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Senckenberg-Poster

TheEvolutionofAnimals
Manfred Grasshoff and Michael Gudo
with cooperation of S. Hilsberg, W. Oschmann and J. Scholz

One of the most important aspects in word in English scientific papers and From here the evolutionary pathways
biological sciences is the investigation text books). originate and branch off. Not all
of long-term changes in nature. It is branchings of the animal kingdom can
commonly accepted that the universe, This approach provides new visions of be presented in the design, we decided
the planetary system, and all the or- biological research. Common knowl- to include those leading to well-known
ganisms living on earth, are the result edge of biological investigations can animals or to some of specific zoologi-
of a process of continuous modifica- be integrated, but in addition, this ap- cal interest. On these branches you
tion and development. The architec- proach provides answers for the ques- find the most important steps in the
ture of recent organisms provides a tion: is a gradual change of one bau- evolutionary transformation of organ-
key for the understanding and recon- plan into another possible? For exam- isms represented by technical draw-
struction of their history through the ple: can a dinosaur evolve into a bird? ings. At the end of each pathway a
billions of years, of the process called This question can be answered with a most recent or a known fossil repre-
evolution. ‘yes’. We can show that the dinosaur sentative is shown in a naturalistic
bauplan supports the possibility of form. These stand in contrast to the
Evolution is the gradual change of or- this evolutionary transformation. An- model-like, hypothetical and technical
ganisms through generations and other question: Can a dinosaur evolve drawings. All extant animals are
time. We cannot directely observe into a mammal? This has to be an- equally remote of their point of origin.
evolution in nature. So we have to re- swered with a ‘no’. We can show that Each animal by itself has attained its
construct how organisms could have the bauplan does not support this kind own stage of sophistication in the evo-
changed and how their variety of transformation. lutionary process. We no longer share
emerged under anatomical and struc- the traditional anthropocentric view
tural-functional constraints. For any This way of reasoning, its philosophi- of the world, in which man has taken
evolutionary transformation it is con- cal background, and the results on the the position of the top of a tree grow-
sidered that the process has to be con- reconstruction of evolutionary path- ing from so-called lower to so-called
tinuous. Organisms are not engines, ways, were developed by a group of sci- higher animals.
which can be stopped for rebuilding. entists in the Senckenberg-Museum,
Organismic evolution can be likened Frankfurt, in cooperation with several
to a gradual change in the running en- colleagues. The new approach has
gine. been called "The Frankfurt Evolution- CONTENTS
ary Theory" and "Engineering Mor-
The origin of life 1
Evolutionary research should not only phology" (in German: Konstruktions- The explosion of bacterial life 2
consider obvious similiarities. The ar- Morphologie). The origin of eucaryotes 3
chitectural elements by which an or- The diversity of Eukaryotes 4
ganism is designed are much more im- The results are summarized in a The first of multicellular animals 5
portant. In particular, this means the graphic design. It is published as a Gelatinous fibrous body mass 7
mechanical properties of tissues, their poster, and presented in the museum The body-cavities develop 9
arrangement and their functional con- as a wall-hanging in room 206 and in a The worm-coelomates 9
nections. (In German this complex of modified form in room 104. The top The chordates 12
structure and function is called ‘Kon- part of the poster shows the formation Peribranchial-chambers 13
struktion’ or ‘Bauplan’; the latter term of the earth and the development of Vertebral column and tooth 14
the first organisms. The most ances- Jaws, skull and bones 14
is sometimes used as a German loan-
tral animals are located in the center. Evolution in teching courses 16

Evolution of Animals 1
Early Evolution of Earth and Life

The origin of life


Red Planet

About 5 billion years Archaean Atmospheres


Meteorites and Asteroids
ago accumulating and forming Planet Earth CO2 - Atmosphere
Brownish-red Planet
fusing meteorites and
asteroids formed the Organisms produce O2
and control the Greenhouse Effect
hot red planet earth.
After an extensive about 4 billion years ago Bacteria

cooling the rocks crys-


tallized and internal Protocells Biofilms

Archaea
chemical reactions ex- about 3,8 billion years ago

haled various gases Motiloids


(Proto-Eucarya)
through volcanoes, Endosymbioses
about 2 billion years ago
forming the archean
Eucaryotes:
atmospheres which nuceli, fibres,
mitochondria, cilia
chloroplasts
contained water but no
free oxygen. The water
condensed, and a rain
lasting millions of years
filled rivers, lakes, and
Just like soap-bubbles, they are able to for the bacteria. The stabilizing layer
the oceans. The world of water, called
fuse, to repair small holes and to se- is the so called murein-layer.
hydrosphere was created. About 4 bil-
quester small bubbles. In the lipid-
lion years ago physical and chemical
membrane protein-molecules are in- This also was the time when the ge-
conditions in the water permitted the
corporated, which were able to trans- netic substance desoxyribonucleic-
formation of the so-called organic
port matter from the external medium acid (DNA) originated. The DNA car-
chemical compounds. This fluid has
into the inner cavity and vice versa. ries the information for the distinct
been called ‘Ursuppe’, which means
Such membrane-enclosed structures enzymes and molecules, which are im-
‘primeval soup’. These organic com-
were the result of chemical-evolution. portant during metabolism. In most
pounds were formed at various sites of
They indeed had some properties of cases the genetic information is only a
the oceans and their shores, mainly in-
living beings: metabolism, reproduc- single ribbon, ribonucleic-acid (RNA),
volving mineral surfaces, such as rock
tion and merging. We call them pro- and it merges in some cases to a dou-
or clay. In such places solutions be-
tocells. ble ribbon, DNA. Only the eucarya
came concentrated and chemical re-
enclose these genetic information in
actions took place in favourable condi-
The selective permeability of the an additional membrane. They formed
tions. It was the phase of chemical-
membranes provided two options for a nucleus which hold the genetic infor-
evolution on earth.
metabolism. The first option was to mation in a save place.
sequester membrane-parts (vesicles)
A decisive step in the chemical-evolu-
into the inner cavity, in order to trans-
tion, was the origin of lipids. Lipids are
port substances from the outside into
molecules which arrange theirselves in
the inner space. This option allows to
a flexible double-layer, forming mem-
produce organic substances in the in-
branes. Such membranes are barriers
ner of the cell. This is typical for the
between solutions of different kinds.
motiloids (proto-eucarya). The
Such flexible membranes enclose
second option was to produce a slime
small reactive cavities and therefore
and to deposit it on the outer mem-
they are an indispensible element of
brane surface. These substances stabi-
living beings.
lized the outer shape. This is typical

