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Ang Diablo sa Filipinas translated into English with
annotations by Benedict Anderson Carlos Sardiña
Galache and Ramon Guillermo 1st Edition Isabelo De Los
Reyes Digital Instant Download
Author(s): Isabelo de los Reyes
ISBN(s): 9789712730610, 9712730611
Edition: 1
File Details: PDF, 22.04 MB
Year: 2014
Language: english
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Fig. 348. Neuropteridium validum, Feist. Nat. size. From the Karharbári
Coal-field, India. From Arber, after Feistmantel.
Neuropteridium validum. (Feistmantel[1370]). Fig. 348.
1869. Odontopteris Plantiana, Carruthers, Geol. Mag. Vol. vi.
p. 9, Pl. vi. figs. 2, 3.
1878. Neuropteris valida, Feistmantel, Mem. Geol. Surv.
India, Foss. Flor. Gondwana Syst., Vol. iii. p. 10, pl. ii.–vi.
1880. Neuropteridium validum, Feistmantel, Ibid. 2, p. 84.
The specimen represented in fig. 348 illustrates the main features
of Neuropteridium validum. This species is referred to by Dr
White[1371] as N. Plantianum on the ground of priority, and with a view
to perpetuate the name of the English engineer Nathaniel Plant who
discovered the species in a Brazilian Coal-field in the province of Rio
Grande do Sul. Feistmantel’s specific name is however retained as
being much better known. An examination of Mr Plant’s specimen in
the British Museum led me[1372] to speak of the Brazilian species as
identical with N. validum described by Feistmantel from Lower
Gondwana rocks of India. Zeiller[1373] had previously drawn attention
to the resemblance between the two sets of specimens. The frond of
N. validum may exceed 50 cm. in length. The lower pinnules may be
entire and semicircular in form while the upper and larger segments,
which may reach a length of 5 or 6 cm., are characterised by broad
lobes (fig. 348).
This type is represented in the flora of the Talchir-Karharbári series
(Lower Gondwana) of India[1374], in Permo-Carboniferous rocks of
Brazil and Argentine[1375], and in the sandstones of Vereeniging on
the borders of the Transvaal and Cape Colony. It is a characteristic
member of the Glossopteris Flora and occurs in association with
Glossopteris and Gangamopteris.
Neuropteridium intermedium (Schimper). Fig. 349.
This species has been figured by Schimper and Mougeot[1376] from
the Bunter of the Vosges and more fully described by
Blanckenhorn[1377] from the Bunter beds of Commern. The pinnate
leaves reach a length of 65 cm.; the lower semicircular pinnules pass
gradually into broadly linear segments characterised by an auriculate
base and a Neuropteris type of venation (fig. 354, D′, E). In the
example reproduced in fig. 349 from one of Blanckenhorn’s figures,
the fronds are attached to a short and thick rhizome bearing roots
and portions of old petioles.
Fig. 349. Neuropteridium intermedium (Schimp.). (After Blanckenhorn. ¼
nat. size.)
An example of another Triassic species is afforded by
Neuropteridium grandifolium Schimp. and Moug., which agrees very
closely with N. validum in the size and shape of the pinnules. The
occurrence in Lower Mesozoic European rocks of fronds hardly
distinguishable from the older southern species may be regarded as
favourable to the view already expressed, that some at least of the
Permo-Carboniferous plants migrated north of the Equator. The
resemblance between the Vosges Triassic species of
Schizoneura[1378] and the examples of this genus recorded from the
Lower Gondwana rocks of India affords additional evidence of a
northern migration.
Our knowledge of the reproductive organs of Neuropteridium is
practically nil. There is no doubt that Zeiller[1379] and
Blanckenhorn[1380] are correct in regarding the Bunter fronds
assigned by Schimper and Mougeot to the genus Crematopteris as
the fertile leaves of Neuropteridium intermedium or some other
species from the same horizon. These fronds bear crowded pinnules
similar to those of Neuropteridium intermedium, N. Voltzii[1381], and
other species, exhibiting on the exposed surface numerous
carbonaceous spots which may be the remains of sporangia.
Cardiopteris.
