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 It is usual to find more than four megaspores in each
megasporangium in Palaeozoic and not infrequently, as we have
seen, in Mesozoic lycopodiaceous strobili, but in some Palaeozoic
cones, e.g. Bothrostrobus (fig. 216) and Lepidostrobus foliaceus[436],
a single tetrad only appears to have reached maturity.
The occurrence of long simple or branched and sometimes
capitate hairs is a common feature of Carboniferous megaspores
(fig. 191, E, F, I). It is possible that these appendages served to
catch the microspores, thus facilitating fertilisation. A peculiar form of
megaspore has been described by Mrs Scott[437], and assigned by
her to Lepidostrobus foliaceus, the megasporangium of which
apparently contained only four spores. As shown in fig. 191, G, a
large bladder-like appendage characterised by radiating veins is
attached to the thick spore-coat; it is suggested that this
excrescence may be compared with the “swimming” apparatus of the
recent water-fern Azolla. The epithet swimming which it is customary
to apply to the appendages of Azolla megaspores would seem to be
inappropriate if Campbell[438] is correct in stating that spores of Azolla
are incapable of floating.

B. Spencerites.
Spencerites insignis (Williamson). Fig. 192.
1878. Lepidostrobus sp., Williamson, Phil. Trans. R. Soc., p.
340, Pl. xxii.
1880. Lepidostrobus insignis, Williamson, Phil. Trans. R. Soc.,
p. 502, Pl. XV. figs. 8–12.
1889. Lepidodendron Spenceri, Williamson, Phil. Trans. R.
Soc., p. 199, Pl. vii. figs. 20–22; Pl. viii. fig. 19.
1897. Spencerites insignis, Scott, Phil. Trans. p. 83, Pls. xii–
xv.

Another type of lycopodiaceous strobilus, differing sufficiently from

Lepidostrobus to deserve a special generic designation, is that
originally described by Williamson[439], from the Lower Coal-
Measures of Yorkshire, as a type of Lepidostrobus, L. insignis, but
afterwards[440] more fully investigated and assigned to a new genus
by Scott[441]. It should be pointed out that in a later publication
Williamson spoke of the lycopodiaceous axis, which he suspected
might belong to his L. insignis, as possibly worthy of recognition as a
distinct generic type.

Fig. 192. Spencerites insignis (Williamson). (After Miss Berridge.)

Of the two species included by Scott in his genus Spencerites only
one, S. insignis, need be considered. Since the publication of Scott’s
paper our knowledge of this type has been extended by Miss
Berridge[442] and by Prof. Lang[443].
The axis of the strobilus has a stele characterised by a pith of
elongated elements, most of which have thin walls; the xylem
cylinder possesses about twenty protoxylem strands forming more or
less prominent exarch ridges. The cortex exhibits a differentiation
comparable with that in the shoots of Lepidodendron. The
sporophylls are arranged in alternating verticils, each whorl
consisting of ten members: the narrow horizontal pedicel of a
sporophyll, containing a single vascular bundle, as shown in fig. 192,
is expanded distally into a prominent upper lobe bearing a cushion of
small and delicate cells, to which the sporangium is attached, and
prolonged obliquely upwards as a free leaf-like lamina. The lower
blunt prolongation of the sporophylls appears to form a thick dorsal
lobe, but, as Lang has pointed out, it is highly probable that the
present form of the dorsal lobe is of secondary origin, and is “due to
the disappearance of a mucilage cavity from a large sporophyll
base[444].” As Miss Berridge remarks, the vascular bundle of the
sporophyll does not give off a branch to the ventral lobe and
sporangium. In attachment, in shape, and in the structure of the wall
the sporangia differ markedly from those of Lepidostrobi. The
spores, which also constitute a characteristic feature of the genus,
have a maximum diameter of 0·14 mm.; they are described as oblate
spheroids with a broad hollow wing running round the equator (fig.
