morphosis of the foliage-leaves situated at a lower level on the axis
of the plant. He held that the production of floral leaves was the
result of a degeneration in the quality of the sap, the richer portion
being used up at a lower level by the first-formed vigorous foliage-
leaves. Goethe-, however, first established the theory of the under-
lying homology of all foliar organs of the plant. His main idea
was that the fioral leaves represented the final stage in a gradual
ascending metamorphosis of which the foliage-leaves are the
starting-point, and that the production of flowers is due to an
improvement and etherealisation of the sap during its ascent through
the plant. All foliar organs are variants on an ideal type-leaf.
These two great thinkers dealt solely with the individual develop-
ment of the plant; they, like so many biologists since their time,
failed to see that this could throw no real light upon the ultimate
phylogenetic development of the parts concerned.
GoebeP states that the type of which all foliar organs are the
modifications is the foliage-leaf, a view sufficiently idealistic and
resembling Goethe's " type-theory."
We hold, for cogent reasons* which it is beyond the scope of
this article to enlarge upon, that fertile foliar organs (or sporophylls)
preceded in time all other kinds of leaves, and that the latter have
been gradually differentiated from the former by a process of
sterilisation of their tissues.
' Wolff, C. F. Theoria Geuerationis, 1759.
* Versuch die Metamorphose der Pflaiizen zu erkliiren, Gotha,
1790 (" Essay on the Metamorphosis of Plants; " translated
by Emily M. Cox in Seeniann's Journal of Hotany, Vol. I.,
1863).
» " Die vergleichende Entwickelnngsgeschichte der Pflan-
genorgane;" Schenk's Handbuch der Botanik, Vol. III.,
part I, p. 100, 1884.
* See Cclakovsky : " Ueber die allgeineine Entwickelungsges-
chichte des Pflanzenreiches;" Kgl. bohni. Gesellsch.
der WissenschaftPU ; Naturwiss.-Mathem. .Section, i868.
" Ueber den phylogenetischen Entwickelungsgang der liliithe,
etc., part i, p. 7, loc. cit., Piag, 1896.
" Ueber den phylogenetischen Eutwickelungsgang der liliithe,
etc., part 2, p. 162, Prag, 1900.
" Nachtrag zu meiner Schrift iiber die Gynniospernien;"
Englers Bot. Jahrbiich, etc., Vol. XXIV., part 2, 1897.
Epilog zu meiner Schrift iiber die Placenten der Angiosperuieu j '
(appendix) Sitzuugsb. d. Kgl. bohm. Ges. d. Wiss., 1899.
 The Origin of tke Perianth of Flowers 43
Now we must look for the nearest origin of our modern flower
to the cone of the Cycadaceae, which is, in fact, the primeval flower>
consisting of a much-elongated axis clothed with very numerous
spirally-arranged sporophylls. But the chief point of interest lies
in the fact that in all cases a few of the lowermost sporophylls are
perfectly sterile and somewhat altered in shape, although not in
texture or colour, from the fertile ones. They may, in fact, be
regarded as the progenitors of the perianth-leaves of the Angio-
sperms. And it is obvious that they have been derived by simple
sterilisation of the loweriitost sporophylls of the cone. (Hig. 1).
We may, in tracing from this point onwards, the evolution of
the flower, confine our attention almost exclusively to the order
Ranunculacese among the higher plants, as there is a great deal of
nstruction to be gleaned from this group. Here we flnd some of
the primitive characters of the cone still present: e.g. inconstancy
and indefiniteness in the number of members of the various
categories of floral foliar organs (not yet stereotyped into the 5's or
4's or 3's of the more advanced orders); the much-elongated floral
axis, as seen especially in Myostints (Fig. 2) and also in an allied
order, the Magnoliaceae. The new, more advanced characters which
have appeared are:—the coloration of the perianth (yet exceptions to
this occur), and the frequent addition of an extra category or whorl
of perianth-leaves to that already present. Having recognised these
features we will first of all enquire into the origin of the corolla.
Nageli' and subsequently Drude- were the first to point out the
probable origin of the corolla from the outermost whorl of stamens.
Not long afterwards Grant Allen' ingeniously endeavoured to prove
that the petals of flowers are in all cases derived from stamens.