Evolution of Animals 2
A certain group of bacteria was of par- transform Earth into a planet similar
The explosion of bacterial life ticular importance for the evolution of to Venus, several hundreds of degrees
the planet earth, the Cyanobacteria (in hot. Only the metabolism of living be-
A typical bacterium is not much larger
former times called the blue-green al- ings continuously recycles both gases,
than a thousandth of a millimeter.
gae). They contain the enzyme chloro- oxygen and methane. Therefore the
Bacteria do not have a nucleus and
phyll which is capable of producing concentration of both these gases is
their small sizes result in simple
sugar out of water, carbon dioxide and contained in a steady-state equilib-
shapes: rods, spheres or tubes. But it
energy from the sunlight. Hereby oxy- rium (homeostasis).
would be by no means adequate to un-
gen is released into the environment.
derstand these simple shapes as primi-
Bacteria have modified the surface of
tive organisms. From these simple
3.5 billion years ago this oxygen re- the planet earth to a considerable ex-
shapes no inference can be made to
acted directly with other elements. tent, and they have provided an envi-
their metabolism capacities. Bacteria
Iron was oxidized into iron oxid ronment suitable for other forms of
live by the use of distinct matter and
(Fe2O3). In water by help of bacteria, life. Bacteria control the green-
exotic elements, such as sulfur, sulfur-
massive stromatolite-like sediments house-effect and the overall impact
hydrogen, copper, iron and even the
were accumulated, called the Banded of solar radiation through the media-
radioactive uranium.
Iron Formation (BIF). When finally all tion of the reflexion capacities (al-
the iron was oxidized, free oxygen bedo) of earth‘s surface.
Furthermore bacteria can live in ex-
could accumulate in the hydrosphere
treme environments, such as the gey-
and the atmosphere. The atmosphere, lithosphere, and hy-
sers in Yellowstone (USA), in New
drosphere are influenced and partly
Zealand in 150 degree hot sulfuric-acid
Oxygen is a poison to cells, because it generated by organisms. All together
and they are also found in more than
oxidizes organic substances. About 2 constitute the biosphere, a realm of
four kilometers depth in the earth‘s
billion years ago, oxygen caused the living matter. Up till now everything is
crust. Some of these microorganisms
largest mass extinction in earth‘s his- based on the metabolic activity of bac-
have been interpreted as living fossils
tory. Even today many microorganisms teria.
and therefore they were called archaic
are sensitive to oxygen, such as the Ar-
bacteria, Archaebacteria or Archaea.
chaea and a number of anaerobic bac-
Today it is well known that the diver-
teria. Life was able to persist, since at
sity of the Archaea is as complex as
the same time some bacteria were ca-
that of the other bacteria. Accordingly
pable of using oxygen for energy sup-
life is divided into three domains: the
ply. Finally an equilibrium of produc-
Bacteria, the Archaea (both called
tion and destruction of oxygen was at-
procaryotes) and the Eucarya (Eu-
tained.
karyotes).
Planet earth is the only planet in our
More than 90 % of all procaryotes live
solar system, which has free oxygen in
on rock surfaces. Here their slime-
its atmosphere. The UV-rays induce
layer develops a tendency to form the
the production of ozone (O3) in a high
so-called biofilms. In such biofilms
stratum of the atmosphere. In return
cycles of production and destruction
the ozone layer reduced the amount of
of organic substances are established.
UV-rays reaching the earth‘s surface.
When particles were added into this
slime or when the cells actively pro- The atmospheres of the planets Mars
duced carbonate matter, a layered sed- and Venus mainly consist of carbon di-
iment developed, a so-called stroma- oxide. In contrast the atmosphere of
tolite. Stromatolites exist in oceans, the earth mainly consists of nitrogen,
lakes and rivers. They are easily pre- oxygen and methane and a small
severd in the fossil record. The first amount of carbon dioxide. On an inan-
stromatolites are more than 3.5 billion imate planet earth methane would
years old. They have been found in the have been oxidized into carbon diox-
West Australian desert – at a place ide and accumulated in the atmo-
which is called ‘North Pole’. sphere. This greenhouse-gas would

Evolution of Animals 3
The second endosymbiosis opened up
a separate pathway, the evolution of
plants. With the additional incorpora-
tion of cyanobacteria, which are able
to conduct photosynthesis, these cells
had created their own supply for en-
ergy and organic compound. The
chloroplasts produce sugars out of
water and carbon dioxide, which are
the basis for production of distinct
building materials.

The eucaryotes do not only have dis-


tinct cell organelles, but also nuclei,
in which the genetic substance (the
DNA) is located. The DNA became
enveloped by a membrane and was in
However, when bacteria were not im- this way separated from the mechani-
The origin of eucaryotes cal system of the cell.
mediately digested by the motiloid,
The mobile protocells, which we call but rather continued to live, the moti-
loid used the substances produced by Cilia had already developed in this
motiloids, lived in the shadow for
these bacteria and a close interaction early phase of evolution. They are
long time, compared to the rich bacte-
began: a symbiosis. As one partner movable thread-like structures, whose
rial life. In these protocells a cell ske-
lives inside the other, this relation is beat enabled an unicellular organism
leton developed, made up of various
called an endosymbiosis. Such endo- to swim about in water. Cilia are pre-
protein molecules forming rigid rods
symbioses can be observed today in served throughout the whole of evolu-
(the microtubuli) and fibers. Fibers
some fungi and algae. tionary history. For example, ciliated
with tensile strength constructed
cells are incorporated into the epithe-
cords and grids throughout the cell
Two endosymbioses played a crucial lia of many parts of recent organisms,
and under the surface of the mem-
role for further evolution. The first en- such as: into lung, central nervous sys-
brane.
dosymbisis was one in which a kind of tem, gut tissues, or coelom cavities.
Some of the molecules in the fibre grid bacteria were incorporated that could
gain their energy by combustion of or- Cells with one or numerous nuclei,
are able to glide along each other; they
ganic substances with oxygen. In this various fibers, one or several cilia, and
can contract. Since they are anchored
process, energy was stored in special cell organelles, are called eucaryotic
in the fiber grid they deform the entire
compounds, called ATP. This process cells. They are the cells of plants, ani-
cell and generate motion. Shifting and
is called respiration. Such bacteria mals and many other organisms, col-
gliding consumes energy. Energy is
transformed into permanent parts of lectively called Eucaryotes. They in-
supplied by special compounds, in
the cell, the so-called mitochondria. habit our world today and they all live
which it is stored in a chemical form.
Energy turnover increased dramati- at the expense of bacteria.
This chemical energy is transformed
into mechanical work, which creates cally in these cells and new options
heat. Fibers with tensile strength as were opened for further evolution.
well as contractile fibers play a major However, the mitochondria cannot
role in all organismic cells. stop their activity, and they require a
permanent supply of food and oxygen.
The archaean motiloids were able to Organisms with mitochondria have to
flow around bacteria and other parti- eat and to breathe continuously.
cles, to incorporate them into their
body, and to digest and use them as
energy suppliers.