Schimper[1382] applied this generic name to Lower Carboniferous
fronds of a simple-pinnate habit which had previously been
described as species of Cyclopteris. Cardiopteris frondosa may
serve as a typical example. This species, originally described by
Goeppert as Cyclopteris frondosa (fig. 350), is recorded from Lower
Carboniferous rocks in the Vosges district[1383] in Silesia,
Moravia[1384], and Thuringia[1385]. The pinnules, which are attached in
opposite pairs to a broad rachis, vary in length from 2 to 10 cm. and
have a breadth of 2 to 8 cm.; in manner of attachment and venation
they agree with those of Neuropteridium validum. The venation is
very clearly shown in a drawing of some large pinnules figured by
Stur[1386].
The specimen of Cardiopteris frondosa, a portion of which is
shown in fig. 350 on a slightly reduced scale, was originally figured
by Schimper from an unusually good example in the Strassburg
Museum. Schimper’s drawing hardly does justice to the original
specimen.
A frond bearing rather narrower pinnules, alternately placed on the
rachis, which Fritsch has described as Cardiopteris Hochstetterii var.
franconica from the Culm of Thuringia, bears a close resemblance to
Neuropteridium validum but differs in the entire margin of the
pinnules. An Upper Carboniferous species from Russia described by
Grigoriew[1387] as Neuropteris, cf. cordata var. densineura, represents
another form of similar habit.
Fig. 350. Cardiopteris frondosa (Goepp.). (¾ nat. size. After Schimper.)
Schuster[1388] has recently proposed a new generic name
Ulvopteris for a fragment of a pinna from the Coal-Measures of
Dudweiler in Germany bearing large pinnules, which he compares
with those of Cardiopteris and species of Rhacopteris. The specimen
appears to be indistinguishable from some of those already referred
to as conforming to Neuropteridium, and it is difficult to recognise
any reason for the creation of a new generic name.
We cannot hope to arrive at any satisfactory decision in regard to
the precise affinity between Neuropteridium validum and species
referred to Cardiopteris and other genera so long as portions of
sterile fronds are the only tests at our disposal. It is difficult to
determine whether a specimen consisting of an axis bearing
pinnules represents a large pinna of a bipinnate frond or if it is a
complete pinnate leaf. There is, however, no adequate reason for
supposing that the presumably pinnate fronds from the Gondwana
Land rocks are generically distinct from the Lower Carboniferous
European species Cardiopteris frondosa. Granting the probability
that both genera are Pteridosperms and closely allied to one
another, the two generic names may be retained on the ground of
long usage and in default of satisfactory evidence confirmatory of
generic identity. Cardiopteris would thus stand for a type of frond
characteristic of the Lower Carboniferous strata of Europe, while
Neuropteridium is retained for the Southern species N. validum, and
for others from the Trias of the Vosges.
Aphlebia.
This name was proposed by Presl[1389] for large leaf-like
impressions having a pinnate or pinnatifid form and characterised by
a confused irregular type of venation, or by a fine superficial striation
or wrinkling which simulates veins. Gutbier had previously described
similar fossils as Fucoides, and other authors have described
Aphlebiae as species of Rhacophyllum, Schizopteris, and other
genera[1390]. The term Aphlebia is retained, not as denoting a distinct
genus but (i) as a descriptive name for detached leafy structures
similar to those figured by Presl, which are now recognised as
laminar appendages of the petioles of ferns or fern-like fronds, and
(ii) as an epithet for highly modified pinnules which frequently occur
at the base of the primary pinnae of Pecopteroid and Sphenopteroid
fronds (e.g. Dactylotheca plumosa, fig. 293)[1391].
Modified pinnules, similar in their reduced and deeply dissected
lamina to those represented in fig. 293, are frequently found at the
base of the primary pinnae of Palaeozoic species of Sphenopteris
and other genera of Pteridosperms or ferns, including members of
the Coenopterideae. Potonié[1392] gives a list of various types of
Aphlebiae in his paper on these organs. A striking case has recently
been described by Zeiller in a French Upper Carboniferous species,
Sphenopteris Matheti[1393]. It would seem that the larger examples of
Aphlebiae are more frequently associated with the compound leaves
of Pteridosperms than with those of Ferns[1394].
