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 Fig. 232. Davallia aculeata. (⅖ nat. size.)
It is noteworthy that while certain vegetative features may in some
cases be cited as family-characters, such features are not usually of
much value from a taxonomic point of view. While the typical tree
ferns are practically all members of the Cyatheaceae, a few
members of other families, e.g. Todea barbara (Osmundaceae) and
the monotypic Indian genus Brainea (Polypodiaceae), form erect
stems several feet in height; but these differ in appearance from the
Palm-like type of the Cyatheaceous tree ferns. On the other hand,
the thin, almost transparent, leaf of Hymenophyllum tunbridgense
and other filmy ferns is a character shared by several species of
Todea, Asplenium resectum, and Danaea trichomanoides
(Marattiaceae); the filmy habit is essentially a biological adaptation.
The form of frond represented by certain species of Gleichenia,
characterised by a regular dichotomy of the axis and by the
occurrence of arrested buds, is on the whole a trustworthy character,
though Davallia aculeata (bearing spines on its rachis) (fig. 232) and
Matonia sarmentosa have fronds with a similar mode of branching
and also bear arrested radius-buds. A limited acquaintance with
ferns as a whole often leads us to regard a certain form of leaf as
characteristic of a particular species, but more extended enquiry
usually exposes the fallacy of relying upon so capricious a feature.
The form of leaf illustrated by Trichomanes reniforme is met with also
in Gymnogramme reniformis and is fairly closely matched by the leaf
of Scolopendrium nigripes. The fronds of Matonia pectinata (figs.
227, 228) bear a close resemblance to those of Gleichenia
Cunninghami, Adiantum pedatum, and Cheiropteris
[704]
palmatopedata .

The habit, leaf-form, and distribution of Ferns.

The full accounts of the structure and life-history of the common
Male Fern, given by Scott in his Structural Botany and by Bower in
the Origin of a Land Flora, render superfluous more than a brief
reference to certain general considerations in so far as they may
facilitate a study of fossil types.
In size Ferns have a wide range: at the one extreme we have the
filmy fern Trichomanes Goebelianum[705], growing on tree stems in
Venezuela, with leaves 2·5 to 3 mm. in diameter, and at the other the
tree ferns with tall columnar stems reaching a height of 40 to 50 feet
and terminating in a crown of fronds with a spread of several feet. A
common form of stem is represented by the subterranean or
creeping rhizome covered with ramental scales or hairs: the remains
of old leaves may persist as ragged stumps, or, as in Oleandra,
Polypodium vulgare and several other species, the leaf may be cut
off by the formation of an absciss-layer[706] leaving a clean-cut peg
projecting from the stem. As a rule the branches bear no relation to
the leaves and are often given off from the lower part of a petiole, but
in a few cases, e.g. in the Hymenophyllaceae, it is noteworthy that
true axillary branching is the rule[707]. In the typical tree-fern the
surface resembles that of a Cycadean trunk covered with persistent
leaf-bases and a thick mass of roots. Among epiphytic ferns highly
modified stems are occasionally met with, as in the Malayan species
Polypodium (Lecanopteris) carnosum and P. sinuosum[708].
The leaves of ferns are among the most protean of all plant
organs; as Darwin wrote, “the variability of ferns passes all
bounds[709].” The highly compound tri- or quadripinnate leaves of
such species as Pteris aquilina, Davallia and other genera stand for
the central type of fern frond; others exhibit a well-marked
dichotomy, e.g. Lygodium, Gleichenia, Matonia, etc., a habit in all
probability associated with the older rather than with the more
modern products of fern evolution. Before attempting to determine
specifically fossil fern fronds, it is important to familiarise ourselves
with the range of variability among existing species and more
especially in leaves of the same plant. A striking example of
heteromorphy is illustrated in fig. 233. Reinecke[710] has figured a
plant of Asplenium multilineatum in which the segments of the
compound fronds assume various forms. In Teratophyllum
aculeatum var. inermis Mett., a tropical climbing fern believed by
Karsten[711] to be identical with Acrostichum (Lomariopsis)
sorbifolium,—an identification which Goebel[712] questions,—the
fronds which stand free of the stem supporting the climber differ
considerably from the translucent and much more delicate filmy
leaves pressed against the supporting tree. From this fern alone Fée
is said to have created 17 distinct species. In this, as in many other
cases, differences in leaf-form are the expression of a physiological
division of labour connected with an epiphytic existence. Some
tropical species of Polypodium (sect. Drynaria), e.g. P. quercifolium
(fig. 234 and fig. 231, D), produce two distinct types of leaf, the large
green fronds, concerned with the assimilation of carbon and spore-
production, being in sharp contrast to the small slightly lobed brown
leaves which act as stiff brackets (fig. 234, M) for collecting humus
from which the roots absorb raw material. Similarly in Platycerium
the orbicular mantle-leaves differ widely from the long pendulous or
erect fronds fashioned like the spreading antlers of an elk. In
Hemitelia capensis, a South African Cyatheaceous species, the
basal pinnae assume the form of finely divided leaves identified by
earlier collectors as those of a parasitic Trichomanes (fig. 235). In a
letter written by W. H. Harvey in 1837 accompanying the specimen
shown in fig. 235, he says, “Apropos of Hemitelia, be it known
abroad that supposed parasitical Trichomanes ... is not a parasite,
but a part of the frond of Hemitelia.” The delicate reduced pinnae
remain on the stem and form a cluster at the base of the fronds[713].