192) comparable with the air-sacs of the pollen of Pinus. Scott points
out that the spores of Spencerites are intermediate in size between
the microspores of Lepidodendron and the megaspores of
Lycopodium; it is difficult therefore to decide to which category they
should be referred. Spencerites is clearly distinct from
Lepidostrobus; the absence of a ligule, the manner of attachment of
the sporangia, and the form and size of the spores, are characteristic
features.
A comparison of Spencerites with the strobili of Lycopodium
cernuum (figs. 123, 126–129) has recently been made by Lang, who
draws attention to the striking agreement as regards general plan
and even detailed structural features between the Palaeozoic and
the recent type of strobilus. It is interesting to find, as Lang points
out, that in the original account of the fossil cone by Williamson, the
view is expressed that the sporangiophores were confluent. An
examination of the section figured by Williamson[445] led Lang to
confirm this opinion. It would be out of place to enter here into a
detailed comparison of Spencerites insignis and the cone of
Lycopodium, but the resemblances are considered by Lang to be
sufficiently close to suggest that the striking similarity may be
indicative of relationship[446].
It is worthy of notice that the radial section of Spencerites (fig. 192)
presents a fairly close resemblance to a corresponding section
through a cone-scale of Agathis (Kauri Pine)[447]. In each case the
megasporangium is attached by a narrow pedicel to the sporophyll
and the latter has a similar form in the two plants, though the extent
of the resemblance is somewhat lessened by Lang’s more complete
account of the Palaeozoic type. If the Spencerites sporangia
possessed an integument the similarity with the Agathis ovule would
of course be much closer: recent palaeobotanical investigations
have shown that ovules and sporangia are not separated by
impassable barriers.
[Since this Chapter was set up in type a paper has appeared by Dr
Bruno Kubart on a new species of Spencerites spore, S.
membranaceus, from the Ostrau-Karwiner Coal-basin (Austria). The
spores are larger than those of S. insignis and in some the cells of a
prothallus are preserved. Kubart figures a section of a spore
containing a group of seven cells, a central cell, which he regards as
an antheridial mother-cell, surrounded by six wall-cells. Kubart (90).]
 CHAPTER XVI.
Sigillaria.

i. General.
In view of the close resemblance between Lepidodendron and
Sigillaria, another lycopodiaceous plant characteristic of
Carboniferous and Permian floras, a comparatively brief description
of the latter genus must suffice, more particularly as Lepidodendron
has received rather an undue share of attention. Sigillaria, though
abundantly represented among the relics of Palaeozoic floras,
especially those preserved in the Coal-Measures, is rare in a
petrified state, and our knowledge of its anatomy is far from
complete. In external form as in internal structure the difference
between the two genera are not such as enable us to draw in all
cases a clearly defined line of separation.
In the Antediluvian Phytology, Artis[448] figured a fossil from the
Carboniferous sandstones of Yorkshire which he called Euphorbites
vulgaris on account of a superficial resemblance to the stems of
existing succulent Euphorbias. Rhode[449] also compared Sigillarian
stems with those of recent Cacti. The specimen described by Artis is
characterised by regular vertical and slightly convex ribs bearing
rows of leaf-scars in spiral series, like those on the cushions of
Lepidodendron. A few years earlier Brongniart[450] had instituted the
genus Sigillaria[451] for plants with ribbed but not jointed stems
bearing “disc-like impressions” (leaf-scars) disposed in quincunx; the
type-species named by the author of the genus Sigillaria scutellata is
identical, as Kidston[452] points out, with Euphorbites vulgaris of Artis
and with the plant afterwards figured by Brongniart as S.