As examples of the larger types of Aphlebiae reference may be
made to Aphlebia crispa (Gutb.)[1395], which reaches a length of
nearly 60 cm. and has the form of a more or less triangular pinnate
leaf divided into decurrent deeply lobed segments, to a similar
species represented by A. Germari (= Schizopteris lactuca Germ.)
[1396]
which simulates the leaves of endive (Cichorium endivia L.), and
to some large forms figured by Grand’Eury[1397] as species of
Schizopteris.
Aphlebiae such as that figured by Kidston[1398] as Rhacophyllum
crispum, with narrow ultimate segments, might easily be mistaken for
the impressions of an alga.
The term Aphlebia may be applied also to the Cyclopteroid
pinnules on the petioles of some species of Neuropteris,
Odontopteris and Archaeopteris. Goebel[1399] has referred to the
application by Potonié and other authors of the term Aphlebioid to
the pinnules which serve as bud-protecting organs in recent fronds
of Gleichenia (fig. 226, p. 290); he expresses the opinion that it is
superfluous and misleading to make use of a special designation for
structures which are undoubtedly modified pinnules. In the case of
fossils it is, however, convenient to employ the term Aphlebia as a
descriptive name for modified pinnules or stipular structures which
cannot be connected with definite species of fronds. It is clear that
some Aphlebiod leaflets, such as those of Dactylotheca, served as
protective organs for the unexpanded pinnae[1400], and in all
probability the large Aphlebiae served the same purpose as the
fleshy stipules of Angiopteris and Marattia which cover the uncoiled
fronds. The pinnatifid scale-leaves of considerable size (fig. 351)
which occur in the leaf-axils or as ochrea-like stipules on the fronds
of Gunnera (a tropical and subtropical Dicotyledonous genus) bear a
very close resemblance to some Palaeozoic Aphlebiae, e.g.
Aphlebia crispa (Gutb.). The recent and fossil scale-leaves may be
regarded as similar in function as in form; moreover the delicate
coiled fronds of Palaeozoic Pteridosperms or ferns, like those of
some recent flowering plants, may have been kept moist by a
secretion of mucilage. The pinnatifid stipules of Marattia fraxinea (fig.
241, B, p. 317) resemble certain fossil Aphlebiae, and the wrinkled
surface of the recent stipules presents an appearance similar to that
which in some fossil forms has been erroneously described as
veining. It is not improbable that mantle-leaves of such recent ferns
as Polypodium quercifolium (fig. 234, M, p. 303) are comparable with
some fossil Aphlebiae which may have served as humus-collectors
for Palaeozoic epiphytes.
Fig. 351. Scale-leaf of Gunnera manicata. (Slightly reduced. M.S.)
The filiform appendages on the petioles of the recent fern
Hemitelia capensis (fig. 235, p. 304) have often been compared with
the aphlebioid leaflets of fossil fronds.
Potonié who has discussed the nature of Aphlebiae regards them
as vestiges of a once continuous lamina, which formed a winged
border to the branched axes of more primitive forms of fronds. It is
possible that the pinnules between the pinnae on the rachis of
Archaeopteris and the Cyclopteroid leaflets of Neuropteris and
Odontopteris may have the morphological significance attributed to
them by Potonié. In some cases it is probable that the Aphlebiae,
whether vestiges or not, served the purpose of protecting either the
whole frond or individual pinnae. Aphlebiae, though especially
characteristic of Palaeozoic leaves, are occasionally met with in the
form of modified pinnules at the base of the primary pinnae on
Mesozoic ferns, e.g. in Coniopteris hymenophylloides[1401].
In some fern fronds the lowest pinnule of each pinna differs in
shape or size from the normal ultimate segments, but it would be
almost affectation to extend the use of the term Aphlebia to such
pinnules. The Jurassic species Cladophlebis lobifolia (Phill.) is a
case in point[1402]. In this fern, which some authors speak of, without
sufficient reason, as Dicksonia lobifolia[1403], the lowest pinnule is
large and different in shape from the others.
Fig. 352.
A. Sphenopteris obtusiloba. Pinnule. (Enlarged. After Zeiller.)
B, C. S. obtusiloba. (⅞ nat. size. After Zeiller.)
D. Pecopteris arborescens. (Slightly enlarged. After Zeiller.)
E. Sphenopteris furcata (= Diplotmema furcatum). (Slightly enlarged.
After Zeiller.)