Fig. 233. Polypodium Billardieri Br. (¼ nat. size.) Middle Island, New
Zealand. From specimens in the Cambridge Herbarium.
 Fig. 234. Polypodium quercifolium. (Much reduced: M, Mantle-leaves.)
In many species the sporophylls are distinguished from the sterile
fronds by segments with little or no chlorophyllous tissue, as in
Onoclea struthiopteris[714] in which, each year, the plant produces a
funnel-shaped group of sterile leaves followed later in the season by
a cluster of sporophylls; or, as in many other genera, the fertile
leaves are distinguished also by longer petioles and thus serve as
more efficient agents of spore-dissemination. In Ceratopteris the
narrow segments of the taller fertile leaves are in striking contrast to
the broader pinnules of the submerged foliage leaves. Leaf-form is in
many cases obviously the expression of environment; the
xerophilous fern Jamesonia[715] from the treeless paramos of the
Andes[716] is characterised by its minute leaflets with strong revolute
margins and a thick felt of hairs on the lower surface; in others,
xerophilous features take the form of a covering of overlapping
scales (Ceterach), or a development of water-tissue as in the fleshy
leaves of the Himalayan fern Drymoglossum carnosum. In the
Bracken fern Boodle[717] has shown how the fronds may be classed
as shade and sun leaves; the former are spreading and softer, while
the latter are relatively smaller and of harder texture (fig. 236, a and
b). Even in one leaf six feet high, growing through a dense bush of
gorse and bramble, the lower part was found to have the features of
a shade leaf, while the uppermost exposed pinnae were xerophilous.

Fig. 235. Hemitelia capensis R. Brown. Nat. size. a, Pinna of normal frond.
[From a specimen in the British Museum. M.S.]
 Fig. 236a. Pteris aquilina.
Part of leaf from greenhouse. (¼ nat. size.) After Boodle.
PTERIS

The resemblance between some of the filmy Hymenophyllaceae

and thalloid Liverworts[718] is worthy of mention as one of the many
possible pitfalls to be avoided by the palaeobotanical student. The
long linear fronds of such genera as Vittaria and Monogramme might
well be identified in a fossil state as the leaves of a grass-like
Monocotyledon, or compared with the foliage of Isoetes or Pilularia.
The resemblance of some fern leaves with reticulate venation to
those of Dicotyledons has led astray experienced palaeobotanists; it
is not only the anastomosing venation in the leaves of several ferns
that simulates dicotyledonous foliage, but the compound leaves of
many dicotyledons, e.g. Paullinia thalictrifolia (Sapindaceae) and
species of Umbelliferae, may easily be mistaken for fronds of ferns.