Evolution of Animals 4
isms, decompose them outside their
The diversity of eukaryotes: body and feed on the degradation
Precambrian explosion of life products, which are taken up in the
form of small drops or as single large
A remarkable array of evolutionary de-
molecules.
velopments roots in this early level of
eukaryotic cells. We may speak of a
Protoctista is a collective term for
Precambrian explosion of life which
quite a number of very different struc-
took place about 2 billion years ago.
tural types of organisms. All of these
(For comparison, the so-called Cam-
divide their cells concentrically and
brian explosion of life, about 600 mil-
completely by incision. In some types
lion years ago, is in reality the begin-
the cells remain in connection, glued
ning of the rich fossil record). The var-
together by tough gelatinous sub-
ious types of eukaryotes differ from
stances. Among such types are the
each other in the contents of their
large kelps of the sea, the red and the
cells, the way in which they gain their
green algae. But many other Protoc-
food, and how they divide and grow.
tista are unicellular. They show a large
variety and represent a huge biomass
Plants, as we know them today, owe
as a whole. Many have chloroplasts
their existence to the second endo-
and are called unicellular plants; oth-
symbiosis shown in our drawing, in
ers ingest particles, they are called
which cyanobacteria were additionally
Protozoa, called unicellular animals.
incorporated into eucaryotic cells.
Three of them are shown in our
The cyanobacteria remained almost
poster, a Paramaecium, a flagellate with
unchanged. They became the well-
chloroplasts, and a foraminiferan.
known green chloroplasts of the
plant cells. All of these eucaryotes
The microscopically small protoctists,
with chloroplasts are called plants, al-
together with fungi and bacteria, are
though they comprise a variety of
almost omnipresent in water and on
types. Photosynthesis with the aid of
land. On surfaces of various kinds,
chlorophyll has never stopped, and to-
they live in thin layers, the so-called
day plants still produce most of the or-
biofilms, in which they interact by
ganic material.
chemical turnover and growth. Al-
ready in archaean times, biofilms
Plants in a stricter sense have a special
played a major role in evolutionary
kind of growth. The cell-nucleus di-
processes.
vides into two, the cell enlarges, and
produces internally a sieve-like plate
The land-plants with their incomplete
with holes, so that on either side a nu-
cell-division are exceptional among all
cleus is placed. The terrestrial plants
eucaryotes, (as well as the animals with
and among the algae the Charo-
their specifically structured body).
phyceae are built according to this
The conspicuousness of plants and an-
type; all of their cells are imperfectly
imals makes them so important for us,
divided. The sugar produced by assim-
but we should not forget: the huge
ilation is chemically transformed into
masses of unicellular organisms, fungi,
cellulose which forms thick walls
and bacteria, including the bacteria-
around the cells and enables the plant
derived chloroplasts of plants, with
to construct large sized bodies.
their energy-turnover are the major
driving force for the megamachine of
Fungi are, in contrast to common as-
life on earth.
sumptions not plants at all. Their cell
walls are made up of chitin. Fungi
chemically attack dead or living organ-

Evolution of Animals 5
The body was stabilised by the matrix
of jelly and fibers, and could increase
in size. The matrix expanded more
and more, and filled the cytoplasm,
until in each region of cytoplasm at
least one nucleus and some endoplas-
matic reticulum were retained. These
parts of cytoplasm surrounded by the
membrane of the expanded tube sys-
tem are, by definition, cells, – or if they
had several nuclei, cell complexes. The
Gallertoids: body was compartmentalized into fi-
primitive connective brous matrix and cells. The surface re-
tissues mained covered by cytoplasm with
cilia (later separate cells with cilia), as
the gelatinous matrix may not be ex-
posed to the open water, where it
would swell and pour out.

The result is a body construction in


which many cells and cell-complexes
are fused together with primitive con-
nective tissues. This structural design,
the cell. As a consequence, there are as it is typical for the animal body, had
The Evolution of multicellular two regions in the cell: the inner por- gradually slipped in.
animals tion in strict sense, the so-called cyto-
plasm, and the inner portion of the In the inner of these first ancestral an-
For the understanding of the evolu- imals, some cells specialized in con-
tube system, which may be open to the
tionary history of animals we have to traction, later forming muscles. Be-
surface. The tubes are flexible, they
find answers for the question: How did cause of chemical and mechanical con-
can be widened or narrowed, can
the body construction which is typical straints, contractile fibers could not
merge or branch. Various molecules
for the animals (and only for them) extend themselves again. They have to
are incorporated into the membrane
evolve? In the center of the poster a se- be extended by outer forces. There-
of the tubes. Here the chemical turn-
ries of four technical drawings shows fore, they arranged automatically in a
over takes place and various chemical
these transformations. With these an- counteracting, so-called antagonistic
compounds can be secreted into the
cestral organisms the threshold to the way. This was only posssible in such a
tubes, stored or excreted at the sur-
animal kingdom was overcome. They gelatinoid system. Pressure (by muscle
face. A typical example for the latter
are not drawn at the same scale. The contraction) on the one side tries to
procedure are gland cells.
first (in the back) stands for a small move the body mass to the counter-
unicellular animal, less than a millime- The development towards multicellu- side. The pressure is forwarded to the
ter in size. The last (in front) for a lar animals (the Metazoa) started when side where muscle fibers have to be ex-
larger animal, about some centimeters compounds originating from sugar and tended. The final result are grids of
in size. The last stage represents the proteins were retained in the endo- connective tissue fibers and muscles, –
starting point for all other evolution- plasmic reticulum. These deposits a typical feature of the animal body.
ary transformations in the animal have gelatinous and fibrous properties.
kingdom. Some of the polypeptids are scleropro- We call these archaic animals Galler-
teins (chain-molecules), the basic sub- toids (this German word means: ani-
During this process, the internal tubu- mals with a gelatinoid body design).
stance of collagenous fibers. They
lar system, the endoplasmatic reticu- The first evolutionary steps began in
have a remarkable tensile strength,
lum, played a major role. It had small unicellular animals, less than a
and are the basic matter of connective
evolved already in the archaic proto- millimeter in size. They increased as a
tissue throughout all animals.
cells, when the membrane from the consequence of the new mechanical
surface turned inside, forming tubes in properties, which expanded their