With the last-named authors he considers the calyx to have been
derived from bracts. The great German botanist PrantI* held, on
the other hand, that the corolla, like the calyx, arose from the meta-
morphosis of bracts, and that staininodial structures only (including
nectaries and small petaioid organs bearing incomplete or rudi-
mentary anthers) bad been transformed from the outermost stamens.
Having especial regard to the Ranunculaceae, he always drew a
sharp distinction between these staminodes on the one hand and the
corolla on the other. For instance, he regards the nectar-bearing
"petals" of Rantincidus as staminodes and possessing a totally
• Mecliaiiisch-physiologisclie Theorie der Abstauimitngslebre;
p. 496 1884.
» '• Die sj'stematisclie and geograpbisrhe Anordiiung der
Pliaiierogaiueii;" Scheuk's Haiidbucb, \'ol 1II., part 2,1887
• The Colours of Flowers, 18S2.
•• "Beitrage zur Morphologie and Syaleinatik der Kauuiicu*
lacese; " Bugler's Bot. Jahrbucber, Vol. IX., 1S87.
44 ^^- C. Worsdell—
different origin from true petals. He ascribes to Actaea, along
with Ciinicifuga, the luiiversal possession of staminodes. Celakovsky
from whose works all that this article contains is gleaned, is of
opinion that the sharp distinction made by Prantl cannot well be
maintained; for instance, the tiny petals of some species of Actaca
{Euactaca Prantl) are without nectaries, while others {Ciinicifuga)
produce them; the same holds good for Coptis. Further, the fact
that many petals not possessing nectaries, and to which Prantl
assigned the character of staminodes, exhibit rudiments of anthers,
while the petals of Adonis do not is of no decisive importance, for
such transitional forms between stamens and petals do not occur in
all species of those genera to which staminodes are ascribed, as is
the case in our own Actaea spicnta (Fig. 3). It is, therefore, far
from clear why these latter should have been derived from stamens
and the petals of Adonis enjoy a different origin. Other important
facts, for which there is no room here, tend to render Prantl's view
improbable. And we have only to turn to any "double" flower in
the garden to observe the gradual transitions taking place under our
very eyes, between stamens and true petals to be convinced that
staminodes and petals have precisely the same origin, and that what
to-day occurs abnormally may have been phylogenetically the normal
process.
We have hitherto assumed that staminodes are of staminal
origin; the real ruison d Hre of this view lies in the fact that
nectaries usually exhibit a more or less distinct 2-lipped structure,
the upper (inner) lip being homologous with the upturned basal lobe
of a peltate anther,^ while the non-secretory staminodes very fre-
quently bear rudiments or portions of pollen-sacs.
Without entering further into the question we may conclude
with Prantl that nectaries and other staminodes are of staminal
origin; but it seems quite permissible to proceed farther and add
that in the same way as the petals of Ranunculus and Calliantheimim
have been derived from the expansion and enlargement of the outer
or lower lip of the nectaries {e.g., of Helleborus (Fig. 4)—so also
the petals of Adonis have been derived from small pollen-bearing
or sterile staminodes and, therefore, ultimately from stamens.
A very pretty piece of evidence in favour of the view just stated—
that petals are of staminal origin—can be adduced from the behaviour
of the flowers in the Cleinatis-^roup. The perianth of most species of
this genus is single and of the nature of a caly.v ; within, the
numerous stamens are spirally arranged. In Clematis integri/olia,
' This is shewn by Celakovsky' as a result of his investigations
into the origin of the perianth of Narcissus as seen in the
phenomena presented in many doubleflowersof that plant.
 On the Origin of the Perianth of Flowers. 45
according to Eichler, four of the stamens, viz., the outermost ones,
are conspicuously larger than the rest, and, moreover, alternate
regularly with the four sepals (Fig. 5). Now, if we turn to the
species C. hnlearicHS {Atriigeite calyciim) the four large stamens of
the last species are exactly replaced by four small petals (Fig. 6).
Eichler observed in cultivated specimens of Atragene aipiiia that
not only were the four large stamens of C. integrifolia replaced by
petals, but that also all the intervening stamens belonging to the
same spiral cycle as the four large ones, as well as the whole of
those of the next inner cycle, had become transformed into tiny
petals; the four alternating with the sepals considerably exceeding
in size any of the other stamens.