Fig. 236b. Pteris aquilina.

Leaf from the same plant grown out of doors. (¼ nat. size.)
After Boodle.
RECENT FERNS
 The dichotomously lobed lamina of some Schizaeas, e.g. S.
dichotoma and S. elegans (fig. 222), bears a close resemblance to
the leaves of Baiera or Ginkgo[719]. The original description by
Kunze[720] of the South African Cycad Stangeria paradoxa as a
Polypodiaceous fern illustrates the difficulty, or indeed impossibility,
of distinguishing between a sterile simply pinnate fern frond and the
foliage of some Cycads. The deeply divided segments of Cycas
Micholitzii[721] simulate the dichotomously branched pinnae of
Lygodium dichotomum, and the leaves of Aneimia rotundifolia (fig.
223) and other species are almost identical in form with the Jurassic
species Otozamites Beani, a member of the Cycadophyta.
There are certain facts in regard to the geographical distribution of
ferns to which attention should be directed. Mr Baker in his paper on
fern distribution writes: “With the precision of an hygrometer, an
increase in the fern-vegetation marks the wooded humid regions[722].”
If in a collection of fossil plants we find a preponderance of ferns we
are tempted to assume the existence of such conditions as are
favourable to the luxuriant development of ferns at the present day.
On the other hand, we must bear in mind the wonderful plasticity of
many recent species and the fact that xerophilous ferns are by no
means unknown in present-day floras.
Ferns are admirably adapted to rapid dispersal over comparatively
wide areas. Bower[723] estimates that in one season a Male Fern may
produce about 5,000,000 spores: with this enormous spore-output
are coupled a thoroughly efficient mechanism for scattering the
germs and an unusual facility for wind-dispersal. When Treub[724]
visited the devastated and sterilised wreck of the Island of Krakatau
in 1886, three years after the volcanic outburst, he found that twelve
ferns had already established themselves; the spores had probably
been carried by the wind at least 25 to 30 miles. It is not surprising,
therefore, to find that many ferns have an almost world-wide
distribution; and, it may be added, in view of their efficient means of
dispersal, wide range by no means implies great antiquity. Prof.
Campbell[725] has recently called attention to the significance of the
wide distribution of Hepaticae in its bearing on their antiquity; the
spores are incapable of retaining vitality for more than a short period,
and it is argued that a world-wide distribution can have been
acquired only after an enormous lapse of time. If we apply this
reasoning to the Osmundaceae among ferns, it may be legitimate to
assume that their short-lived green spores render them much less
efficient colonisers than the great majority of ferns; if this is granted,
the wide distribution of Osmundaceous ferns in the Mesozoic era
carries their history back to a still more remote past, a conclusion
which receives support from the records of the rocks.
The Bracken fern which we regard as characteristically British is a
cosmopolitan type; it was found by Treub among the pioneers of the
New Flora of Krakatau; in British Central Africa, it greets one at
every turn “like a messenger from the homeland[726]”; it grows on the
Swiss Alps, on the mountains of Abyssinia, in Tasmania, and on the
slopes of the Himalayas. The two genera Matonia (fig. 228) and
Dipteris, which grow side by side on Mount Ophir in the Malay
Peninsula, are examples of restricted geographical range and carry
us back to the Jurassic period when closely allied types flourished
abundantly in northern latitudes. Similarly Thyrsopteris elegans,
confined to Juan Fernandez, exhibits a remarkable likeness to
Jurassic species from England and the Arctic regions.
The proportion of ferns to flowering plants in recent floras is a
question of some interest from a palaeobotanical point of view; but
we must bear in mind the fact that the evolution of angiosperms,
effected at a late stage in the history of the earth, seriously disturbed
the balance of power among competitors for earth and air. The
abundance of ferns in a particular region is, however, an unsafe
guide to geographical or climatic conditions. Many ferns are
essentially social plants; the wide stretches of moorland carpeted
with Pteris aquilina afford an example of the monopolisation of the
soil by a single species. In Sikkim Sir Joseph Hooker speaks of
extensive groves of tree ferns, and in the wet regions of the Amazon,
Bates[727] describes the whole forest glade as forming a “vast
fernery.” In a valley in Tahiti Alsophila tahitiensis is said to form “a
sort of forest almost to the exclusion of other ferns[728].” In the
abundance of Glossopteris (figs. 334, etc.) fronds spread over wide
areas of Permo-Carboniferous rocks in S. Africa, Australia, and
India, we have a striking instance of a similar social habit in an
extinct fern or at least fern-like plant.
Acrostichum aureum, with pinnate fronds several feet long, is an
example of a recent fern covering immense tracts, but this
species[729] is more especially interesting as a member of the
Filicineae characteristic of brackish marshes and the banks of
tropical rivers in company with Mangrove plants and the “Stemless
Palm” Nipa. This species exhibits the anatomical characters of a
water-plant and affords an interesting parallel with some Palaeozoic
ferns (species of Psaronius) which probably grew under similar
conditions.

