changing, but also that it has been changing from the earliest geologic
times. What, then, is to be said about the living creatures that have
existed in the changing environment during all these geologic ages?
Have they been rigid, unyielding forms? By no means! We know that they
can be modified by altering the conditions at present; and a study of
the fossils in the rock formations of the different ages of the world
shows conclusively that animals and plants have altered in the past with
the changing environment. The living creatures in the _Silurian ages_
differ from those in the succeeding _Devonian ages_, and these latter
differ from those in the still later _Carboniferous ages_; and so on,
to the present. Changing physical geography and climate are associated
with changing forms in animal and plant life. The growing amplitude and
complexity of a continent are associated with increasing complexity and
specialization of its living forms. Just as the North American Continent
of the Tertiary period differs from that of the Silurian ages, so also
do the animal and plant forms of the Tertiary period differ from those
of the Silurian ages. Just as there has been a continuity in the growth
of the Silurian continent to that of the Tertiary ages, and the present,
so, also, there has been a continuity of living creatures from Silurian
to Tertiary and present times. Changing conditions of life have compelled
modifications in living forms, and those creatures that were unable to
adapt themselves to the altering conditions of life have perished, while
those that did adapt themselves, through useful variations, lived and
progressed in organization.
A study of the fossils in the rock formations (see page 95) of different
ages reveals the fact that thousands of species have lived and flourished
in one age and then perished, never to appear in succeeding ages. A study
of the fossils also reveals that life-forms have passed on from age to
age, ever changing with the changing continent, some advancing to higher
and higher levels, while others remained lowly.
The facts enumerated above may be instructively illustrated by a hasty
reference to the history of organic life as unfolded by the rocks of
different ages.[7]
=Archæan Era.= No evidence of life has been discovered in the Archæan
rocks, but, inasmuch as with the dawning of the Palæozoic time, the
waters of the sea were peopled with plants and animals living in great
numbers and considerable variety, it is evident that the ancestors of
these creatures must have lived during the Archæan ages.
TABLE OF STRATIFIED ROCKS AND THE SUCCESSIVE APPEARANCE OF TYPICAL ANIMAL
LIFE-FORMS.
+----------+-----------------------+------------------------------------+
| | |=Man=, Mastodon, Saber-toothed |
| | Quaternary. | Tiger and other Vertebrates. |
| | | Invertebrates. |
| +---------+-------------+------------------------------------+
| | | |Equus and other Vertebrates. |
| | | Pliocene. | Pliohippus. Protohippus. |
| | | | Invertebrates. |
| | +-------------+------------------------------------+
|Cenozoic. |Tertiary.| Miocene. |Miohippus and other Vertebrates. |
| | | | Mesohippus. Invertebrates. |
| | +-------------+------------------------------------+
| | | |=Monkeys.= |
| | | |=Lemurs.= Primitive Carnivora and |
| | | Eocene. | Herbivora. Orohippus. |
| | | | Invertebrates. Eohippus. Fishes, |
| | | | Amphibians, Reptiles. Land Birds. |
| | | | Monotremes. Marsupials. |
| | | |=Prim. Monodelphs.= |
+----------+---------+-------------+------------------------------------+
| | |Marsupials. Monotremes. Reptiles. |
| | | Reptilian Birds. Amphibians. |
| | Cretaceous. | Teleosts and other Fishes. |
| | | Belemnites and other |
| | | Invertebrates. |
| +-----------------------+------------------------------------+
| | |Reptilian Birds, Reptilian Mammals. |
|Mesozoic. | Jurassic. | Rays, Chimæroids and other |
| | | Vertebrates. Invertebrates. |
| +-----------------------+------------------------------------+
| | |Primitive =Marsupials=, Primitive |
| | Triassic. | Monotremes, Reptiles, Amphibians, |
| | | Fishes. Ceratites and other |
| | | Invertebrates. |
+----------+--------------+--------+------------------------------------+
| | |Permian.|Primitive =Reptiles=, Amphibians, |
| | | | Fishes. Invertebrates. |
| |Carboniferous.+--------+------------------------------------+
| | | |Primitive =Amphibians=. |
| | | | Fishes, Invertebrates. |
| +--------------+--------+------------------------------------+
| | |Primitive =Crossopterygii=, |
| | Devonian. | Ganoidei, Dipnoi. Goniatites and |
| | | other Invertebrates. Sharks. |
|Palæozoic.+--------------+--------+------------------------------------+
| | | Upper. |Primitive =Sharks=. Ostracoderms. |
| | | | Invertebrates. |
| | Silurian. +--------+--------------------------------+---+
| | | Lower. |Blastids, Scorpions, Centipedes,| I |
| | | | Sea-urchins and other | n |
| | | | Invertebrates. | v |
| +--------------+--------+--------------------------------+ e |
| | |Marine =Worms=, Molluscs, | r |
| | | Trilobites, Brachiopods, | t |
| | Cambrian. | Crinoids, Star-fishes, | e |
| | | Corals, Graptolites, | b |
| | | Cystids, Sponges, | r |
| | | Foraminifera. | a |
+----------+-----------------------+--------------------------------+ t |
| Archæan. |=Protozoans=, though there is | e |
| | no evidence of life. | s |
+----------------------------------+--------------------------------+---+
=Cambrian Period.= The Cambrian rocks have furnished many different
species of marine animals. Seaweeds are the only plants found fossil
in the Cambrian rocks. Some plants and animals may possibly have dwelt
upon the land, but if so they have failed to leave any record of their
existence. The animals are all _Invertebrates_, but not of extremely low
forms; they have progressed since their appearance in an earlier period.