pachyderma[453]. Brongniart in 1822 figured another type of stem
characterised by the absence of ribs and by prominent spirally
arranged cushions bearing relatively large leaf-scars like the upper
part of the specimen shown in fig. 203; this he named Clathraria
Brardii, a well-known and widely distributed Carboniferous and
Permian species now spoken of as Sigillaria Brardi (figs. 196, A–C;
203). A third type of stem figured by Brongniart as Syringodendron
striatum[454] agrees with Sigillaria scutellata in having ribs, but differs
in the substitution of narrow oval ridges or depressions for leaf-scars;
this is now recognised as a partially decorticated Sigillaria, in which
the vascular bundle of each leaf is represented by a narrow ridge or
depression. The name Syringodendron, originally used by Sternberg,
is conveniently applied to certain forms of Sigillarian stems which
have lost their superficial tissues. A fourth generic name, Favularia,
was instituted by Sternberg[455] for Sigillarian stems with ribs covered
with contiguous leaf-scars of hexagonal form and prominent lateral
angles (fig. 193, A; fig. 200, G).
 Fig. 193.
A. Sigillaria elegans Brongn.
B. Sigillaria rugosa Brongn. Middle Coal-Measures.
C. Omphalophloios anglicus Kidst. Barnsley.
D. Sigillaria elegans Brongn.
E. Sigillaria tessellata Brongn.
(A, B, C, E, about ¾ nat. size. Dr Kidston’s Collection.)

The generic or sub-generic title Rhytidolepis, also instituted by

Sternberg, is applied to ribbed Sigillarian stems such as S.
scutellata, S. rugosa (fig. 193, B), S. mammillaris (fig. 195), or S.
laevigata (fig. 196, D). Goldenberg[456] proposed the name
Leiodermaria for smooth Sigillarian stems with leaf-scars not in
contact with one another (fig. 196, C).
The shoot system of Sigillaria consisted of a stout stem tapering
upwards to a height of 100 feet[457] or more as an unbranched
column, with its dome-shaped apex[458] covered with linear grass-like
leaves or, in some species, such as Sigillaria Brardi[459], S.
Eugenii[460], etc., the main trunk was occasionally divided by
apparently equal dichotomy. The younger portions of the stem or
branches were in some species clothed with leaves separated by a
narrow zigzag groove surrounding their hexagonal bases, while in
other forms each leaf was seated on a more or less prominent
cushion having the form illustrated by Sigillaria McMurtriei (fig. 194)
or by the example represented in fig. 200, H; or as in the ribbed
species shown in figs. 193, B, and 195, the leaves in vertical series
were separated from one another by longer portions of the ribs. As in
Lepidodendron the cushions are frequently characterised by irregular
transverse wrinklings and other[461] surface-ornamentation which in
some instances at least may have been produced as the result of
post-mortem shrinkage of superficial tissue. From the rarity of shoots
with the foliage attached, it would seem that the leaves persisted for
a comparatively short time and were cut off by an absciss-layer
leaving behind a well-marked leaf-scar area. The linear leaves,
reaching in rare cases a length of one metre (e.g. S.
lepidodendrifolia) but usually much shorter, possessed a single
median bundle, and the lower face was characterised by two
stomatal grooves and a median keel. It is not uncommon to find leaf-
bases of Sigillaria detached from the stem and preserved as
separate impressions. The term Sigillariophyllum used by
Grand’Eury[462] may be applied to detached leaves, though it is by no
means easy to distinguish between the foliage of Sigillaria and
Lepidodendron. A comparison of a typical species of Sigillaria, such
as S. rugosa (fig. 193, B) or S. Brardi (fig. 196, A–C) with a typical
Lepidodendron reveals obvious differences in the form of the leaf-
cushion, but in some cases the distinction becomes purely arbitrary.
Fig. 194. Sigillaria McMurtriei Kidst. From a specimen from the Upper
Coal-Measures of Radstock, in the British Museum (V. 952). Nat.
size.

Fig. 195. Sigillaria mammillaris. (Rhytidolepis form.) From a specimen in

the Manchester Museum. p, parichnos; l, ligule-pit; t, leaf-trace; c,
cushion; s, leaf-scar.