The Anatomy of Ferns.

The text-book accounts of fern-anatomy convey a very inadequate
idea of the architectural characters displayed by the vascular
systems of recent genera. When we are concerned with the study of
extinct plants it is essential to be familiar not only with the commoner
recent types, but particularly with exceptional or aberrant types. The
vascular system of many ferns consists of strands of xylem
composed of scalariform tracheae associated with a larger or smaller
amount of parenchyma, surrounded either wholly or in part (that is
concentric or bicollateral) by phloem: beyond this is a pericycle, one
layer or frequently several layers in breadth, limited externally by an
endodermis, which can usually be readily recognised. The vascular
strands are embedded in the ground-tissue of the stem consisting of
thin-walled parenchyma and, in most ferns, a considerable quantity
of hard and lignified mechanical tissue. The narrow protoxylem
elements are usually characterised by a spiral form of thickening, but
in slow-growing stems the first-formed elements are frequently of the
scalariform type.
A study of the anatomy of recent ferns both in the adult state and
in successive stages of development from the embryo has on the
whole revealed “a striking parallelism[730]” between vascular and
sporangial characters in leptosporangiate ferns. For a masterly
treatment of our knowledge of fern anatomy from a phylogenetic
point of view reference should be made to Mr Tansley’s recently
published lectures: within the limits of this volume all that is possible
is a brief outline of the main types of vascular structure illustrated by
recent genera.