Evolution of Animals 6
range of locomotion and to new food three-dimensional grids. All in all The Vendozoa are fossils, more pre-
resources. The size of such a fibrous- more than twenty types of connective cisely, they are casts of organisms,
gelatinous body is almost unlimited; tissues are known for humans alone. known only from Vendian strata,
jelly fish are structured mainly on this Much more can be expected across the about 600 million years old. The
basis, and some of them attain a size of range of other animals. rounded, often cushion-like forms are
more than a meter. The stability of the probably the relics of organisms that
matrix permitted various body shapes, When skeletal elements developed – had the body structure of gallertoids.
(not only the ellipsoids shown on the and this happened often in the animal
poster). But more decisive was the fact kingdom – the body structure always
that folds and grooves at the surface as a whole determines where and how
could be formed that eventually closed skeletal elements are constructed.
to form canals. In the canals, the water Such hard parts consist of organic
stream was controlled by the beat of compounds like horn or chitin, and
cilia, improving the ingestion of parti- minerals like carbonate or siliceous
cles into the canal wall cells. matter. Such skeletons hold parts of
the body in a definite shape without
Sexual reproduction and vegetative permanent energy supply.
multiplication, both of which occur in
unicellular organisms, must have con- Shifting, enlarging, reduction of these
tinued during the development of elements and/or specialisation of the
A part of the Protozoa, the unicellu-
multicellular animals. The Gallertoid cells determine further evolution. The
lar organisms, can probably be derived
was able to bud off parts of its body for diversity of the animal kingdom, based
from early gallertoids, without folds or
vegetative multiplication, and cells on these relatively few structural ele-
grooves, which remained small and
could continue to exchange material in ments, is impressive.
continued to move by the beat of cilia.
contact with other individuals. Even-
This explanation might be considered
tually some cells specialized to form
especially for the Ciliata, to which the
sperm and egg-cells. In many animals, Animals which are close to Paramaecium belongs, and a number of
budding and fission have been devel- the Gallertoids Flagellata.
oped for specific modes of vegetative
multiplication, apart from sexual re- There are no recent or fossil animals
production. known which completely correspond
to the Gallertoids. However a number
The structural elements and their me- of recent and fossil animals stand rela-
chanical properties, as outlined above, tively close to the ancestral Galler-
persist throughout the entire history toids, can be interpreted on the basis
of animal evolution. They are the in- of this body structure or can be de-
variant structures which provide the rived directly from ancestral stages.
constraints for bauplans and evolu-
tion. In all following evolutionary Trichoplax is about a millimeter in
transitions these existing structural el- diameter, flat, creeping on hard sub-
ements and constraints are only modi- strate. This animal ingests particles
fied. Tissues are rearranged, enlarged into the cells of its underside. Trichop-
or reduced. Cells specialize in frame of lax most likely evolved from an ances-
their capacities. Thes develop into tral gallertoid which had not yet devel-
muscle cells, nerve cells, connective oped canals.
tissue cells, secretorial cells or ga-
metes. The primitive connective tis-
sues differentiate into the defined
types of connective tissues with dis-
tinct mechanical properties. Some of
them are stable against pure and sim-
ple shear or pressure. Others have
elastic properties, some form two- or

Evolution of Animals 7
Evolutionary pathways
The various shapes of the ancestral Sessile way of life
Gallertoid determined all the evolu- The larger the body, the less efficient
tionary pathways of the animal king- is the beat of cilia for locomotion.
dom. In all pathways the structural- Larger gallertoids began a sessile life
functional constraints are preserved. style, this means, they settled perma-
Evolution is only possible on the basis nently on the ground. This was possi-
of what already exists. Evolution is ble as food collection in canals worked
characterized by diversification and, independently of locomotion. For
sometimes at certain levels of evolu- sessile gallertoids, two further evolu-
tion, explosive like radiations can be tionary directions were possible: in-
observed in the fossil record. creasing the number of canals and col-
lecting fine particles, or enlarging a
few canals and collecting larger parti-
Organisms with gelatinous cles.
fibrous body mass
Stromatopora
Gallertoids of a compact body shape, The fossil Stromatopora had no in-
stabilized by the fiber-grid, continued ternal skeleton, but they produced a
to swim by the beat of cilia. The fol- calcareous substrate. Their soft body
lowing evolutionary lineages de- attained a flat shape by reducing
scended from such forms. their gelatinous tissues down to a thin
layer with few canals.
Ctenophores
The ctenophores (comb jellies) are ba-
Sponges
sically similar to gallertoids, as their
body consists largely of gelatinous fi- In gallertoids that stayed with collect-
brous tissues. However, their cilia ing fine particles, the canals multip-
are no more evenly distributed, but ar- lied and arranged in such a way that
ranged to form an efficient ciliary openings for ingestion and egestion
propulsion system. They are united resulted. The efficiency of driving the
on plates, each plate consisting of long water stream was improved, when in
partly fused cilia, resembling a comb, canal wall cells the cilium grew to a
long flagellum. The tiny protrusions Corals and their relatives
and the plates stand in eight rows. A
canal runs under each comb row. Two on these cells, (present in all surface Sessile gallertoids with wider canals
tentacles, which developed from pro- cells) became longer and arranged could catch larger particles. Gastric
trusions of the body, catch particles around the flagellum in the form of a cavities for digestion developed be-
for food. collaret of threads. The finest parti- hind the openings of such canals.
cles, even bacteria, are caught by it. Folds in the canal wall transported and
These cells are called choanocytes, managed the particles, the folds en-
they are placed in a central part of the larged, forming deep pouches into the
canals, the choanocyte chambers. A body mass. The tissue sheets between
skeleton developed, made up of col- the pouches are called mesenteries.
lageneous fibers and calcareous or sili- The flexible tissue of the foremost
ceous sclerites, preventing the sponge part of the canal formed a gullet, the
body from collapsing, when the body pharynx, and tentacles developed in
was hollowed out by more and more front of it. The system, stabilized by
canals. its inner water pressure and held in
shape by the tensile strength of the

Evolution of Animals 8
mesenteries, could be protruded over
the body surface. Such is a typical
coral polyp. The former gallertoid had
been transformed into a coral, the an-
cestor of all later corals.

