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Fig. 30. 1. Rill-mark (after Williamson). 2. Trail made by a seaweed
dragged along a soft plaster of Paris surface (after Nathorst). 3. Tracks
made by Goniada maculata, a Polychaet (after Nathorst). 4. Burrow of
an insect. 4a. Section of the gallery (after Zeiller).
There is, however, another category of impressions and casts of
common occurrence in sedimentary rocks which requires a brief
notice. Very many of the fossil algae described in text-books and
palaeobotanical memoirs have been shown to be of animal origin,
and to be merely the casts of tracks and burrows. A few examples
will best serve to illustrate the identity of many of the fossils referred
to algae with animal trails and with impressions produced by
inorganic agency.
Dr Nathorst of Stockholm has done more than any other worker to
demonstrate the true nature of many of the species of Chondrites,
Cruziana, Spirophyton, Eophyton, and numerous other genera. In
1867 there were discovered in certain Cambrian beds of
Vestrogothia, long convex and furrowed structures in sandstone
rocks which were described as the remains of some comparatively
highly organised plant, and described under the generic name
Eophyton[226]. By many authors these fossils have been referred to
algae, but Nathorst has shown that the frond of an alga trailed along
the surface of soft plaster of Paris produces a finely furrowed groove
(fig. 30, 2) which would afford a cast similar to that of Eophyton. The
same author has also adduced good reasons for believing that the
Eophytons of Cambrian rocks may represent the trails made by the
tentacles of a Medusa having a habit similar to that of Polydonia
frondosa Ag. Impressions of Medusae have been described by
Nathorst from the beds in which Eophyton occurs; and the
specimens in the Stockholm Museum afford a remarkable instance
of the rare preservation of a soft-bodied organism[227]. By allowing
various animals to crawl over a soft-prepared surface it is possible to
obtain moulds and casts which suggest in a striking manner the
branched thallus of an alga. The tracks of the Polychaet, Goniada
maculata Örstd.[228], one of the Glyceridae, are always branched and
very algal-like in form (fig. 30, 3). Many of the so-called fossil algae
are undoubtedly mere tracks or trails of this type. In the fossil-plant
gallery of the British Museum there are several specimens of small
branched casts, clearly marked as whitish fossils on a dark grey rock
of Lower Eocene age from Bognor; these were described by Mantell
and Brongniart[229] as an alga, but there is little doubt of their being of
the same category as the track shown in fig. 30, 3.
FOSSILS SIMULATING ALGAE.
A. DIATOMACEAE (Bacillariaceae).
This family occupies a somewhat isolated position among the
algae, and is best considered as a distinct subdivision rather than as
a family of the Phaeophyceae or Brown algae, with which it
possesses as a common characteristic a brown-colouring matter.
Single-celled plants consisting of a simple protoplasmic body
containing a nucleus and brown colouring matter (diatomin)
associated with the chlorophyll. The cell-wall is in the form of two
halves, known as valves, which fit into one another like the two
portions of a pill-box. The cell-wall contains a large amount of silica,
and the siliceous cases of the diatoms are commonly spoken of as
the valves of the individual, or the frustules. Diatoms exhibit a
characteristic creeping movement, and are reproduced by division,
also by the development of spores in various forms[251].
The recent members of the family have an exceedingly wide
distribution, occurring both in freshwater and in the sea. Owing to the
lightness of the frustules, they are frequently carried along in the air,
and atmospheric dust falling on ships at sea has been found to
contain large numbers of diatoms[252]. The siliceous valves are
abundant in guano deposits, and they have been found also in
association with volcanic material. Diatomaceous deposits are now
being formed in the Yellowstone Park district; “they cover many
square miles in the vicinity of active or extinct hot spring vents of the
park, and are often three feet, four feet, and sometimes five to six
feet thick[253].” The gradual accumulation of the siliceous tests on the
floor of a fresh-water lake results in the formation of a sediment
consisting in part of pure silica. Such deposits, often spoken of as
kieselguhr or diatomite, and used as a polishing material, occur in
many parts of Britain, marking the sites of dried-up pools or lakes. At
the northern end of the island of Skye there occurs an unusually
pure deposit of diatomite overlain by peat and turf, and extending
over an area of fifty-eight square miles. Many of the individuals in
this deposit were in all probability carried into the lake by running
water, while others lived in the lake and after death their tests
contributed to the siliceous deposit[254]. The late Dr Ehrenberg
published numerous papers on diatomaceous deposits in different
parts of the world, and in his great work, Zur Mikrogeologie[255], he
gave numerous and beautifully executed illustrations of such
siliceous accumulations. In many of the samples he figures one sees
fragments of plant tissues, spores of conifers and ferns, associated
with the diatom tests. The occurrence of the pollen grains of
coniferous trees in lacustrine and marine deposits is not surprising in
view of their abundance in Lake Constance and other lakes. It is
stated that the pollen of conifers in the Norwegian fiords plays an
important part in the nourishment of the Rhizopod Saccamina[256].