Fig. 237.
A. Matonia pectinata (petiole).
B. M. pectinata (stem).
C. Gleichenia dicarpa (stem): p, petiole; pp, protophloem; position
of protoxylem indicated by black dots.
D. Matonidium.
E. Trichomanes reniforme: pp, protophloem.
(C, E, after Boodle; D, after Bommer.)
To Prof. Jeffrey[731] we owe the term protostele which he applied to
a type of stele consisting of a central core of xylem surrounded by
phloem, pericycle, and endodermis. While admitting that steles of
this type may sometimes be the result of the modification of less
simple forms, we may confidently regard the protostele as
representing the most primitive form of vascular system. The genus
Lygodium affords an example of a protostelic fern; a solid column of
xylem tracheae and parenchyma is completely encircled by a
cylinder of phloem succeeded by a multi-layered pericycle and an
endodermis of a single layer of cells. In this genus the stele is
characterised by marginal groups of protoxylem; it is exarch. An
almost identical type is represented by species of Gleichenia, but
here the stele is mesarch, the protoxylem being slightly internal (fig.
237, C). Trichomanes scandens (fig. 238) has an exarch protostele
like that of Lygodium; but, as Boodle[732] has suggested, the
protostelic form in this case is probably the result of modification of a
collateral form of stele such as occurs in Trichomanes reniforme (fig.
237, E). A second type of stele has been described in species of
Lindsaya[733] in which the xylem includes a small group of phloem
near the dorsal surface. This Lindsaya type is often passed through
in the development of “seedling” ferns and may be regarded as a
stage in a series leading to another well-marked type, the
solenostele. The solenostele[734], a hollow cylinder of xylem lined
within and without by phloem, pericycle, and endodermis, occurs in
several genera belonging to different families, e.g. Dipteris, species
of Pteris, species of Lindsaya, Polypodium, Jamesonia, Loxsoma,
Gleichenia and other genera. In a smaller number of ferns the stele
consists of what may be called a medullated protostele similar to the
common form of stele in Lepidodendron: this type is found in species
of Schizaea and in Platyzoma (fig. 239). It is important to notice that
in the solenostele and as a rule in the medullated protostele when a
leaf-trace passes out from the rhizome stele the vascular cylinder is
interrupted by the formation of a foliar gap (Platyzoma[735], fig. 239, is
an exception). This fact has been emphasized by Jeffrey[736] who
draws a distinction between the Lycopodiaceous type of stele, which
is not broken by the exit of leaf-traces, and the fern stele in which
foliar gaps are produced: the former he speaks of as the
cladosiphonic type (Lycopsida) and the latter as the phyllosiphonic
(Pteropsida).
 Fig. 238. Stele of Trichomanes scandens: px, protoxylem; s, endodermis.
From Tansley, after Boodle.

Fig. 239. Platyzoma microphylla. l.t., leaf-trace; i.e., internal endodermis.

(After Tansley; modified from Boodle.)
The transition to a hollow cylinder of xylem from a protostele may
be described as the result of the replacement of some of the axial
conducting tracheae by parenchyma or other non-vascular tissue
consequent on an increase in diameter of the whole stele and the
concentration of the true conducting elements towards the
periphery[737].
The occurrence of the internal cylinder of phloem, pericycle, and
endodermis in a solenostele is rendered intelligible by a study of fern
seedlings and by a comparative examination of transitional types
connecting protosteles and solenosteles through medullated
protosteles and steles of the Lindsaya type. A further stage in stelar
evolution is illustrated by what is termed the dictyostele, the
arrangement of vascular tissue characteristic of Nephrodium Filix-
mas, Cyathea (fig. 240), Polypodium vulgare and many other
common ferns.