The octocorals correspond to this


structural type. They developed addi-
tionally calcareous sclerites embed-
ded in the gelatinous fiber grid. All
other corals and sea-anemones, the
fossil Rugosa, and the various Hexa-
corallia, evolved form solitary po-
lyps that descended from the ances- The various internal functions of the
tral coral by reduction of the common The body-cavities develop coelom-chambers have far-reaching
tissues. consequences. The internal water
In ancestral gallertoids with elonga-
pressure stretches the tissues and sta-
ted body shapes, swimming by the
bilizes the body. The connections to
aid of cilia was more and more re-
the central canal were cut. The central
placed by a propulsion through lateral
canal became the intestinal tract, the
bending of the entire body, called un-
gut, and the coelomic-cavities were
dulation. In general, undulation puts
separated from it. The transverse walls
the internal parts of any body under
between them, made up of fibrous tis-
stress: during bending, the lateral parts
sue and muscles, are a kind of tensile
are compressed on the concave side
cords, holding the diameter of the
and stretched on the convex side, and
In a different evolutionary lineage pol- body in a slender shape. In an engi-
in the middle line the two forces neu-
yps with four pouches in the gastric neering sense, such a body is a system
tralize each other. In ancestral galler-
chamber, developed a peculiar method of flexible membranes and a fluid-fill-
toids, it was the jelly/fiber mass (pene-
of vegetative multiplication. The up- ing. We call it a coelomate body or-
trated by canals) that was exposed to
per parts of the polyps were budded ganization.
this stress. As a response to this stress
off, swam for a while settled down, and the lateral canals widened their diame-
grew into a new coral polyp. When the This body organization consists of
ter. They were fluid-filled and hence
period of swimming was extended, connective tissues, muscles, covering
more easier deformed than gelatinous
eggs and sperm began to ripen, and cell layers, and fluids in gut and coelom
body structures. Such animals had to
new polyp developed out of fertilized chambers. The fluids constitute a hy-
expend less energy for moving than
eggs. The alternation of polyp and draulic system which works as a
others, and widening of lateral canals
medusa was set on its way. hydroskeleton for the protagonistic
was favoured. However, the slender
and antagonistic musles.
elongated body shape had to be main-
Examples of large medusae are the
tained. The continuous stress caused a A number of functional properties of
jelly fish of Scyphozoa and Cubozoa.
regular widening of those canals that the coelomic-cavities were decisive for
The Hydrozoa have mostly small me-
ran from the middle to the sides, one the further evolution. In the coelom-
dusae but may form large polyp colo-
behind the other. The resulting fluid- fluid the metabolic end-products of
nies.
filled chambers were placed one be- the surrounding tissues were col-
hind the other, in a so-called lected, and conveyed outwards
metameric arrangement. They are the through small canals. Furtheron cells
so-called coelomic cavities. These for reproduction, i.e. sperm and eggs,
animals are called Coelomates, and developed in the coelom walls and left
they make up the largest part of the the body through the same canals.
animal kingdom. There was plenty of space in the co-
eloms and increasing quantities of
eggs could be produced.

Evolution of Animals 9
The gut remained in the center, run- Polychaetes, bristle worms
ning through the transverse tissue Hydroskeletons: The worm-
In the evolutionary course of the ma-
sheets, which held the gut in its place. coelomates
rine bristle-worms (Polychaeta) the
A tissue sheet remained over and un-
Various kinds of animals descended bulges of the body segments with their
der the entire gut, the so-called me- bristles enlarged more and more.
from the ancestral worm-coelomates.
sentery. Accodingly the gut may have They formed the so called parapo-
There are numerous animals, inaccu-
moved independently from the loco-
rately summarized as ‘worms’, but also dia, appendages used for swimming or
motory movements of the body.
arthropods and molluscs. burrowing. Recent polychaetes show a
remarkable variety of parapodia and
Nutritive molecules and oxygen had to
In most evolutionary pathways the bristles.
be transported beween muscles and
coelom cavities enlarged and made the
connective tissues. Therefore parts of
body highly mobile. As a consequence Many polychaetes have an evertible
the connective tissue became liquid in
these animals were not only able to pharynx armed with teeth. Addition-
the form of internal tracks, which
make undulating movements, but also ally, some polychaetes developed ten-
eventually organized themselves to
peristaltic movements (long/thin tacular crowns around their mouth re-
form real blood vessels. Blood is fluid
and short/thick). The latter can easily gion. These animals live in the sedi-
connective tissue.
be observed in the earth-worm. The ment. Many have adopted a sessile life
thin body wall bulged out in the seg- style and they protrude their tentacles
The gut transported its contents only
ments and additionally small bristles into the water.
in one direction. As a consequence, a
formed.
mouth and an anus, and a fore- and a
rear-end of the body developed. In
The ancestral worm-coelomates used
Coelomates, we can distinguish a back
their mobile mouth region to ingest
(the dorsal side) and a belly (the ven-
nutrients. A strong gullet, the phar-
tral side). The right and left sides are
ynx, developed behind the mouth.
equal. The entire organism is bi-
The pharynx is highly flexible and mo-
laterally symmetrical.
bile.

Comparing the evolutionary pathways


Surface cells of the body secreted a
outlined up to now, four different
layer of collagenous fibers, the so-
PostScript-Fehl
types of body stabilization can be dis-
called cuticula. It is thin, flexible, and
tinguished: (1) The ancestral Galler-
highly tearproof. It supports the circu-
toids and the recent ctenophores are
lar muscles by holding the body shape.
stabilized by gelatinous fibrous tissues,
The result was a complete hydraulic
(2) the sponges by internal skeletal ele-
system composed of thin tissue mem-
ments, (3) the coelenterates by mesen-
branes: the cuticula, the connective
teries in a central gastric cavity and (4)
tissues and muscles form a functional
the coelomates by a metameric ar-
unit. It enclosed the coelom-cavities
rangement of coelom cavities. In the
in which the gut made its peristaltic
coelomates and in the coelenterates
movements to transport nutrition in-
the ancestral organization can be
side.
modified by skeletal elements.
Animals with such a body construc-
In subsequent evolutionary pathways
tion can colonise different environ-
of the coelomates the coelom cavities
ments. Various evolutionary pathways
are modified in various ways. In some
were now opened.
cases it is completely reduced. At an
early stage of evolution two lineages
were separated: the coelomates with a
central body axis, the chordates, and
the coelomates with a metameric hy-
droskeleton, which we call worm-coe-
lomates.