Siliceous Sponges are not uncommon. The Cœlenterates are represented
by Graptolites and Corals. Echinoderms are rare, and are principally
represented by Cystids, a very primitive type; true Crinoids and
Star-fishes appear before the close of the period. That marine worms
existed is indicated by borings and tracks in the sands, which have
since consolidated into rocks. Mollusca burrowed in the mud or crawled
over the ocean bed. Brachiopods existed, and Arthropods were represented
by primitive types. The most characteristic of Cambrian fossils are
_Trilobites_ and _Lampshells_.
The former are extinct, and are not represented in the modern ocean; the
latter, although greatly reduced in variety and numbers, are still found
in various parts of the sea. During all this period, no backboned animals
existed; then there were no fishes or amphibians, no reptiles or birds,
and no mammals; nothing but invertebrates. The Cambrian fauna shows
steady progress, being decidedly more advanced in the upper divisions
than in the lower ones.
=Lower Silurian (Ordovician) Period.= The life characteristics of this
period are very similar to those of the Cambrian. Brachiopods have
developed extensively, though they have not yet reached their height
of development. The Trilobites attained their greatest development in
this period; but after the Lower Silurian these creatures commenced to
decline in variety and numbers, and finally underwent extinction near the
close of the Palæozoic era. In America no plants have been discovered
above the grade of seaweeds, but a few of the higher Cryptogams are
doubtfully reported in Europe. The character of the flora in a later
(Devonian) period makes it highly probable that land plants were well
advanced in the Lower Silurian period. Professor Scott states that the
remains of land plants may be discovered at any time, though this must
remain a matter of chance, inasmuch as all known Ordovician rocks are
marine, and therefore not a favorable circumstance for the preservation
of land plants. Foraminifera and Radiolaria were abundant in the seas
of the Lower Silurian period. Sponges and Brachiopods are numerous and
varied. Among Cœlenterates the Graptolites are very numerous and varied;
the few and doubtful Cambrian Corals are succeeded by a considerable
number of Ordovician genera and species. They were characteristically
different from the reef-builders of modern seas. The Echinodermata have
greatly increased in importance, being more numerous and varied. The
_Cystids_ reach their maximum development in this period. The Crinoids
greatly increase in numbers and variety. The Star-fishes greatly expand;
and a new and higher order of Echinoderms, the _Sea-Urchins_, make their
first appearance in the later Ordovician period, but under very primitive
forms.
Among Arthropods the Trilobites increase greatly in numbers and variety,
and attain their maximum development in this period. The occurrence of
a Centipede in Ordovician rocks indicates the interesting fact that
terrestrial animal life had already begun.
One of the most striking differences between the Lower Silurian and
Cambrian periods is the great advance made by the Molluscs, in variety
and numbers, during the former period. Molluscs exhibit the most
significant change in the great expansion of the _Cephalopods_, a few of
which had appeared in the uppermost Cambrian rocks. In the Lower Silurian
period the Cephalopods became one of the predominant types in the marine
life. In modern times nearly all Cephalopods are naked (_Cuttlefish_ and
_Squid_), only a few having a shell (_Nautilus_ and _Argonaut_). The
naked forms are higher creatures than those with shells. In the Lower as
well as in the Upper Silurian periods no naked forms existed, but only
the lower forms with shells (Nautiloids). In modern times and throughout
later geological periods only Nautiloids with coiled shells were in
existence. In both of the Silurian periods the shells were all straight,
and the animals were called Orthoceratites. Such animals were extremely
abundant in those times, and often reached an enormous size. Specimens
have been found which were ten inches in diameter and over fifteen feet
long. They were the most formidable animals of those early ages, and
were the rulers and scavengers of the seas; so that the Lower and Upper
Silurian periods are known as the =Age of Molluscs=.