Fig. 240. Cyathea Imrayana. (From Tansley after de Bary.) (Sclerenchyma

represented by black bands.)
If a solenostele is interrupted by leaf-gaps at intervals sufficiently
close to cause overlapping, a transverse section at any part of the
stele will show apparently separate curved bands of concentrically
arranged xylem and phloem, which on dissection are seen to
represent parts of a continuous lattice-work or a cylinder with the
wall pierced by large meshes. The manner of evolution of the
dictyostele has been ably dealt with by Gwynne-Vaughan[738] and
other authors. In a few ferns, e.g. Matonia pectinata[739], a transverse
section of the stem (fig. 237, B) reveals the presence of two or in
some cases three concentric solenosteles with a solid protostele in
the centre: this polycylic type may be regarded as the expression of
the fact that in response to the need for an adequate water-supply to
the large fronds, ferns have increased the conducting channels by a
method other than by the mere increase of the diameter of a single
stele. Fig. 237, A, shows the vascular tissue of a petiole of Matonia
in transverse section.
The two genera of Osmundaceae, Todea and Osmunda, are
peculiar among recent ferns in having a vascular cylinder composed
of separate strands of xylem varying considerably in shape and size,
from U-shaped strands with the concavity facing the centre of the
stem and with the protoxylem in the hollow of the U, to oval or more
or less circular strands with a mesarch protoxylem or without any
protoxylem elements (fig. 221, A, B). These different forms are the
expression of the change in contour or in structure which the parts of
the lattice-work undergo at different levels in the stem[740]. Beyond
this ring of xylem bundles is a continuous sheath of phloem of
characteristic structure. A transverse section of a stem of Osmunda
regalis may show 15 or more xylem strands; in O. Claytoniana there
may be as many as 40. In Todea barbara (fig. 221, B) the leaf-gaps
are shorter, and in consequence of the less amount of overlapping
the xylem cylinder becomes an almost continuous tube. The recent
researches of Kidston and Gwynne-Vaughan[741] have resulted in the
discovery of fossil Osmundaceous stems with a complete xylem ring,
the stele being of the medullated protostele type; in another extinct
member of the family the stele consists of a solid xylem core. The
Osmundaceous type of stele is complicated in O. cinnamomea (fig.
221, A) by the occurrence of local internal phloem and by an internal
endodermis, a feature which leads Jeffrey to what I believe to be an
incorrect conclusion that the vascular arrangement found in
Osmunda regalis has been evolved by reduction from a stele in
which the xylem was enclosed within and without by phloem. New
facts recently brought to light enable us to derive the ordinary
Osmundaceous type from the protostele and solenostele. It is worthy
of remark that the Osmundaceae occupy a somewhat isolated
position among recent ferns; their anatomy represents a special
type, their sporangia differ in several respects from those of other
leptosporangiate ferns and in some features Osmunda and Todea
agree with the Eusporangiate ferns. The possession of such
distinguishing characters as these suggests antiquity; and the facts
of palaeobotany, as also the present geographical range of the
family, confirm the correctness of this deduction.
Before leaving the stelar structure of leptosporangiate fern stems,
a word must be added in regard to a type of structure met with in the
Hymenophyllaceae. In this family Trichomanes reniforme (fig. 237,
E) may be regarded, as Boodle suggests, as the central type: the
stele consists of a ring of metaxylem tracheae, the dorsal portion
having the form of a flat arch and the ventral half that of a straight
band. This flattened ring of xylem encloses parenchymatous tissue
containing scattered tracheae some of which are protoxylem
elements. In Trichomanes radicans the rhizome is stouter than in T.
reniforme and the stele consists of a greater number of tracheae.
The stele is cylindrical like that shown in fig. 238, but the centre is
occupied by two groups of protoxylem and associated parenchyma.
In Hymenophyllum tunbrigense the stele is of the subcollateral type;
the ventral plate of the xylem ring has disappeared leaving a single
strand of xylem with endarch protoxylem and completely surrounded
by phloem. Trichomanes muscoides possesses a still simpler stele
consisting of a slender xylem strand with phloem on one side only.
Reference has already been made to the occurrence in this family of
the protostelic type. The Hymenophyllaceae afford a striking
illustration of the modification in different directions of stelar structure
connected with differences in habit, and of the correlation of demand
and supply as shown in the varying amount of conducting tissue in
the steles of different species.
The leaf-trace in a great number of ferns is characterised by its C-
shaped form[742] as seen in transverse section: this in some genera,
e.g. Matonia (fig. 237, A), is complicated by the spiral infolding of the
free edges of the C; in other ferns (e.g. some Cyatheaceae) (fig.
278, C) the sides of the C are incurved, while in some species the
xylem is broken up into a large number of separate strands.
An elaborate treatment of the leaf-traces of ferns was published a
few years ago by MM. Bertrand and Cornaille[743] in which the
authors show how the various systems of vascular tissue in the
fronds of ferns maybe derived from a common type. As Prof.
Chodat[744] justly remarks this important work has not received the
attention it deserves, the neglect being attributed to the strange
notation which is adopted[745].
The roots of ferns are characterised by a uniformity of plan in
marked contrast to the wide range of structure met with in the stem
and to a less extent in the leaves. The xylem may consist of a plate
of scalariform tracheae with a protoxylem group at each end, or the
stele may include six or more alternating strands of xylem and
phloem.