 Fig. 196.
A–C. Sigillaria Brardi. (A after Germar; B, C after Zeiller.)
D. Sigillaria laevigata.
E. Lepidodendron Wortheni (D and E after Zeiller).

Immediately above the centre of the upper boundary of a

Sigillarian leaf-scar a ligule pit may often be detected, as shown in
fig. 195, l, and in some cases, e.g. a specimen figured by Germar[463]
(fig. 196, A) as Sigillaria spinulosa (identical with S. Brardi), some
circular scars with a central pit surrounded by a raised rim occur on
the surface of the stem, either singly or in pairs, near the leaf-scars;
these, it is suggested, may represent the position of adventitious
roots or, as Germar thought, of some deciduous spinous processes.
The leaf-scars are frequently hexagonal in shape, with the lateral
angles either rounded (fig. 200, F) or sharply pointed (fig. 200, G, H);
each scar bears three smaller scars as in Lepidodendron, a central
circular, oval or crescentic leaf-trace scar and larger oval or slightly
curved scars formed by the two parichnos arms (fig. 195, p). The
larger size of the parichnos arms, the individual cells of which may
often be detected as a fine punctation, is a distinguishing feature of
the genus, but otherwise the structure is very similar to that in
Lepidodendron. As shown in figs. 195, 200, F, G, the three scars
may occur nearer the upper than the lower margin of the leaf-base
area.
Lepidodendron Wortheni[464] (fig. 196, E), described from North
America by Lesquereux[465], by Zeiller[466] from France, and by
Kidston[467] from the Upper and Middle Coal-Measures of England,
may be quoted as a Lepidodendron bearing a close resemblance to
Sigillaria. The shoots bear cushions two or three times as long as
broad and without the usual median division, but with numerous
irregular and discontinuous transverse wrinklings. Lepidodendron
Peachii Kidston[468] affords another example of a form agreeing both
with Sigillaria and with Lepidodendron. An Upper Devonian type
described by White[469] as Archaeosigillaria primaeva affords a
striking instance of the combination on one stem of Sigillarian and
Lepidodendroid leaf-cushions.
The difference between the original surface of a Sigillaria stem
and that of partially decorticated specimens is seen in figs. 196, C
and D; in fig. C the bark of Sigillaria Brardi shows the characteristic
wrinklings of the superficial tissue, while at a slightly lower level the
leaf-scars are replaced by the parichnos casts, a, and fine
longitudinal striations represent the elongated phelloderm cells laid
bare by the exfoliation of the surface-layers. Similarly, in the rib of
Sigillaria laevigata (fig. 196, D) the parichnos arms, p, and the
longitudinal striations are exposed at the lower level, while the
surface is smooth and bears rows of widely separated leaf-scars.
 Fig. 197. Carica sp. From the Royal Gardens, Kew. (Much reduced.) M.S.
The older part of a Sigillarian stem may present an appearance
very different from that of the younger shoots. The leaf-cushions may
be stretched apart as the result of elongation and increase in girth,
while in some cases the arrangement of the leaf-scars may vary on
the same axis as the result of inequalities in growth or changing
climatic conditions. The contiguous arrangement of the leaf-scars
and narrow cushions characteristic of the Clathrarian form of stem,
as was first demonstrated by Weiss[470], and afterwards illustrated by
Zeiller[471] and Kidston, may be gradually replaced (on the same
specimen) by a more distant disposition of the leaf-scars separated
by a smooth intervening surface of bark. The specimen of S. Brardi
reproduced in part in fig. 203, and first figured by Kidston, affords an
example of three “species” on one piece of stem, S. Brardi Brongn.,
S. denudata Goepp. and S. rhomboidea Brongn.[472]
The piece of Carica stem, represented in fig. 197, illustrates the
danger of trusting to the disposition of leaves as a specific criterion.