=Upper Silurian Period.= In this period great progress is made in the
history of life on the globe. Upper Silurian life is the continuation and
advance of the organic system that flourished in the Ordovician, certain
groups expanding, others diminishing; and some new groups now appear
for the first time. Sponges are still common. Among Cœlenterates the
Graptolites have greatly diminished; Hydroid Corals have become important
features of the seas and in the formation of the reefs. True Corals
increase largely and play a more important part than in the preceding
period. Honeycomb and Chain Corals are quite characteristic of this
period.
Among Echinoderms there is a diminution of the Cystids, and a marked
increase of the Crinoids. A new class of Echinoderms, the Blastids,
now make their first appearance. The Sea-urchins and Star-fishes
have increased in variety and abundance. Brachiopods still exist in
multitudes, but under changed forms. Among Molluscs the Orthoceratites
are still the rulers of the seas. Among Arthropods the Trilobites are
still numerous, though decidedly less so than in the preceding period.
The land animals are insects of low types, mainly allied to cockroaches.
Scorpions are also present as fossils in the Upper Silurian rocks. These
animals prove the existence of a contemporaneous land vegetation.
During all the many millions of years that constitute the Silurian
periods, hosts of species lived and died; hosts of variations were
induced in the living creatures by the ever-changing environment; so
that at the close of this time the Silurian rocks show that some of the
life-forms, lowly as they were, had yet climbed higher in the scale of
organization.
The later Silurian rocks reveal the remains of _an entirely new branch
of living forms, a higher branch of animals_ (=Vertebrates=) than
had ever before existed. These remains are those of =Ostracoderms=
and =primitive Sharks=. But at that early time they held a very
subordinate position among the hosts of living creatures; the Molluscan
Orthoceratites were still the rulers of the seas; it was still the Age of
Molluscs.
=Devonian Period.= The next set of ages succeeding the Silurian time
is known as the Devonian period. Many of the kinds of creatures living
in the Silurian ages are also found in these ages, but under changed
forms. The Devonian seas had a great abundance and variety of Corals
and Crinoids. The Chain Corals have perished, but the Cup and Honeycomb
Corals still live on with modified appearance. The Graptolites are almost
extinct. The Orthoceratites still live on, much reduced in numbers and
size. The Molluscan Cephalopods have been revolutionized. New forms have
grown out of the variations of the past, so that now in Devonian times
we have the introduction of the great Molluscan _Ammonite family_, under
the forms known as _Goniatites_, which are characteristic of this and the
succeeding Carboniferous ages. The Trilobites still continue under new
forms, but much reduced in size and number. Brachiopods are abundant and
diversified.
In this age, for the first time, land plants become conspicuous. The
Devonian forests consisted of the highest flowerless plants, such as
ferns, horsetails, and club mosses; and also the lowest flowering plants,
such as cycads, pines, cypresses, etc.—plants which have imperfect
and inconspicuous flowers. In all ages, as now, land vegetation has
been closely related with _insects_. Insects, though rare as fossils,
are found in connection with the forests in the Devonian period. They
are among the lower orders of the class, and are somewhat allied to
cockroaches and dragon flies.
The most characteristic feature of the Devonian age is the expansion
of that new and higher class of animals, =the backboned animals=; for
during the earliest Devonian or latest Silurian times the =Fishes= made
their appearance. At first many of them were comparatively small in size,
they were few in numbers, and of strange, unfishlike forms. Such were
the _Ostracoderms_, which, though generally called fishes, belong to a
type much below the true fishes, and more nearly allied to the Lampreys,
for they were devoid of jaws and paired fins. There was a great variety
and wealth of true fishes in the Devonian period. The _Elasmobranchii_
(Sharks) were well represented, _though very generalized ones_.
_Crossopterygii_, _Dipnoi_ and _Ganoidei_ were important elements of the
fish fauna of this period. At this period there was an entire absence of
those highly specialized fishes (_Teleosts_) which in modern times make
up the vast majority of fishes, both marine and fresh-water.