DIATOMACEOUS OOZE.
(fig. 32, C.) Several species have been figured by Heer from beds of
Muschelkalk, Keuper and Rhaetic age. He describes the wall as
thick and firm (fig. 32, C. ii.) and probably composed of silica, with a
hollow interior. The specimen shown in fig. 32, C. was found in the
Rhaetic beds, and named by Heer Bactryllium deplanatum; it has a
length of 4·5 mm.; the surface is transversely striated and traversed
by a single longitudinal groove. Stefani[269] has given reasons in
favour of removing Bactryllium from the plant to the animal kingdom;
he points out that the specimens are too large for diatoms, and
moreover that they are asymmetrical in form and possessed a
calcareous and not a siliceous shell. He would place the fossil
among the Pteropods, comparing it with such genera as Cuvierina
and Hyalaea. In view of Stefani’s opinion we cannot attach any
importance to this supposed diatom, especially as it has generally
been regarded as at best but an unsatisfactory genus.
a. Siphoneae.
Thallus consisting of simple or branched cells very rarely divided
by septa, and containing many nuclei. In certain genera the
branches form a pseudoparenchymatous tissue by their repeated
branching, and as a result of the intimate felting together of the
branched cells. Reproduction is effected either by the conjugation of
similar gametes or by the fertilisation of an egg-cell.
Vaucheria and Botrydium are two well-known British genera of this
order, but most of the recent representatives live in tropical and
subtropical seas. The most striking characteristic feature of this
division of the Chlorophyceae is the fact that the thallus of a
siphoneous alga consists of an unseptate coenocyte; the plant may
be extremely small and simple, or it may reach a length of several
inches, but in all cases the body does not consist of more than one
cell or coenocyte.
From a palaeontological standpoint the Siphoneae are of
exceptional interest. It is impossible to do more than refer to a few of
the living and fossil genera. There are numerous fossil
representatives already known, and there can be little doubt that
further research would be productive of valuable results.
As examples of the order, a few genera may be described
belonging to the three families Caulerpaceae, Codiaceae, and
Dasycladaceae.
α. Caulerpaceae.
Thallus unseptate, showing an extraordinary variation in the
external differentiation of the plant-body. Reproduction is effected by
means of detached portions of the parent plant.
The genus Caulerpa, represented by a few species in the
Mediterranean and by many tropical forms, has already been alluded
to as a striking example of a plant which appears under a great
many different forms[271]. As a recent writer has said, “Nature seems
to have shown in this genus the utmost possibilities of the
siphoneous thallus[272],” Fragments of coniferous twigs, the tracks
and burrows of various animals and other objects have been
described by several authors as fossil species of Caulerpa. As an
illustration of the identification of a very doubtful fossil as a species
of Caulerpites, reference may be made to such a form as C.
cactoides Göpp.[273] from Silurian and Cambrian rocks. There are
several examples of this fossil in the Brussels Museum which
probably owe their origin to some burrowing animal, and may be
compared with Zeiller’s figures of the tunnels made by the mole-
cricket (fig. 30, 4)[274].
Mr Murray, of the British Museum, has recently described what he
regards as a trustworthy example of a fossil Caulerpa from the
Kimeridge Clay near Weymouth[275]. Specimens of the fossil were
first figured in a book on the geology of the Dorset coast as casts of
an equisetaceous plant[276].