Similarly, in the ribbed forms the degree of separation of the leaf-
scars is by no means uniform in a single species[473]. Some authors
have adopted a two-fold classification of Sigillarian stems proposed
by the late Prof. Weiss[474] of Berlin, who divided the Sigillariae into
(A) Sub-Sigillariae, comprising Leiodermariae and Cancellatae, and
(B) Eu-Sigillariae, including Favulariae and Rhytidolepis.
Grand’Eury[475] adopts the terms Rhytidolepis and Leiodermaria for
ribbed and smooth stems respectively, the type to which the name
Clathraria was applied by Brongniart being in some cases at least
the young form of Leiodermarian stems. While recognising the
artificial distinction implied by such terms as Rhytidolepis,
Leiodermaria, and other sub-generic titles, we may conveniently
speak of the two main types of Sigillaria stems as ribbed and
smooth.
Still older stems of Sigillaria are not uncommon from which the
leaf-scars and other superficial tissues have been exfoliated, leaving
exposed a longitudinally fissured surface of secondary cortex
characterised by pairs of considerably enlarged parichnos strands
(fig. 198) which are sometimes partially or wholly fused into one
(Syringodendron state of Sigillaria). The single or double nature of
the elliptical or circular parichnos areas is doubtless due to the
degree of exfoliation, which may extend sufficiently deep into the
cortex to reach the level of the parichnos before the single strand
has bifurcated (cf. Lepidodendron, p. 100). In the Museums of
Manchester, Newcastle, and other places casts of large Sigillaria
stems may be seen, which illustrate the differences in breadth and
regularity of the vertical ribs, and in the size and shape of the
parichnos areas in different regions of a partially decorticated stem.
A cast of a ribbed species in the Manchester Museum, having a
length of 185 cm. and a breadth of 56 cm., shows in the upper
portion straight vertical grooves and broad ribs bearing pairs of
parichnos scars 11 mm. long; in the lower portion the ribs tend to
become obliterated and the parichnos scars, 2 cm. in length, may be
partially fused and arranged in much less regular vertical series. A
feature of these older ribbed Sigillarian stems is the increase in the
number of the ribs from below upwards. Kidston[476] has described a
specimen in the Sunderland Museum, 6 feet 6 inches long, with a
circumference at the slightly bottle-shaped base of 5 feet. On the
lower portion of the stem there are 29 broad ribs; about one-third the
height many of these bifurcate, producing as many as 40 ribs in the
upper part where the cast has a circumference of 3 feet. The
increase in number of the ribs is due in part to bifurcation, but also to
the intercalation of new ones. As Kidston points out, this example
shows that as a stem grew in length additional leaves were
developed at the apex. A similar stem, which illustrates very clearly
the increase in the number of ribs from below upwards, may be seen
in the Newcastle Museum.
Grand’Eury[477] has described an example of an old stem of a
ribless species of Sigillaria, Syringodendron bioculatum, bearing
single and double parichnos areas of nearly circular form and with a
diameter of 1–2 cm. In a specimen figured by Renault and Roche[478]
(Syringodendron esnostense) from the Culm strata in France, the
parichnos scars reach a length of 3 cm. As seen in the fragment of a
ribbed Sigillaria represented in fig. 198, the large parichnos areas
exhibit a distinct surface pitting in contrast to the fine longitudinal
striation of the rib; the difference in surface-appearance is due to the
nature of the tissue, which in the parichnos consists of fairly large
parenchymatous elements with groups of secretory cells[479], and in
the exposed cortex of elongated elements. The vertical line in the
middle of fig. 198, which occurs in the middle of the rib, has probably
been formed by splitting of the bark.
 Fig. 198. Sigillaria with large parichnos areas. (⅓ nat. size.) M.S.