II. Marattiales (Eusporangiate isosporous Filicales).

The Marattiaceae, the single family of ferns included in the
Marattiales, comprise the genera Angiopteris, Archangiopteris,
Marattia, Danaea, and Kaulfussia, which are for the most part
tropical in distribution. These genera are characterised by
eusporangiate sori or synangia, the presence of stipules at the base
of the petioles, and by the complex arrangement of the vascular
tissue. In view of the fact that many fossil ferns show a close
resemblance to the recent Marattiaceae, the surviving genera are
briefly described. The prothallus is green and relatively large.
Angiopteris. This genus occurs in Polynesia, tropical Asia, and
Madagascar; it is characterised by a short and thick fleshy stem
bearing large bipinnate leaves which occasionally show a forking of
the rachis[746], a feature reminiscent of some Palaeozoic fern-like
fronds. One of the large plants of Angiopteris evecta in the Royal
Gardens, Kew, bears leaves 12 feet in length with a stalk 6 inches in
diameter at the base. The sessile or shortly stalked and rather
leathery linear or broadly lanceolate pinnules have a prominent
midrib and dichotomously branched lateral veins. The surface of an
old stem is covered with the thick stumps of petioles enclosed by
pairs of fleshy stipules (fig. 241, A) and bears numerous fleshy roots,
which hang free in the air or penetrate the soil. The young fronds
(fig. 220, A) exhibit very clearly the characteristic circinate vernation.
The proximal part of each primary pinna is characterised by a
pulvinus-like swelling. The sporangia, in short linear elliptical sori
near the edge of the pinnules, consist of free sporangia (fig. 242, A–
D) provided with a peculiar type of “annulus”[747], in the form of a
narrow band of thicker-walled cells, which extends as a broad strip
on either side of the apex. An examination of sections through the
sporangia of Angiopteris in different planes[748] illustrates the difficulty
of determining the precise nature of the annulus in a petrified
sporangium which is seen only in one or two planes. Many of the
sporangia from the English Coal-Measures, compared by authors
with those of Leptosporangiate ferns, are in all probability referable
to the Marattiaceous type.

Fig. 241.
A. Angiopteris evecta. (Considerably reduced.)
B. Marattia fraxinea. Stipule. M.S.
The vascular system[749] of the stem constitutes a highly complex
dictyostelic or polycylic type which may consist of as many as nine
concentric series of strands of xylem surrounded by phloem, with
large sieve-tubes and a pericycle which abuts on the
parenchymatous ground-tissue without any definite endodermal
layer. A peculiarity in the vascular strands is that the first-formed
elements of the phloem lie close to the edge of the xylem, the
metaphloem being therefore centrifugal in its development. The
ground-tissue is devoid of mechanical tissue and is penetrated by
roots, a few of which arise from the outer vascular strands while
others force their way to the surface from the more internal
dictyosteles. Leaf-traces, consisting of several strands, are given off
from the outermost cylinder and a segment of the second dictyostele
moves out to fill the gap formed in the outermost network, while the
gap in the second cylinder receives compensating strands from the
third. A few layers below the surface of the petiole there is a ring of
thick-walled elements (s, fig. 243), and in both petiole and stem
numerous mucilage ducts and tannin-sacs occur in the ground-
tissue. It has been shown by Farmer and Hill[750] that in some of the
vascular strands in an Angiopteris stem a few secondary tracheae
are added to the primary xylem by the activity of the adjacent
parenchyma. The vascular bundles in the petiole form more or less
regular concentric series; they have no endodermis and are
characterised also by the large size of the sieve-tubes (st, fig. 243).