Sphenopteris.
Sphenopteris is one of the many generic names which we owe to
Brongniart[1404]. It is the generic designation used for a great number
of Palaeozoic and later fronds, most of which are those of true ferns
while some Palaeozoic species are undoubted Pteridosperms. The
genus, which is purely provisional, includes members of widely
different families possessing pinnules of the same general type, such
as is represented in some recent species of Davallia, Asplenium,
and other ferns.
The fronds of Sphenopteris may be bipinnate, tripinnate, or quadripinnate;
the rachis may be dichotomously branched or the branching may be of the
pinnate type characteristic of most recent ferns. The pinnules are small; they
vary considerably in shape even in a single frond, but the chief characteristics
are: the lobed lamina, contracted and often wedge-shaped at the base (fig.
352), the dichotomously branched veins radiating from the base or given off
from a median rib at an acute angle. The lamina may be divided into a few
bluntly rounded lobes (fig. 352, C) or deeply dissected into linear or cuneate
segments (fig. 352, A, B, E).
Examples of Sphenopteroid leaves have already been described
under the genera Coniopteris, Onychiopsis, Ruffordia, etc. Among
the numerous examples of Sphenopteris species from the
Carboniferous rocks mention may be made of Sphenopteris
obtusiloba Brogn.[1405] (fig. 352, A–C), which occurs in the Middle and
Lower Coal-Measures of Britain[1406]. This type is characterised by
the almost orbicular, oval or triangular pinnules which may reach a
length of 15 mm.; they are occasionally entire, but more usually
divided into 3 to 5 rounded lobes. The forked veins radiate from the
base of the pinnule. The rachis may be dichotomously branched.
Fructification unknown.
The species S. furcata Brongn.[1407], characteristic of the Middle
and Lower Coal-Measures of Britain (fig. 352, E), is referred to under
Stur’s genus Diplotmema[1408] in which it is included by some authors
solely because of the dichotomous habit of branching of the pinnae.
The pinna represented in fig. 353 illustrates a similar type of
pinnule. This species, which is very common in the Calciferous
Sandstone of Scotland, was described by Lindley and Hutton as
Sphenopteris affinis[1409].
The fronds of Sphenopteris affinis were discovered by Mr
Peach[1410] in a fertile condition, but he regarded the reproductive
organs as those of a plant parasitic on the Sphenopteris fronds.
Kidston[1411] substituted Stur’s genus Calymmatotheca for
Sphenopteris on the ground that the sporangia figured by Peach
under the name Staphylopteris Peachii bear a close resemblance to
the organs which Stur described as valves of an indusium in his
species Calymmatotheca Stangeri[1412]. An examination of Stur’s
specimens by Miss Benson[1413] and by Prof. Oliver and Dr Scott has
confirmed Stur’s interpretation of the appendages at the tips of the
fertile pinnae as valves of an indusial or cupular structure. The
superficially similar bodies on the fertile pinnae of S. affinis are
however true sporangia, and cannot legitimately be included in the
genus Calymmatotheca as described by Stur. For this reason Miss
Benson institutes a new genus Telangium, the type-species of which,
T. Scotti from the Lower Coal-Measures of Lancashire, is based on
petrified material. The Scotch species Sphenopteris affinis (=
Calymmatotheca affinis of Kidston) is also transferred to Telangium;
the sporangia are considered by Miss Benson to be microsporangia.
This with other species is no doubt correctly included in the
Pteridosperms. A complete frond of Sphenopteris affinis, showing a
regular dichotomy of the main axes, is represented by an admirable
drawing in Hugh Miller’s Testimony of the Rocks[1414].
Fig. 353. Sphenopteris affinis, Lind. and Hutt. From the Calciferous
Sandstone of Burdiehouse (Scotland). (Sedgwick Museum,
Cambridge.) M.S.
Some of the Palaeozoic species of Sphenopteris probably
represent the fronds of true ferns, but others are known to have been
borne by Pteridosperms. S. Hoeninghausi (fig. 290, C, p. 399) is the
foliage of Lyginodendron, and Scott[1415] speaks of three species, S.
dissecta, S. elegans, and S. Linkii as the leaves of Heterangium.