The true fishes soon developed to so great an extent in size and numbers
that they swarmed in the Devonian seas and quickly became the rulers of
the age; hence Devonian time is known as the =Age of Fishes=.
These early Devonian Fishes were of a more lowly organization than modern
ones, and were what is called generalized forms; that is, they combined
in themselves the characters of two distinct classes. They had distinct
_amphibian characters_. From these generalized Fishes were afterwards
formed, through many transitions, the _Amphibians_ and the _Fishes_ as
branches from a common trunk. This illustrates what is a very general
law, viz.: that the first-introduced examples of a class are not typical
forms of that class, but intermediate forms or connecting links with
other classes. Certain footprints recently discovered in the upper
Devonian rocks of Pennsylvania indicate that the Amphibia, the lowest of
air-breathing vertebrates, commenced their career in the latter part of
the Devonian period; but they were inconspicuous among the monarch fishes.
=Carboniferous Age.= As in the Devonian Age so in the Carboniferous
times, we find flowerless plants, such as ferns, horsetails, and club
mosses; but in this coal-forming age they culminate, and have become
gigantic in size, especially the two latter. At this time we again meet
with the next higher order of plants, as cycads, pines, cypresses, etc.
These plants of the coal period were remarkable, generalized types,
connecting classes now widely separated. During this period the insects
increase in variety and numbers along with the advancing vegetation.
It is interesting to find that as the highest flowering plants are not
yet in existence, so those highest orders of insects, the flower-loving
and honey-loving ones, such as bees, butterflies and moths, are not yet
in existence. Corals and Echinoderms, Molluscs and Crustaceans, etc.,
continue through the ages with ever shifting forms. Brachiopods have
greatly diminished. Foraminifera for the first time assume considerable
importance in the earth’s economy.
The Molluscan Goniatites continue through this age with changing forms
and advancing organization, and are very numerous. The Trilobites, which
are characteristic of Palæozoic times, continue under new forms through
this age, and then perish. In this age are introduced for the first time
typical Crustaceans of the long-tailed kind, such as shrimps.
Devonian fishes still prevail. The Elasmobranchii are numerous and
varied, and some of them are highly specialized. _Pleuracanthus_ is a
remarkable shark which has many features in common with the lung-fishes
(Dipnoi), such as the character of the pectoral fins, the shape of the
tail, the bones which form a roof for the skull, while the skin is naked.
The Dipnoi continue, though in diminished numbers.
The Crossopterygians are much less abundant. The Ganoidei increase in
numbers and varieties.
The fact of greatest interest concerning the Carboniferous period is the
_expansion of the true land-breathing, backboned animals, such as the_
=Amphibians=. _Their expansion marks a distinct step forward in the scale
of life._
The Carboniferous Amphibia all belong to the extinct order of
_Labyrinthodonts_ (_Stegocephala_), in which the skull is well covered
with a roof of sculptured bones. The Amphibia of the lower Carboniferous
rocks are of small or moderate size, not exceeding eight feet in length
and mostly much smaller. A great number of these Amphibians are known,
most of them like the Salamanders in shape, but some are snake-like
in form, being long and slender. An example of the Stegocephala is
_Archegosaurus_.
The upper Carboniferous rocks represent the later Carboniferous period
(=Permian Period=). The life of this period is transitional between that
of the Palæozoic and of the Mesozoic eras. Here we meet with the last
of many types which had persisted from Cambrian times, associated with
forms which are prophetic of the characteristic types of the Mesozoic
era. We also meet with types in the Permian that are peculiar to the
period. In this section of the Carboniferous period many of the genera
of the fishes are the same, while new ones are introduced. Among the
lung-fishes the genus _Ceratodus_ is introduced, a creature very closely
allied to the modern lung-fish of Australia. The Amphibia are still
represented by the Stegocephala, several of the older genera persisting,
while many new forms appear for the first time; several of the latter
surpass the earlier Carboniferous genera greatly in size. The transitions
from many of the Devonian fishes to the Carboniferous Labyrinthodonts
are so gradual that it is sometimes difficult to say whether we are
dealing with Fish or Amphibian. In other words, the Devonian fishes are
generalized fishes, that is, connecting links between Fish and Amphibian.