To this fossil Murray has assigned the name Caulerpa Carruthersi,
and given to it a scientific diagnosis. The best specimens have the
form of a slender central axis, giving off at fairly regular intervals
whorls of short and somewhat clavate branches; they bear a
superficial resemblance to such a recent species as Caulerpa
cactoides Ag. An examination of several examples of this fossil leads
me to express the opinion that there is not sufficient reason for
assigning to them the name of a recent genus of algae[277]. To use
the generic name of a recent plant without following the common
custom of adding on the termination “ites” (i.e. Caulerpites) is as a
general rule to be avoided in dealing with fossil forms; and there are,
I believe, no satisfactory grounds for referring to these fossils as
trustworthy examples of a Mesozoic alga.
In the present case I am disposed to regard the Caulerpa-like
casts as of animal rather than plant origin. The clavate branches
have the form of very deep moulds in the hard brown rock which
have been filled in with blue mud. It is hardly conceivable that the
branches of a soft watery plant such as Caulerpa could leave more
than a faint impression on an old sea-floor. The specimens occur in
different positions in the matrix of the rock and they are not confined
to the lines of bedding; in none of the examples is there any trace of
carbonaceous matter in association with the deep moulds. On the
whole, then, this Kimeridge fossil cannot, I believe, be accepted as
an authentic example of a Mesozoic Caulerpa.
It is not improbable that some of the supposed fossil algae may be
casts of egg-cases or spawn-clusters of animals. In Ellis’ Natural
History of the Corallines[278] there is a drawing representing a number
of disc-like ovaries attached to a tough ligament, and referred to the
mollusc Buccinum, which bears a certain resemblance to the
Weymouth fossil. A similar body is figured by Fuchs[279] in an
important memoir on supposed fossil algae.
It is not suggested that the Caulerpa Carruthersi of Murray should
be regarded as the cast of some molluscan egg-case attached to a
slender axis, but it is important to bear in mind the possibility of
matching such extremely doubtful fossils with other organic bodies
than the thallus of a Caulerpa. In an example of an egg-case in the
Cambridge Zoological Museum, referred to a species of Pyrula,
there is a hard, long and slender axis, bearing a series of
semicircular chambers divided into radial compartments. The whole
is hard and horny and might well be preserved as a fossil.
β. Codiaceae.
The members of this Order present a considerable diversity of
form as regards the shape of the plant-body; the thallus of some
species is encrusted with carbonate of lime. The order is widely
distributed in tropical and temperate seas.
Among the recent genera Penicillus and Codium may be chosen
as important types from the point of view of fossil representatives.
Codium.
The thallus of Codium consists of a spongy mass of tubular cell-
branches which are differentiated into two fairly distinct regions, an
outer peripheral layer in which the branches have long club-shaped
terminations, and an inner region consisting of loosely interwoven
filaments.
Codium Bursa L. and C. tomentosum Huds. are two well-known
British species, the former presents the appearance of a spongy ball
of cells, and in the latter the thallus is divided up into dichotomously
forked branches[280]. In this genus the thallus is not encrusted with
carbonate of lime, at least in recent species.
Penicillus.
The recent genus Penicillus is one of those algae formerly
included among animals. Fig. 33, O, has been copied from a drawing
of a species of Penicillus given by Lamouroux[283] under the generic
name of Nesea in his treatise on the genera of Polyps published in
1821. He describes the genus as a brush-like Polyp with a simple
stem.
The thallus consists of a stout stem terminating in a brush-like tuft
of fine dichotomously-branched filaments. The apical branches are
divided by regular constrictions into short oval or rod-like segments
which may be encrusted with carbonate of lime. A few of the
segments from the terminal tuft of a recent Penicillus are shown in
fig. 35, E. Each of these calcareous segments has the form of an
oval shell perforated at each end, and the wall is pierced by
numerous fine canals. Penicillus is represented by about 10 recent
species, which with one exception live in tropical seas.
The recognition of Penicillus, or a very similar type, in a fossil
condition is due to Munier-Chalmas[284]. This keen observer has
rendered great service to palaeobotany by directing attention to the
calcareous algae in the Paris basin beds, and by proving that many
of the fossils from these Tertiary deposits have been erroneously
included by previous writers among the Foraminifera[285]. It is greatly
to be desired that Prof. Munier-Chalmas may soon publish a
monograph on the fossil Siphoneous forms of which he possesses a
unique collection.