Evolution of Animals 10
Arthropods Oligochaetes, earth worms smaller and by vegetative budding co-
For polychaetes with large and flat In another evolutionary pathway, dis- lonies were formed, encrusting hard
bodies, who lived close to the ground, tinct from polychaetes and arthro- substrates or growing upright like cor-
a quite different evolutionary pathway pods, the parapodia were reduced. In als. In Brachiopoda, a very large ten-
was opened. This pathway led to the some of these organisms the body was tacular crown developed which was
largest diversity in the animal king- long and almost round. They have few encased by a pair of shells. The shape
of these animals resembles that of
dom, the arthropods. The body was small bristles and continued with pe-
mussels.
flattened and the parapodia were ristaltic movements. Examples are
shifted to the ventral side. These ani- the Oligochaeta, among which the
mals scraped particles from the earth-worms are well known. In the
ground and transported them between evolution of the Hirudinea, the
the parapodia to the mouth. The leeches, the coelom was largely filled
parapodia grew longer and finally be- by muscles and a cell mass, called par-
came articulated extremities, i.e., enchyma. In this way the gut was
legs. These animals kept their body tightly connected to the body muscles
more or less stiff, and the body wall be- and enabled to generate enough force
gan to harden in parts. But hardening for sucking.
was possible nowhere else than in the
cuticula, which became thicker and Nematodes, roundworms
was impregnated with chitin. The evo- Other worms with an almost round
lutionary level of the arthropods was body specialized in undulating move-
attained. The chitinous cuticula can ments. As a result, the cuticula
not grow, so these animals have to thickened to stabilize the body. The
mould from time to time. The cuticula muscle-fiber-grid, mainly the circular Platyhelminthes, flatworms
is split off from the underlying tissues, muscles, was reduced up to some Evolution proceeded quite differently
the body is soft for a short while, it longitudinal muscles, so that the origi- in those worm-coelomates that used
grows, and secretes a new cuticula, nal metameric coelom cavities were the flexibility of transversal walls to
which is finally hardened. The diver- fused to one large cavity. This is the flatten their body. This enabled them
sity of arthropods encompasses three body structure we find in the nema- to creep along hard surfaces, and even
quarters of the animal kingdom. The todes, the roundworms. to hold on to such a surface. In addi-
Chelicerata, the spiders and their tion, with the production of slime,
kin, have one pair of extremities in Bryozoa and Brachiopoda, they could wrap around a prey and
front of the mouth, the chelicerae. Pogonophores overcome it.
All others have one or two pairs of an- Some of the worms, living on the
tennae in that position, i.e., the fossil The flattening of the body was im-
ground, protruded a body segment at
trilobites, the crabs and their rela- proved when the coelom cavities were
their front end developed into a ten-
tives, the centipedes, millipedes, reduced and eventually completely
tacular crown around the mouth. The
and the insects. filled by muscle fibers and a cell mass,
segmentation of the body was reduced
and the animals assumed a sessile life the parenchyma.
style.

The Pogonophores have long and


small tentacles and a long and ex-
tremely thin body, in which the gut
was reduced. Nutrition in the form of
microparticles and molecules is in-
gested through the outer epithelium.
Body segmentation and bristles per-
sisted only at the hind end.

In the evolutionary course to the


Bryozoa the individuals became

Evolution of Animals 11
The dense tissue could form suckers at plants could be attacked. Further
any place and the mouth shifted more transformation resulted in the fossil The chordates
to the ventral side of the body. This is and living molluscs: the dorsal plates
The evolution of chordates is an alter-
the body structure of the Plathelm- of chitons are a secondary arrange-
native development in comparison to
inthes. The Turbellaria, the flat- ment, not reflecting directly the old
that of the worm-coelomates de-
worms, have a sucking gut. The metameric design. The worm-mol-
scribed above. It started in those an-
Cestodes, the tape-worms, reduced luscs are descendants of similarly
cestral coelomates that kept their
their gut completely. They live parasit- structured forms, characterised by a
mouth region relatively stiff. The side
ically in the intestinal tracts of ani- loss of the shell. A high and spirally
canals formed slit-like openings in the
mals. rolled up shell characterizes the Ga-
gut, the gill-slits, finally forming a gill-
stropoda, the snails.
basket, the so-called branchial gut.
Mollusca
These animals conveyed a water
The evolutionary development to the stream into the gut by the beat of cilia.
molluscs started in a quite similar way The water left the body through gill-
as that of the flatworms. However, a slits and food particles were caught by
muscle-fiber-grid on the ventral side sticky slime. The mass of slime and
enlarged, forming a cushion, movable particles was transported into the gut.
in itself and able to adhere to a hard
substrate and to creep along it: the For locomotion, undulating move-
creeping foot of the molluscs. The ments were sufficient. As a conse-
coelomic cavities were shifted to the quence, the foremost part of the body
dorsal side, and above them hard skel- developed a rostral filter system
etal plates of horn and calcareous de- (and finally a head), and the larger part
posits stabilized the back side. behind it became a locomotory sys-
tem. This divison of the body was de-
These animals could tightly press their cisive for all further evolution of chor-
mouth to the substrate and graze food. dates.
On the lower side of the mouth horn
bosses and teeth developed as a re- Lengthwise arranged (longitudinal)
sponse to the stress of being pressed to In the Bivalvia, the mussels, the shell muscles, alternately contracting on
the ground: the rasping tongue, the is divided in two halves. Among the the right and left sides, allowed the
radula, evolved. Almost no surface Cephalopoda, the fossil ammonites body to bend and undulate. Under the
has ever been resistant against this and the living Nautilus have a cham- stress of these actions, connective tis-
strong working rasp. The kinds of nu- bered coiled shell, and in the squids sue fibers and all muscles were ar-
trition available to these animals could the shell is shifted internally into the ranged in an efficient system. In the
be extended dramatically, and even shell cavity. mid-line a portion of the ancestral fi-
brous-gelatinous mass remained and
became enveloped by sheets of con-
nective tissues. This was in fact a tube
that could be bent but not compressed
and not shortened. This axial rod is
called the notochord, from which
the name chordates was derived. It
holds the length of the body without
energy input, and the originally
present circular muscles, which had
balanced the forces of the longitudinal
ones, could be reduced. These became
arranged in a new way and formed a
thick skin, including fibers and surface
cells, as is typical for chordates.

Evolution of Animals 12
Graptolites, Pterobranchia
Further steps in evolution led to ani-
mals that wriggled themselves into the
sediment and stayed in it permanently.
They lived in the sediment and pro-
truded their front end into the ambi-
ent water for food collection. The col-
lar could extend like a funnel, and
eventually split to form tentacles.
The small Pterobranchia, still
equipped with two gill slits, and the
fossil Graptolites, constructed stiff
tubes of collagen.