When the Amphibians finally separated from the fishes they were not the
highly specialized forms of more recent times; but they were generalized
Amphibians, having some reptilian characteristics. As time passed on
and the creatures continued to modify, some varied more and more in the
direction of true amphibians, and others more and more in the direction
of reptiles, until in the Permian stage early, generalized Reptiles, of
lizard-like form, appeared in large numbers. The most interesting of
these primitive Reptiles are the _Theromorpha_ (beast forms), which
present many remarkable approximations to the structure of the mammals.
As yet no snakes or turtles, no alligators or crocodiles had come into
existence. In spite of the fact that great advance in animal life was
made during Carboniferous times, there was little of the life in those
ancient woods that we associate with the forests of the present; they
were gloomy wastes of shade, without the presence of bright flowers, no
humming of the bees, no song of birds, and few sounds save the gurgling
of running streams, the sighing of the wind through the leaves, the
splash of waves upon the shore, and the bursting of the thunder clouds.
The interest and importance of the Carboniferous Amphibians as the first
land backboned animals, is so great as to cause the Carboniferous period
to be spoken of as the =Age of Amphibians=.
=Triassic Period.= In this period the ferns and horsetails continue
under new forms; but the next higher orders of plants, the Cycads and
Conifers, now predominate. The Goniatites are replaced by the Molluscan
_Ceratites_, the latter being characteristic of the Triassic period.
One of the most characteristic changes from the Palæozoic to the Mesozoic
era consists in the great reduction of the Brachiopods.
The =Vertebrata= of this period are of extraordinary interest and show
great progress. The fishes exhibit the least progress. The Dipnoan
_Ceratodus_ is very characteristic, continuing from the Permian. The
Crossopterygians have greatly declined. The Ganoidei continue to be
the dominant fish-type, and are most like the existing gar-pikes. The
Amphibia (Stegocephala) culminate in this period, multiplying and
diversifying greatly, and far surpass in size the Carboniferous and
Permian genera, and then become extinct.
It is in the =Reptilian= class that we find the most remarkable changes.
The abundance and diversity of the reptiles in this and the two
succeeding periods are incomparably greater than those in the Permian
age. The Triassic rocks have representatives of almost all the orders of
Mesozoic reptiles, though often these are comparatively small and rare
forms.
The reptiles with mammalian characters (Theromorpha), which first
appeared in the Permian period, culminate in the Triassic, especially in
southern Africa, and then become extinct. No birds, or reptiles which can
be regarded as the ancestors of birds, have been found in this period.
In this period one of the greatest advances in the progress of life is
indicated by the first appearance of _low mammals_,—_mammals having very
decided reptilian characters_, and belonging to the orders of generalized
=Monotremes= and =Marsupials= (_Dromatherium_; _Microlestes_).
=Jurassic Period.= The _Fishes_ have advanced greatly beyond those of
the Triassic rocks. The Sharks have advanced practically to their modern
condition, and a new order of Elasmobranchs, the broad and flat _Rays_,
are introduced.
The _Chimæroids_ occur in this period, and were more numerous than in
modern times. Dipnoans have become very scarce. The Crossopterygians are
greatly reduced. The Ganoidei are still the dominant type; some of these
latter approximate the Teleosts so closely that it seems arbitrary to
call them Ganoidei. In Europe the _Reptiles_ culminate in this period
and show extraordinary development in variety, huge size, number and
degree of organization. They were rulers in every department of nature.
They were the rulers in the air in place of birds; rulers on the land in
place of mammals; and rulers in the sea instead of sharks and whales.
Immense land reptiles (_Dinosauria_) as large as our largest mammals,
and in some cases larger than the elephant, moved sluggishly over the
land. Some walked on all fours; others were occasionally or usually
bipedal, and walked upright like birds, and had many structural features
in common with the latter; some were herbivorous, feeding on plants
and even reaching into the branches of trees for their food, others
were carnivorous, feeding on their fellow-creatures. Huge reptiles
(_Ichthyosauria_) swam about in the sea in great numbers. Immense
bat-like forms (_Pterosauria_) sailed through the air like birds, being
literally flying dragons.
These reptiles, which had branched off from the generalized Amphibians,
were themselves very generalized creatures. It is interesting to remember
that the Monotreme and Marsupial mammals found in this age were very low
reptilian mammals; they were not typical, specialized Monotremes and
Marsupials, like the modern creatures, but very generalized forms, being
probably connecting links with low generalized Insectivora, which were
the first of the true mammals.
These early Monotremes and Marsupials were quite small animals, varying
in size from a mole to a rabbit. They were insignificant creatures among
the mighty giant reptiles, but they carried in their warm blood the
promise of future mammalian supremacy.