Grand’Eury’s description of fossil forests of Sigillariae in the rocks
of the St Étienne[480] district affords a striking picture of these
arborescent Pteridophytes; he speaks of the stems of some of the
trees as swollen like a bottle at the base, characterised by the
Syringodendron features and terminating below in short repeatedly
forked roots of the type known as Stigmariopsis. Other specimens of
Sigillaria stumps show a marked decrease in girth towards the base;
this tapered form is regarded by Grand’Eury as the result of the
development of aerial columnar stems from underground rhizomes.
The nature of the root-like organs of Sigillaria is dealt with in the
sequel: a brief reference may, however, be made to the occurrence
of stumps of vertical trunks which pass downwards into regularly
forked and spreading arms. These arms lie almost horizontally in the
sand or mud like the underground rhizomes of Phragmites and other
recent plants growing in swampy situations where water is abundant
and where deeper penetration of the soil would expose them to an
insufficient supply of oxygen[481]. It is certain that Sigillaria had no
tap-root, but was supported on spreading subterranean organs
bearing spirally disposed long and slender rootlets which absorbed
water from a swampy soil.
 Fig. 199. Partially decorticated stem of Sigillaria showing two zones of
cone-scars. From a cast in the Sedgwick Museum, Cambridge.
M.S. (⅕ nat. size.)
The regularity of the leaf-scar series on a Sigillarian stem may be
interrupted by the occurrence of oval scars with a central scar and
surrounding groove (fig. 193, E); these occur in zones at more or
less regular intervals on the stem, as seen in the partially
decorticated cast represented in fig. 199. Zeiller has pointed out that
the rows of oval or circular scars, which mark the position of
caducous stalked strobili, may occur between the leaf-scars in
vertical series, each of which may include as many as 20 scars,
while in other cases a single series of such cone-scars may encircle
the stem[482]. The zones are usually of uneven breadth, as in S.
Brardi, and their occurrence produces some deformation of the
adjacent leaf-scars.
By the earlier writers Sigillaria was compared with succulent
Euphorbias, Cacti, and Palms; Brongniart[483] at first included
undoubted Sigillarian stems among Ferns, but after investigating an
agatized stem from Autun, he referred Sigillaria to the
Gymnosperms[484] on the ground that it had the power of producing
secondary wood. It was then supposed that Lepidodendron
possessed only primary xylem, and that the presence of a vascular
meristem in Sigillaria necessitated its separation from the
lycopodiaceous genus Lepidodendron and its inclusion in the higher
plants. By slow degrees it was recognised, as in the parallel case of
the genus Calamites, that the presence or absence of secondary
vascular tissue is a character of small importance. Williamson,
whose anatomical researches played the most important part in
ridding the minds of palaeobotanists of the superstition that
secondary growth in thickness is a monopoly of the Phanerogams,
spoke in 1883 of the conflict as to the affinities of Lepidodendron and
Sigillaria as virtually over but leaving here and there “the ground-
swell of a stormy past[485].” In 1872 the same author had written: “If
then I am correct in thus bringing the Lepidodendra and Sigillariae
into such close affinity, there is an end of M. Brongniart’s theory, that
the latter were gymnospermous exogens, because the cryptogamic
character of the former is disputed by no one; we must rather
conclude as I have done that the entire series represents, along with
the Calamites, an exogenous group of Cryptogams in which the
woody zone separated a medullary from a cortical portion[486].”
In 1879 Renault[487] expressed the opinion that Brongniart by his
investigation of the anatomy of Sigillaria elegans had established in
a manner “presque irréfutable” that Sigillaria must be classed as a
Gymnosperm showing affinity with the Cycads.
 In 1855 Goldenberg[488] described some strobili which he regarded
as those of Sigillaria and recognised their close resemblance to a
fertile plant of Isoetes. He was led to the conclusion, which had little
influence on contemporary opinion, that Sigillaria is related to Isoetes
and must be classed among Pteridophytes. To these long and
narrow strobili Schimper gave the name Sigillariostrobus[489]. In 1884
Zeiller[490] supplied confirmation of Goldenberg’s view by the
discovery of cones borne on pedicels with Sigillarian leaf-scars, thus
demonstrating the generic identity of cones and vegetative shoots,
which Goldenberg had connected on the evidence of association.