The longitudinal muscles enlarged Enteropneusts, acorn worms


more and more. The coelomic cavities The evolution of worm-shaped chor-
lost their original stabilizing function. dates and their descendants began in
They were shifted to the ventral side, those ancestral chordates that used to
became smaller and merged to just a live near the bottom, where food par-
few cavities or only one cavity, in ticles accumulate which have sunk
which the gut and finally other inter- down in the water column.
nal organs found their place. The body
was slightly flattened and a fold in the The foremost part of the body, above
skin made up a kind of fin in the mid- the mouth, was made up of flexible tis-
line of the body. This shape improved sues, and could be used for wriggling
undulatory locomotion. In further and digging in the sediment, stirring
evolutionary steps, hard skeletal parts up particles. This part enlarged and fi-
could develop in the skin and in the in- nally developed a trunk-like digging
ner connective tissues. organ, the so-called proboscis. The
rear part of the body lost its locomo-
This specific chordate bauplan stands tory function and the notochord and
in remarkable contrast to that of the muscles were reduced. A transi-
worm-coelomates, in which most of tional part remained between the two
the body-mass is arranged around the portions. Each of them had one coelo-
large body cavity, and in which skeletal mic cavity.
plates could develop only at the outer
surface. The enteropneusts, the acorn-
worms, show this body structure: they
The branchial gut region developed in have a short stiffened structure in the
quite different directions. Some collar, which can be interpreted as the
evolved jaws and skull, some extended remnants of the old notochord.
the branchial gut enormously, and oth-
ers became borrowing and creeping
worms and finally developed tentacles.

Evolution of Animals 13
Echinoderms
Peribranchial-chambers
The evolution of echinoderms started
in similar sessile animals with three The gill slits cause a mechanical prob-
coelomic cavities. Tissues near the lem: the larger and more numerous
body surface were stiffened by calcare- they are, the more they weaken the
ous material in the form of plates. The body wall. Therefore gills arches are
fossil mitrates were an early side- stabilized by stiffer tissue, mostly of
branch of this develoment. They had a cartilage. An additional stabilization
row of elongated openings, inter- developed when the body grew wider
preted as gill slits. The first true echi- and around the branchial gut, so that a
noderms root in sessile animals with peribranchial chamber resulted. Wide
five tentacles. The base of such a ten- flat body shapes developed, stiffened
tacle crown is cup-shaped, and it is a by plates of bone, as in the fossil jaw-
movable soft hydraulic system. With less fishes, some examples of which
increasing body size, this tentacular are the Heterostraci or Osteostraci.
cup could be supported on its back
side, when the next body region ex- Chordates with a vertebral
tended around it. This part could be column and tooth elements
stiffened by plates and extended more
Some chordates with elongated body
and more. Finally the tentacles ap-
shapes developed a vertebral column
peared as sunk into a plate-stiffened
around the notochord and special
surrounding. This is the coelom-in-
tooth-like elements in the mouth and
coelom encapsuled body structure we
gullet. Examples of such organisms are
find in living echinoderms: In a body
the fossil conodonts which had com-
coelom is a small coelom (rest of the The living lancelet Branchiostoma
plicated arrangements of various bars
reduced proboscis) and a large ring- has a slender shape, a large branchial
or tooth elemets. The function of
like coelom with extensions into the gut and has a notochord over the
these conodont apparatuses is not yet
tentacles. The sea-lilies still have a whole length of the body. At its mouth
well understood, but in their micro-
tentacular crown on a stalk; the star- small tentacles developed, used for
structure these elements are quite
fish (sea-stars), the brittle-stars, the food uptake.
similar to the teeth of vertebrates. The
sea-urchins, and the sea-cucumbers,
conodonts do not have any recent rel-
descended from upper body parts that Tunicates
atives, but in regard to recent findings,
were budded off and became indepen- A large bran- their position close to the vertebrates
dent of their stalk. chial gut contin- is most likely.
ued to collect
food without lo-
comotion; such ani-
mals could assume a sessile
life style. This development caused
far reaching changes, by which the
tunicates evolved. The system for
locomotion, notochord and mus-
cles, were reduced, however, it is
still present in the free swimming
larvae. The branchial gut extended to
an enormous size, hanging in the thick
tough peribranchial chamber, the wall
of which, the so-called tunica, con-
tains horn and chitin. The sea-squirts
are permanently sessile; the salps have
a thin-wall and have returned to free
swimming.

Evolution of Animals 14
toral girdle and pelvic girdle) devel-
oped into the shoulder girdle and the
pelvic girdle. They both provided
joints for the legs and surfaces for the
attachment of the leg-muscles. To-
gether with the vertebral column a
carrying system was constructed,
which allowed the complete body to
be lifted over the ground. With this
evolutionary step the stage of tetra-
pods was attained.
Fishes
Chordates with jaws, skull and Today we can distinguish different tet-
In the evolutionary course of fishes
vertebral column rapods. The amphibians have to lay
bony scales developed in the skin tis- their eggs in the water, where their lar-
Ancestral chordates with vertebral sues. These scales were arranged in vae develop. The reptiles are able to
columns additionally developed jaws such a way, that they limited the range live on land permanently. Even those
and skull. The branchial basket was al- of lateral bending. The head was stabi- who went back into the water have to
ternately expanded and compressed. lized by large bony plates, and the car- come onto land for laying their eggs.
These movements drove a water cur- tilaginous capsule, enclosing the brain As an example for the reptiles a dino-
rent through the gill slits, so that and the sensory organs was replaced saur is shown in the poster (Triceratops
breathing and feeding was improved. by bony material. Finally a bony skull is the logo of Senckenberg-Museum).
Even larger particles were swallowed was formed. The lateral, dorsal and The dinosaurs are a certain group of
by the pharynx. Some of the rostral ventral fin folds of the ancestors dif- reptiles which have their closest rela-
branchial arches were enlarged and ar- ferentiated into certain fin-types. tives in the recent (and fossil) birds.
ranged as a cartilaginous jaw. The re- Some reptiles, the so-called therapsids
sult was an efficient apparatus for In the fossil record the first fishes that
are the ancestors of the mammals.
catching and holding large particles were representatives of these evolu-
From these early mammal-like reptiles
and even living prey. tionary stages are the placoderms.
a huge number of different organisms
Their head was reinforced by large
evolved.
The movements of this primordial bony plates and their body shape was
jaw-apparatus were oriented up and stabilized by a cartilaginous vertebral Man also belongs into the group of the
down – which means vertically –, while column and by large scales in the skin. mammals. Man evolved from schim-
the movements of the body for swim- The cartilaginous fishes, another evo- panzee-like ancestors and developed
ming were oriented left and right – lutionary pathway, not figured in the an upright walking position. As a con-
which means laterally. These different poster, have only small tooth-like sequence the brain enlarged enor-
movements were mechanically sepa- scales all over their entire body. A mously. This brain allows thinking,
rated from each other by the last gill bony skeleton developed only in the combining and teaching the following
arch (i.e. visceral arch). This skeletal pathway to the bony fishes. generations. Such a brain is unique in
element enlarged more and more, un- the entire animal kingdom, and on the
til the complete trunk musculature Tetrapods
basis of its intellectual capacities, man
found its attachment at its surface. We Most fishes developed lungs from a constructs his own view of the world.
call this last visceral arch the shoulder bulge of the foregut and finally swim-
girdle. Since it separated the head bladders. These fishes
from the trunk, it played a crucial role were able to breathe
for further evolution of fishes and air. Some of them
tetrapods. Around the head a massive had a body organiza-
cartilaginous capsule developed and tion which allowed
the notochord became surrounded by them to transform
various cartilage elements, suppress- their fins into legs and
ing the range of lateral bending. to carry the complete
body on land. The
two bony girdles (pec-