Birds appear for the first time in this period, but very different
from modern birds. Not only did many of the reptiles of this time have
bird-like characters, but all the birds of this period were distinctly
_reptilian_ birds (_Archæopteryx_). Birds came off from primitive
reptiles as branches from a stem (see Diagram of Development). It was
only with the passing epochs that these early birds changed more and more
from their reptilian characters and assumed more and more the features
of modern birds. Many fossil connecting links between reptiles and birds
have been discovered, and afford most useful illustrations of those
changes in creatures that we call _Evolution_.
=Cretaceous Period.= In the Cretaceous period the aspect of plant life
has changed greatly. Now, for the first time, we meet with ordinary
hard-wood trees, such as beech, oaks, hickory, maples, poplar, etc., but
of very different genera and species from those existing to-day.
In this period, for the first time, we find the highest order of
Cephalopods, viz., the naked ones, allied to cuttlefishes and squids.
They are known as the _Belemnites_. Sometimes the fossil ink-bags of
these creatures are found so that they can be drawn in their own fossil
ink.
The Ammonites proper, which were introduced in the Triassic period
and culminated in Jurassic time, become extinct at the close of the
Cretaceous.
The =Vertebrates= form the most characteristic features of the Cretaceous
fauna. A revolution has occurred among the Fishes. Sharks of modern
type are numerous. Crossopterygians and Ganoidei are rare. There has
been an immense expansion of _Teleosts_ or Bony Fishes, which now become
the dominant fishes. Most of the Cretaceous Teleosts belong to modern
families and even genera.
The _Reptiles_, in this period, continue to be the dominant types of the
land, air, and sea, and it is difficult to decide whether the Cretaceous
or the Jurassic is to be regarded as the culminating period of Reptilian
history. The flying Reptiles (Pterosauria) of this period are remarkable
for their great size, far exceeding, in this respect, those of the
Jurassic time. The land Reptiles (Dinosauria) are in greater numbers than
in the preceding period. The sea Reptiles are less numerous than in the
Jurassic, but are of greatly increased size.
Cretaceous _Birds_ are much more numerous and advanced than the Jurassic
ones.
The _Mammals_ of the Cretaceous are much more abundant and varied than
those of Jurassic times, but they are nearly all of very small size, and
continue to play a very modest rôle. The lower Cretaceous mammals differ
but little from those of the Jurassic, except for the larger number of
genera. The mammals of the latest Cretaceous time are much more numerous
and diversified than those of the early Cretaceous. They also show
affinities with the mammals of the next succeeding or Tertiary period.
The birds and mammals, though they have varied with the passing time and
changing surroundings like all other living creatures, are very different
from those around us now. They have thrown off some of their reptilian
characters, yet they are still distinctly and pronouncedly reptilian.
The Triassic, Jurassic and Cretaceous periods together constitute the
Mesozoic Era, or =Age of Reptiles=; for never before and never in
succeeding ages were reptiles so huge, and varied, and masters in all the
realms of nature.
=Tertiary Period.= In reaching Tertiary time we enter upon the threshold
of our modern world. The reptiles have dwindled to a few low forms,
such as alligators and crocodiles, to the small lizards, to turtles and
tortoises, and to those low-grade reptiles, the snakes, which now for the
first time are the most numerous of the class. In these ages, as now, the
Teleost fishes vastly predominate, and the Ganoid and Crossopterygian
fishes are nearly extinct. In Tertiary times, as in the present, all the
reptilian birds had disappeared, and only typical birds remain. In other
words, the bird-class had now separated from the reptilian class, and the
connecting links became extinct.
In this age for the first time the highest flowering plants are abundant,
and now for the first time also the highest orders of insects are
abundant, such as bees, ants, butterflies, etc. On account of the greater
warmth and moisture in the Tertiary period, insect and plant life were
fuller then than now. That the climate was warmer in those times is shown
by the following facts: In America, during the early Tertiary period,
figs, evergreens and palms grew in Dakota, showing a temperature there at
that time equal to the temperature in Florida at present. In the middle
Tertiary period trees like the Redwood of California, and Magnolias,
were abundant in Greenland.