Now what has here in Atragene nlpina, been noticed as an
abnormal phenomenon occurs normally and typically in Adonis
vernalis where, to the 8 petals succeeding the sepals, and which are
alone present in A. atitnmnalis, is added a second inner whorl of 8
or, more frequently, a slightly larger number of petals (Fig. 7). In
Adonis aestivalis and other annual species 6, 5, or even fewer petals
are present; thus the remaining petals, which, in A. aiitumnalis make
up the 8-merous character of the corolla, occur here as stamens.
There seems, at least to our mind, little doubt in these cases as to
the origin of the petals. Other similar cases could be cited. But
what we have written is probably sufficient to demonstrate the origin
of the corolla from the andrcecium.
Let us now turn to the calyx and try to discover its region of
birth. It is a striking and beautiful fact, as illustrating the division
of labour that takes place in the economy of the flower in this group
of plants, that in the absence of a double perianth, the single one
(which, on the analogy ot the flowers of the great generality of plants,
must necessarily be regarded as a calyx) assumes the characters,
viz., the colour and texture, of the corolla in the double perianth.
Such a petaioid calyx is exhibited by Caltha, Trollius paltnattis,
Isopyrtim (sect. Eneinion,) Tratitvetteria, Cletnatis sp.. Anemone sp.
(Fig. 9), Thalictrum sp., Hydrastis. As regards the generality of its
species, Trollius is in this respect an interesting genus. Its corolla
consists of small staminodial structures; the outermost of these
have, on Celakovsky's view, been transformed at some time or other
in the past history of the plant into extra sepals, so that in T.
europaeus, T. asiatiais, etc., the petaioid calyx, unlike its 5-merous
counterpart in the white-flowered T. laxus, has become polymerous
(Fig. 8).
Now, we have to decide which is the more primitive: the
ordinary green calyx of Ranunculus and the majority of Dicotyle-
donous plants, or the petaioid calyx of the cases just mentioned.
46 W. C. Worsdell-
Celalcovsky points out the following facts as proving the latter to
he the original form. When a typically coloured corolla is present
the calyx is usually, or very often, jjreen and bract-like; when the
corolla is represented by inconspicuous nectaries or staminodes the
calyx is seen to exhibit the attractive characters typical of the
petals. This latter fact is aptly and beautifully illustrated in the
relation which a South American species of Ranunculus, R. npiifolius
Pers. (Aphanosteinwa), holds to the other species of the genus. In
most species, as in R. acris, a well-developed coloured corolla is
present, and the calyx is green ; but in the South American species
the petals are represented by extremely small, sub bilabiate stami-
nodial structures greatly resembling the nectaries of Xanthorrhiza,
while, in distinct correlation therewith, the calyx is more or less
petaloid in character. The occurrence of a similar petaloid calyx in
conjunction with a staminodial corolla is seen in many species of
Helteborus (H. viridis and a few others, with their green calyx, fur-
nishing an exception to the rule), in Nigella, Eranthis, Xauthorrhizn,
Coptis, and very strikingly in Delphinium and Aconituni, where the
(in most cases) blue-coloured calyx admirably simulates, in its
zygomorphic character and adaptability to the visits of insects, the
highly modified zygomorphic corollas of many of the more advanced
orders of plants.
Now, on the view we are considering, the staminodes are
necessarily of earlier origin than the typically coloured petals
because more nearly related to the stamens, and derived from the
latter ; hence it follows that the petaloid calyx (the almost constant
accompaniment of these staminodes) must be of earlier origin than
the green calyx of Ranunculus acris, &c., and that of most"
Dicotyledons.