Zeiller’s more recent work[491] and the still later researches of
Kidston[492] have added considerably to our knowledge of the
morphology of Sigillarian cones. Grand’Eury’s remark made so
recently as 1890[493] that opinion in regard to the Gymnospermous
nature of Sigillaria is losing ground every day, bears striking
testimony to the pertinacity with which old beliefs linger even in the
face of overwhelming proof of their falsity.
It is remarkable, in view of the abundance of vegetative shoots,
how rarely undoubted Sigillarian strobili have been found; this may,
however, be in part due to a confusion with Lepidostrobi which so far
as we know do not differ in important respects from
Sigillariostrobi[494].
There can be no doubt that Sigillaria usually produced its cones on
slender pedicels which bore a few leaves or bracts in irregular
verticils, or in short vertical series on comparatively stout stems, an
arrangement reminding us of the occurrence of flowers on old stems
of Theobroma and other recent Dicotyledons. As Renault[495] pointed
out the fertile shoots are axillary in origin.
Dr Kidston[496] is of opinion that certain species of Sigillaria bore
cones sessile on large vegetative shoots characterised by two
opposite rows of cup-like depressions like those in the Ulodendron
form of Lepidodendron Veltheimianum (fig. 157). He has described
the Ulodendron condition of two species, Sigillaria discophora
(König) and S. Taylori (Carr.); the cup-like depressions may have a
diameter of several centimetres and are distinguished from those of
Bothrodendron by the almost central position of the umbilicus. The
specimens which he figures as S. discophora are identified by him
with the stem figured by König as Lepidodendron discophorum and
by Lindley and Hutton[497] as Ulodendron minus. We have already
dealt with the nature of Ulodendron shoots, expressing the opinion
that in spite of the often quoted specimen described by D’Arcy
Thompson[498], in which a supposed cone occurs in one of the cups,
there is no satisfactory case of any undoubted cone having been
found attached to the large Ulodendron scars. It is more probable
that the Ulodendron depressions represent the scars of branches,
either elongated axes, or possibly in some cases deciduous
tuberous shoots which served as organs of vegetative reproduction.
A specimen figured by Kidston as Sigillaria Taylori from the
Calciferous sandstone of Scotland[499] bears a row of slightly
projecting “appendicular organs” attached to a Ulodendron axis; but
these furnish no proof of their strobiloid nature. The main question is,
are these Ulodendron shoots correctly identified by Kidston as
Sigillarian? The surface of the specimens shows crowded
rhomboidal scars surrounded in some cases by a very narrow border
or cushion; the general appearance is, as Kidston maintains, like that
of Sigillaria Brardi in which the leaf-scars are contiguous (e.g. fig.
203, upper part). None of the leaf-scars exhibit the three
characteristic features, the leaf-trace and parichnos scars, but only
one small scar appears on each leaf-base area. In a more recent
paper Kidston figures a small piece of a stem from Kilmarnock,
which he identifies as Sigillaria discophora, showing the three
characteristic scars on the leaf-base area. There is no doubt as to
the Sigillarian nature of this specimen, but it is not clear if the piece
figured is part of a Ulodendron shoot[500].
Prof. Zeiller[501] retains the older name Ulodendron minus Lind. and
Hutt. in place of König’s specific designation and dissents from
Kidston’s identification of Ulodendron minus and U. majus of Lindley
and Hutton as one species; he is also inclined to refer these
Ulodendron axes to Lepidodendron. In spite of the superficial
resemblance to Sigillaria of the specimens described by Kidston, and
which I have had an opportunity of examining, I venture to regard
their reference to that genus as by no means definitely established.
We must recognise the difficulty in certain cases of drawing any