Although _Reptilian Mammals_ (_Multituberculata_) lived in the preceding
era, true or typical Mammals first came to view in the earliest Tertiary
times. They soon succeeded the vanished giant reptiles as the rulers
of the world; so that this and the succeeding (Quaternary) period are
known as the =Age of Mammals=. At this period, for the first time,
appear generalized _primitive flesh-eating mammals_ (_Creodonta_), and
_primitive grass-eating ones_ (_Condylarthra_ and _Amblypoda_); also
_primitive Primates_ (_primitive Lemuroids_, and, later, _primitive types
of Monkeys_, e. g., _Anaptomorphus_).
The evolution of mammals, compared with that of other animal groups, has
been so rapid that each stage of even the Eocene has its own mammalian
fauna, differing from those of the succeeding and preceding stages. All
through the Tertiary period, as through all the preceding ages, the
species that are advancing in life are undergoing greater and greater
specialization. Animals at first closely related finally become more
and more separated from one another. This is well illustrated by a
study of the hoofed animals. In the earliest Tertiary period all these
animals seem to unite into one branch but now they consist of many widely
separated sub-branches. As we trace the branch down to the earliest
Tertiary period, we find that even in early Eocene times it divides into
the even-toed and odd-toed ungulates. In the later Miocene times, each
of these again separates,—the former, even-toed ones, into the hog and
hippopotamus families with four toes, and the ruminant family with two
toes; the latter, odd-toed ones, into the elephant family with five toes,
the rhinoceros and tapir families with three toes, and the horse family
with one toe.
[Illustration: FIG. 14.—Illustrating genesis of horse’s feet. Fore-feet
(a) and hind-feet (b) of Orohippus; fore-feet (c) and hind-feet (d) of
Mesohippus; fore-feet (e) and hind-feet (f) of Miohippus; fore-feet (g)
and hind-feet (h) of Protohippus; fore-feet (k) and hind-feet (l) of
Pliohippus; fore-feet (m) and hind-feet (n) of Equus.]
It will be exceedingly instructive to trace briefly the progressive
specialization of the horse family through the Tertiary period. A
wonderful series representing this family has been found in the American
Tertiaries. First of all, the =Eohippus= is found in the earliest Eocene
rocks. This little animal was about the size of a fox and had four
perfect toes (hoofed) and a fifth, much smaller, imperfect one (splint)
on the fore-feet; the hind-feet had three perfect hoofed toes. Later on,
in the middle Eocene rocks, was found the =Orohippus= (Fig. 14), with
three toes behind (b), and four in front (a)—the fifth imperfect toe
(splint) being lost. This animal was also about the size of a fox. Still
later on were found, in the Miocene rocks, the =Mesohippus= (c, d) and
the =Miohippus= (e, f), with three toes behind (d, f) and three toes and
a splint in front (c, e). The splint in the Miohippus is much smaller
than in the Mesohippus, and also in the former the two side toes have
become smaller and farther removed from the ground. Both of these animals
were about the size of sheep. Further on, in the early Pliocene rocks,
appears the =Protohippus= (g, h), with three toes on all the feet, but
the middle toe considerably larger and longer than the side toes. This
animal was about the size of an ass. Then, finally, came the =Pliohippus=
(k, l) and =Equus= (m, n) in the latest Pliocene rocks. Here the middle
toe is greatly enlarged and the side toes are reduced to useless splints.
Thus by degrees, through slight modifications not only of the feet, but
also of the skull, teeth, brain, etc., and by adaptations from age to
age, was formed the modern horse. In like manner, by gradual changes and
adaptation to surroundings, all the early generalized mammals pass into
the higher specialized ones.
=Quaternary Period.= The mammals of this age differ from those of the
Tertiary period and from living species. Never before this period,
and never since, have mammals been so large. They culminate in this
period and then decline. The great mastodon, the great cave-bear, and
saber-toothed tiger, the Irish elk, and gigantic sloths and armadillos
lived in this age.
Most interesting of all, the remains of _man_ now first appear associated
with these extinct animals. Man must have been at first an apparently
insignificant creature among the mighty mammals that surrounded him, and
must for a time have contended more or less doubtfully with them for
mastery. But as he increased in numbers and intelligence he became more
and more the ruler of the brute creation; so that this age in which he
rules may fittingly be spoken of as the Psychozoic Era, or =Age of Man=.
[Illustration: FIG. 15.—A snake and two lizards; a, snake; c, Bipes
(lizard); b, Cheirotes (lizard).
From Shaler’s First Book in Geology. By courtesy of the publishers, D. C.
Heath & Co.]
Fig. 15 is introduced to show how easy may be the transitions from one
order of animals to another. Two long, slender Lizards are shown and
also a Snake. Bipes has four limbs, but they are very small and weak.