Grand’Eury[1416] has recorded the occurrence in French Coal-
Measures of seeds in association with other Sphenopteroid fronds.
Mariopteris, Diplotmema, Palmatopteris.
The discovery of sporangia on the fronds of several Palaeozoic
species of Sphenopteris and Pecopteris has led to the institution of
new generic names, which indicate an advance in knowledge
beyond the stage implied by the use of those provisional
designations based solely on the form and venation of the pinnules.
Other names have been created by authors in place of Sphenopteris
and Pecopteris on the ground that a striking feature in the mode of
branching of fronds is sufficiently important to justify generic
recognition even in the absence of fertile specimens. As examples of
designations based primarily on the branch-system of compound
leaves, the genera Mariopteris, Diplotmema, and Palmatopteris may
be briefly considered (fig. 354 A–C). Dr Kidston[1417] is of opinion that
the creation of new genera for purely vegetative characters of fronds
is of no real advantage, and he prefers to retain the older provisional
names for species known only in the sterile condition. On the other
hand, if we are sufficiently familiar with specimens large enough to
enable us to recognise a well-defined morphological character, it
may serve a useful purpose to employ a generic designation for
features which may have a phylogenetic value. A comparative
examination of Palaeozoic, Mesozoic, and recent compound fronds,
including both Pteridosperms and true ferns, brings to light certain
distinguishing features characteristic of the older types which, as
Potonié maintains[1418], point to the derivation of the pinnate habit
from a primitive dichotomous system of branching. For a more
complete discussion of this question reference should be made to
Potonié’s suggestive papers. Among recent ferns Matonia and
Dipteris, two survivals from the past, afford instances of fronds with a
branching system of the dichotomous type.
Similarly, in Gleichenia, Lygodium, and more rarely in species of
Polypodiaceae (e.g. Davallia aculeata, fig. 232) dichotomy is a
striking feature of the fronds. In the great majority of recent ferns the
fronds have assumed a pinnate habit. Among Palaeozoic fern-like
fronds dichotomous branching of the main rachis and of the pinnae
is much more common. Potonié draws attention to several other
features which distinguish Palaeozoic fronds from the majority of
later species: the frequent occurrence of pinnules borne directly on
the main rachis (fig. 354, D), and of modified pinnules or Aphlebiae
on the rachis and petiole, are characters to which he attributes an
evolutionary significance. The main point is that a comparative
examination of leaf-form affords evidence in favour of the view that
the modern type of frond, with its naked rachis bearing two rows of
pinnae, has been derived from a less specialised type in which the
distinction between the parts of the leaf is much less evident. The
primitive leaf was probably a dichotomously branched axis provided
with a continuous lamina which eventually became broken up into
separate lobes or pinnules.
As the dichotomy of the frond became less regular, a pinnate habit
was acquired, as is clearly seen in many Palaeozoic types which
constitute connecting links between forked and pinnate fronds (fig.
354, D). The Aphlebiae may be remnants of the once-continuous
lamina on the petiole, and the normal pinnules borne on the rachis
may be regarded as the attributes of fronds in which the division of
physiological labour had not reached the stage which characterises
the leaves of recent ferns.
Mariopteris.
This name, which is due to Zeiller[1419], is applied by him to
Palaeozoic fronds characterised by a double bifurcation of the rachis
of the primary pinnae. Mariopteris muricata (= Pecopteris muricata
Schloth.) may be taken as the type of the genus. This species is
common in the Lower and Middle Coal-Measures of Britain and rare
in the Upper Coal-Measures[1420]. It is described by Kidston[1421] as
one of the most polymorphic and widely distributed Coal-Measure
species. The pinnules as seen in fig. 364, B, are of the
Sphenopteroid type. No fertile specimens are known, but it is
significant that Grand’Eury[1422] has recorded the association of
Mariopteris muricata and seeds.
The main rachis gives off alternate naked branches, each of which
bifurcates at its apex into two short naked axes, and these are again
forked, the ultimate branches having the form of bipinnate pinnae
provided with large Sphenopteroid pinnules (fig. 354, B). Zeiller
includes in Mariopteris some species which Stur[1423] referred to his
genus Diplotmema. Possibly some of the Palaeozoic fronds with a
zigzag rachis may have been climbers like Lygodium.