Evolution of Animals 15
Scientific theory, Darwinism Certain evolutionary pathways, and
texbooks on animal bauplan
Baumunk, B.-M. & Riess, J. (Hrsg.) (1994):
Darwin und Darwinismus: eine Ausstellung Gudo, M., Gutmann, M. & J. Scholz (Hrsg.)
zur Kultur- und Naturgeschichte. Katalog; ei- (2002, in Vorb.): Concepts of Engineering-
ne Publikation des Deutschen Hygiene-Mu- and Functional Morphology: Biomechanical
seums Dresden; 265 S. – (Akademie) Berlin. approaches to recent and fossil organisms.
Senckenbergiana lethaea. – Stuttgart.
Janich, P. & Weingarten, M. 1999. Wissen-
schaftstheorie der Biologie. 315 S. – (Wilhelm Gutmann, W. F. (1994). Morphologie & Evo-
Fink) München. lution. Symposien zum 175jährigen Jubiläum
der Senckenbergischen Naturforschenden
Weingarten, M. (1993): Organismen - Objek-
Gesellschaft. Evolutionssymposion. Sen-
te oder Subjekte der Evolution? Philosophi-
ckenberg-Buch 70, 454 S. – (Kramer) Frank-
sche Studien zum Paradigmawechsel in der
furt am Main.
Further reading Evolutionsbiologie; 314 S. – (Wissenschaftli-
che Buchgesellschaft) Darmstadt. Storch, V. & Welsch, U. 1991. Systematische
The evolution of animals is presented here in Zoologie. – Gustav Fischer, Stuttgart, New
a new way, which is not like that of the stan- Chemo-Evolution and the origin of York, 731 S.
dard textbooks. The engineering-like expla- life on earth Westheide, W. & Rieger, R. 1996. Spezielle
nation of evolution and the reconstruction of Zoologie – Erster Teil: Einzeller und Wirbel-
Kempe, S., Kazmierczak, J. & Degens, E. T.
evolutionary pathways contrasts sharply with lose Tiere. 909 S. – (Gustav Fischer) Stutt-
(1989): The soda ocean concept and its bea-
morphometric and genetic phylogenetic gart, Jena, New York.
ring on biotic evolution, 29-43. – In: Crick, R.
trees. However, explanations are provided,
E. (ed.): Origin, Evolution, and Modern As-
which are missing in most traditional presen-
pects of Biomineralization in Plants and Ani-
tations.
mals; 536 S. – (Plenum Press) New York.
The results which are the basis of this poster
Wächtershäuser, G. 1997. The origin of life
Acknowledgements
have been developed since 1970 by the group
and its methodological challenge. – Journal of
„Kritische Evolutionstheorie“. Continous A number of colleagues did read the text and
Theoretical Biology, 187: 483-94.
discussion with philosophers and testing im- its english version and made valuable sugge-
proved the coherence of the methodology. Dullo, C., Mosbruger J. & Oschmann, W. stions. Dr. Sven Baszio, Prof. Dr. Euan Clark-
The listed references should give an idea for (2000). Geobiologische und Paläobiologi- son, Doris von Eiff and Dr. Eberhard Gisch-
further readings. sche Prozesse als Antrieb der Evolution des ler. To all of them we extend our thanks.
Systems Erde. – Kleine Senckenberg Reihe
For financal support we thank PASS IT Con-
36. – (Kramer) Frankfurt am Main.
sulting GmbH.
Frankfurt theory of Evolution and
Engineering Morphology
Endosymbiosis and theory of the
Scharf, K.-H. & Gutmann, W. F. (Hrsg.)
biosphere Evolution in teaching courses
(1993): Evolution von Organismen. – Praxis
der Naturwissenschaften, 42: Biologie: 1-49. Margulis, L. (1981): Symbiosis in Cell Evoluti- The poster „The Evolution of Animals“
on. – (Freeman Co.) San Francisco. is available together with this text either
Gutmann, W. F. (1995): Die Evolution hy-
Mollenhauer, D. (1994): Endocytobiosen. An- in German or in English. Certain materi-
draulischer Konstruktionen: Organismische
lässe zu Revisionen in Taxonomie und Phylo- als for teaching are also avaiable (over-
Wandlung statt altdarwinistischer Anpas-
genetik: 339-363. – In: Gutmann, W.F., Mol- heads, slides) and guided tours on the
sung. Senckenberg-Buch 65, 220 S. – (Kra-
lenhauer, D., Peters, D. S. (Hrsg.) (1994): Mor- evolution of animals through the Sen-
mer) Frankfurt am Main.
phologie und Evolution. – Senckenberg-Buch ckenberg Museum can be offered
Edlinger, K. Gutmann, W. F. & Weingarten, (Phone: 069/7542-357).
70; 454 S. – (Kramer) Frankfurt am Main.
M. (1991): Evolution ohne Anpassung. - Auf-
Krumbein, W. E. & Lapo, A. V. 1996. Vernad- Please sent your orders to:
sätze und Reden der Senckenbergischen na-
sky's Biosphere as a Basis of Geophysiology. Schriftentausch der Senckenbergischen
turforschenden Gesellschaft, 37: 92 S. –
115–134. In: Lovelock, J., Margulis, L., Saun- Naturforschenden Gesellschaft,
Frankfurt am Main.
ders, P., Whitfield, M., Goodwin, B. & Ho, Senckenberganlage 25,
M.-W. (Hrsg.): Gaia in Action-Science Of 60325 Frankfurt am Main,
The Living Earth. – (Floris Books) Edin- Phone: 069/7542-246,
burgh. e-mail: sjessel@sng.uni-frankfurt.de.

Levit, G. S. 1999. Biogeochemistry-Biosphe-


re-Noosphere: The Growth of the theoreti-
cal system of Vladimir Ivanovich Vernadsky.
Studien zur Theorie der Biologie Band 4. –
(VWB) Berlin.

Evolution of Animals 16

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