Finally, we may call into view another set of phenomena which
are exhibited by the interesting genus Anemone. Our own obser-
vations shew that in some species, as/I. virginiana, A.pennsylvanica,
and A. ranunculoides there is a single perianth-whorl of five
coloured leaves, these being arranged, as in Calthoe, &c., in a two-
fifth spiral (fig. 9), the perianth is thus in this case, as in the other
genera above mentioned, obviously a calyx. In other species, as
A.hortensis, A. nemorosa, A. Halleri, there is a distinct tendency
shewn towards the arrangement of the sepals in two distinct whorls,
while at the same time one or two extra sepals have been added
above, with the result that there obtain, as in the Common Wood
Anemone, two whorls of three or four members each. A. sylvestris
affords a transitional case, where the five sepals are, as it were,
hesitating whether to be whorled or spirally arranged on the axis,
 On the Origin of the Perianth of Flowers. 47
Now consider the very striking case of A. japonica or of A.
steUntn. Here, to the original and primitive 5 mei ons calyx, a
considerable number of members have bOen added, .so that the
perianth frequently consists of as many as twenty sepals,' and it is,
moreover, clear that these have been added from the stamens, hy
metamorphosis of the latter, for, in the centre of the flower, transi-
tional forms between stamens and sepals are always present and we
receive, as it were, ocular demonstration of the method of pro-
duction of these sepals which from without inwards form a con-
tinuous spiral of similarly-constituted petaloid leaves (fig. 10). But
the most important fact has yet to be stated. In A.japouica two or
three of the outermost sepals (as is also the case in Trollius) are
always slightly dilTerentiated from the rest, owing to their having,
in whole or in part, a somewhat darker, purplish coloration,- and
occasionally they exhibit a decidedly greenish tinge. Now, in that
section of the genus known as Kmwltonia (exhibiting also a poly-
merous calyx) these two or three oitteniiost sepals are entirely green
and brnct-like. Prantl accounts for the polymerous perianth of these
forms by assuming that the single perianth of other species (which,
on his view, is of bracteal origin), first multiplied its parts and
subsequently became differentiated, as regards colour and texture,
into two distinct portions. But, as Celakovsky aptly suggests, this
multiplication could hardly have taken place ex nihilo ; on the con-
trary, the extra sepals must necessarily have been derived by
metamorphosis of the stamens; and further, if the perianth be
originally of bracteal derivation it would seem strange that single
perianths of a green bract-like consistence are of such rarity in the
order. But, as regards tbis, and all other questions bearing on the
8ubject of this article, we leave our readers to judge of the respec-
tive value of the theories of the two authors whose names have been
chiefly introduced. Yet after all the Tmpressive facts above detailed
it seems to our mind fairly clear that in the great order Ranun-
culaceae, calyx and corolla have sprung from the self-same source, viz.
the andracium ; that they constitute one and the same morphological
structure, which, following the economic needs of the plant, appears
now as green protective leaves, now as brightly-coloured, delicately-
textured organs for the allurement of insects which are the destined
agents in the fertilization of the flower.
And if this be the history of the evolution of the perianth in
the group of plants we have been considering, it is surely but
' The same pheiioiiienon exists normally in the genus Clematis,
in such species as C.florida and C. patens where the calj-x
may consist of as many as ten sepals.
' In the polymerous calyx of/I. nVn/aru we have noticed that
the five outermost sepals are of a dark purple colour below.
 On the Origin of the Perianth of Flowers.
rational to conclude that the same story will be true of all other groups
of flowering-plants as well. Celakovsky, from the investigation of
certain double flowers of Narcissus, found every transition occurring
between the stamens and the members of both perianth-whorls,
showing how, in fact, the " corona " or " trumpet " represents the
upturned basal lobe of a peltate anther, each member of the perianth
possessing such an appendage. If this be true for Narcissus, it
must necessarily hold good also for the perianth of all other Monocoty-
ledons, including such a differentiated type as that of the Alismaceae
or the Commelynaceae.
Hence we see that everywhere both calyx and corolla may lay
claim to a similar place of birth, viz., in the andrcecium, however
distinct and dissimilar from each other in almost every character
they may at times appear.' W. C. WORSDELL.

STAMEN
DIAGRAM ILLUSTRATING ORIGIN OP CALYX AND COROLLA

s = sepals. p = petals. n = uectarie.s. sd = staminodes

spl = sporophyll. st.spl = sterile sporophyll.
* In Calycanthus the most gradual transitions occur from the
8tamens downward through the petals aud sepals to the
bracts.
R. MADLBY, Printer, 161, Whitfield Street, Fitzroy Square.