26 ISTEODDCTORY ESSAY.
exceptions to this rule see the precodiog section under the heading " F e m a le
S p i k e l e t s . "
Male Flowers.—flowers of both soxes in Calamus are small and unobtrusive.
T h e male flowers are most frequently larger aad more elongated than the female
ones, but even the largest, as in 0. erecius, C. longisetus, 0. arhrescens, C. Flaijellum,
0. Thwaitesii, etc., rarely exceed I era. iu length; they are green or yellowish and
are always of a coriaceous or pergamentaceous texture.
T h e calyx of the male flower is shortly tubular, cupular or urceolate, and is
always more or less deeply 3-toothed or 3-lobed (PLATE I, fig. 4 ) . The corolla is
always considerably longer than the calyx, and is divided almost to the base into
3 narrow segments ( PLATE I, figs. 4 and 5 ) . The stamens are 6 in number and
most frequently are uniseriate, and have subulate filaments with a more or less
distinctly inflexed apex. In C. pachystemoms and allied species the stamens ai-e
d i s t i n c t ly 2-seriate, three of them being, moreover, shorter than the other three, while
t h e i r filaments are not inflexed at the tip, so that the anthers in bud as well as
duriog floweriQg are erect and basifixed. When the filaments are inflexed the
anthers are versatile and deeply divided at the base (PLATE I, fig. 17, and PLATE II,
fig. 5 ) . la all the male flowers that I have examined I have always found a
rudimentary ovary composed of 3 small elongated bodies or rudimentary carpels
( PLATE I, fig. 6n, and PLATE II, fig. on). I bave never met with the faintest sign of
a nectary iu the interior of a male flower.
Female Flowers.—The female flowers are usually shorter and stouter tlian the
male ones and are ovate or conic iu shape; their calyx is cupular—urceolate or
subcampanulate and always more or less deeply 3-toothed or 34obed ( Pr,ATE I I , figs.
6 - 1 1 ). The corolla is always more deeply partite than the calyx, but its divisions
v e r y seldom exceed the teeth of the calyx ; in a very few cases, as for instance
in C. jamisis, the corolla of the female flower is conspicuously longer, and in
C. adsperitis is slightly shorter than the calyx.
T l i e stamens of the female flower are six in number and are always sterile;
t h e bases of their filaments aro sliglitly connate to form a cup or urceole which
envelopes the ovary and is crowned by 6 more or less elongated teeth ; these teeth
bear sagittate basifixed but abortive anthers (PL4TE I I , fig. 11).
T h e ovary is globular, ovate-conic or even turbinate, and does not differ
essentially from that of the other Lepidocaryece, of which the chief characteristic is
a " l o r i c a " formed of imbricated scales (PLATE II, fig. I I ) . The ovary is
trilocular, with the 3 cells separated by very thin dissepiments; each cell has a
solitary ovule which is anatropous. basilar, and inserted on the inner angle of the cell.
Host usually only one of the ovules grows to maturity, and of the others only the
remains may be traced in fertilised ovaries. Perhaps the dissepiments of the cells
are frequently incomplete from the first formation of the ovary and the three ovules
stand erect in the centre of the ovary ; this at all events is what has seemed to me to
be the case in dry specimens of C. ZolUngerii.
T h e style is usually very short and comparatively stout; the stigmas are 3
i n number and are usually rather stout, elongate-trigonous, acuminate or subulate,
THE FLOWEES. 27
lamellose inside, spreading and rccurved when the flower opens (PLATE II, figs.
6 , 7 , 8 , 10 and 11), and are usually persistent even when the fruit is mature.
Neuter Flotocrs.—In the female spadix the fertile flower is accompanied by a
neuter one (PLATE I, fig. 1 0 / , and PLATE II, figs. 6, 7 / ) which is frequently very
small and very soon deciduous, but not rarely is similar to a perfect male flower,
though thinner and with attophied anthers and abortive ovary ; this is formed—as in
the male flowers—of three very small bodies, the representatives of three carpels
(PLATE I I , fig. 12«). The neuter flower is always inserted on the outer side of the
involucre in the centre of the special area or niche which I have tenncd the
areola of the neuter flower.
T h e neuter flower is almost always sessile, but I have found that it is provided
with a distinct stalk in C. JJenruanus and C. adspersus. In some species, as
C. salidfolius and C. Rotang, the neuter flowers do not differ in appearance from the
f e r t i l e male flowers, but I do not know if the pollen of their anthers be perfect and
capable of fertilizing. In C. temiis, C. Guruha and again in C. Delessnrtianm and
C. RidUyantfs the neuter flowers are also well developed, and are only slightly smaller
than the female ones; as they are only deciduous when the female flowers are on
Ihe point of expanding, the female spikelets immediately prior to the opening
of the flowers have these arranged in four very distinct and almost similar series.
F r om the manner in which the neuter flower is inserted outside the floral
involucre and on account of the small axial part with which in a f ew cases it is
furnished, we can readily recognise that, along with the con-esponding female flower,
it forms part of a very small secondary contracted spikelet.
T h e neuter flower may therefore be considered to be a male flower which is
rendered functionless by a retrograde process and becomes depauperated owing to
the greater development of the fertile one or perhaps it may, from another point of
view, be considered one that has never attained the full structure fitting it for reproductive
functions,
I t happens sometimes in nature that certain structures, which have made their
appearance at an early date (in the evolutionary sense) under the stimulus of a definite
need of the organism, and have been capablu of fulfilling a definite function, later on,
under altered conditions of existence, have become useless and have consequently
been modified and reduced by atrophy, but still prsiest, possibly because the disuse
began to bo experienced when the malleability of the organism was already greatly
diminished. All this is in accordance with my theory of variability restrained by
the force of heredity in the plasmatical era.*
T h e s e neuter flowers therefore appear to me to be a striking proof of the
presence in an organism of useless structures that do not now exercise any function,
or that perhaps never even in bygone times exercised any function.
T h e neuter flowers of Calami usually never open; they have stamens and an
ovary that, according to the species, are more or less atrophied, and that are
deciduous before the fall of the female flowers. There seem however to be a few
exceptions to this rule, as for example in C. GriJJithianns (PLATE I I . fig. 12) where
the ealyx and corolla apparently expand.
• Beccari: Nelle Foreste di Uorneo loc, eit.
Akn. EOY. BOX. GARD.. CALotxtA VOL. X I .