Fig. 354.
A. Palmatopteris.
B. Mariopteris. (A, B, after Potonié.)
C. Diplotmema Zeilleri, Stur. (After Zeiller.)
C′. D. Zeilleri. Pinnule. (× 3. After Zeiller.)
D. Neuropteris macrophylla. (British Museum.)
D′. N. macrophylla. Pinnule. (Slightly enlarged. After Kidston.)
E. N. heterophylla. Pinnule. (Slightly enlarged. After Zeiller.)
F. N. Scheuchzeri. (Slightly reduced. After Kidston.)
G. Alloiopteris Essinghii. (Enlarged. After Potonié.)
Diplotmema.
This generic name is employed by Zeiller[1424] and other authors in
a more restricted sense than that in which it was originally used by
Stur. The Upper Carboniferous species Sphenopteris furcata
Brongn. (fig. 352, E) may serve as the type. This species occurs in
the Middle and Lower Coal-Measures of Britain[1425]. The main rachis
gives off branches as in Mariopteris, but in Diplotmema each naked
lateral branch is forked at its apex into two opposite pinnae bearing
deeply dissected Sphenopteroid pinnules. Zeiller[1426] and Stur have
recorded fertile specimens of Diplotmema, but in no case have
actual sporangia been discovered. In the species Diplotmema Zeilleri
Stur (fig. 354, C, C′) two Aphlebiae occur at the base of each
secondary axis[1427]. It has been pointed out by Potonié that in
Diplotmema furcatum the equal dichotomy of the lateral branches is
not characteristic of the frond as a whole. In the case of branches
higher on the rachis the dichotomy becomes unequal and the forked
axis is gradually replaced by a simple pinna (fig. 354, A). For this
type of frond, Potonié proposed the generic name Palmatopteris in
place of Diplotmema, which he discards. The long comparatively
slender rachis of P. furcata suggests comparison with the liane
species of Lygodium[1428].
Fig. 355.
A. Cephalotheca mirabilis, Nath. Fertile pinnae. (Partially restored. After
Nathorst.)
B. C. mirabilis. Sterile pinnule. Nat. size. (After Nathorst.)
Cephalotheca.
This genus was proposed by Nathorst[1429] for some peculiar
bipinnate fertile fronds from the Upper Devonian rocks of Bear
Island. The pinnae bear slender forked ultimate segments
represented by a few detached fragments (fig. 355, B), associated
with the rachises. The fertile pinnae are given off in opposite pairs
from the main axis over which they are concrescent (fig. 355, A). A
mop-like cluster of sporangia is borne on the lower surface and close
to the base of a fertile pinna: the exannulate sporangia are
compared with those of Scolecopteris. Nathorst compares
Cephalotheca with a Belgian species of Upper Devonian age
described by Crépin[1430] as Rhacophyton condrusorum and by
Gilkinet[1431] as Sphenopteris condrusorum. A similar fossil is also
described by Baily[1432] as Filicites lineatus from the Kitorkan Grits of
Ireland.
The position of Cephalotheca cannot be definitely determined from
the available data, but it is more probable that it was a seed-bearing
Pteridosperm and not a true fern. Zeiller[1433] has recently expressed
the same opinion.
Thinnfeldia.
The genus Thinnfeldia, founded by Ettingshausen in 1852[1434] on
some Hungarian Liassic specimens, though frequently included in
the Filicales, cannot be said to occupy that position by virtue of any
well-authenticated filicinean features. It is by no means improbable
that many of the species referred to this genus are closely allied to
Palaeozoic Pteridosperms.
Thinnfeldia may be briefly defined as follows:
Fronds simple and pinnatifid, pinnate or bipinnate: rachis broad and
occasionally dichotomously branched. Pinnules often fleshy or coriaceous;
broadly linear, entire or lobed, provided with a midrib from which simple or
forked secondary veins are given off at an acute angle: or the laminae may be
short and broad without a midrib and traversed by several slightly divergent and
forked veins.
No satisfactory evidence of reproductive organs has so far been adduced.
The genus is chiefly characteristic of Upper Triassic, Rhaetic, and
Jurassic floras, though it was in all probability represented in
Permian floras. Several species, many of which are valueless, are
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