Cheirotes has lost, probably through disuse, the hind-limbs and the front
ones are small and weak, almost useless. The snake has no limbs at all.
The two lizards are quite snake-like in external forms. Just as changing
environment, use and disuse, etc., can effect transitions in external
forms by slight gradations, so also may the whole internal structures
undergo marked alterations by slight modifications through long ages.
The brief references we have made to the changing life-forms during the
geologic ages, should be very instructive even to a reader not familiar
with zoölogy. They teach the important lesson that life, in the main, has
ever advanced to higher and higher levels. At one time in the history of
the earth no animals were in existence higher than the Invertebrates;
these lived through the ages with ever-changing forms. Later, in the
course of geologic history, some of the primitive Invertebrates evolved
into the lowest Vertebrates, and Fishes finally appeared, at first
very primitive and generalized. For long ages there were no creatures
on the globe higher than these. Then as the æons slowly rolled on some
of the early fishes evolved finally into a higher group of animals,
the Amphibians, and these for long ages were the highest creatures in
existence. As time slowly passed, some of the primitive amphibians
evolved into primitive Reptiles, and these creatures for long ages were
the highest in existence and monarchs of the world. Some of the primitive
reptiles branched off in one direction leading to modern Birds, and some
advanced in another direction that led through primitive mammals to man
and other modern mammals. And now in modern times man is monarch among
the animals.
We may now begin to realize that truly with the changing and growing
continents during the geologic ages, many life-forms have varied and
advanced to higher levels, culminating in the final appearance of _man_
as well as other living creatures.
SECTION V.
NATURAL SELECTION.
NATURAL SELECTION.
In briefly outlining the transmutations of living forms that took place
during the Geologic Ages, we have said that changing conditions of
life—through an ever-shifting environment—have compelled modifications
in the form, structure, and habits of living creatures; and that those
creatures which were unable to _adapt_ themselves, through _useful
variations_, to the altering conditions of life have perished, while
those that _did_ adapt themselves lived and progressed in organization.
What, then, is the great agency through which some life forms have been
_eliminated_ during the ages, while others have been _selected_ to
continue through these ages? This agency is =Natural Selection=. The
phrase “natural selection” is simply a convenient, condensed statement of
observable and easily verifiable facts, viz.: that animals and plants are
so situated in this world that they can only secure their food and mates
by work, by effort, by struggles, whether consciously or unconsciously,
and whether directly or indirectly; and that in these struggles those
that are best equipped for their life duties are the ones that are
most naturally successful in living and procreating their kind. The
survival of those best adapted to their environment may be spoken of,
in the language of Spencer, as the =Survival of the Fittest=. Darwin’s
phrase—=Natural Selection=—has precisely the same significance and means
that those creatures which are best fitted to their surroundings are the
survivors. The working of Natural Selection may be made clearer by a
brief reference to the =Artificial Selection by Man= of various animals
and plants. Variations frequently take place in domesticated animals and
plants. Some of these variations appeal to man as being of practical
value, others as beautiful, and others again as curious or interesting.
He selects those individuals whose variations he wishes to preserve,
breeding only them together, and in this way accentuates those variations
he desires to perpetuate. In course of time the accumulation of these
differences becomes so marked as to make the animals differ greatly from
the original stock from whence they came. A well-known illustration of
this process is the pigeon. All our domestic pigeons, forming a large
number of well-marked races, such as the fantail, the tumbler, the
pouter, etc., have been produced from the ordinary wild rock pigeon
of Europe. The bird fancier, noticing individual differences in the
offspring of the wild rock pigeon, selected the peculiar individuals, and
bred only them together. By this simple process of artificial selection
and isolation of the chosen or selected individuals, all the races of
pigeons have been produced. Plate I shows the wild rock pigeon _Columba
livia_ and the domesticated pouter. The forms of these pigeons are very
different, yet the wild rock pigeon has been transmuted into the pouter
through the agency of Artificial Selection. The same is the case with
our pigs, dogs, cats, apples, grapes, and other domesticated animals and
plants. Fig. 16 shows the domesticated pig that has been derived from
the wild boar by Artificial Selection. By this process, such distinct
races as the Newfoundland, the Skye Terrier, and the Bulldog have been
produced—creatures that have all come from common ancestors, yet so
different looking, one from the other, that if they had been found in the
wild state, they would not only have been ranked as distinct species, but
as even distinct genera. The same method has given us different races
of horses, cows, sheep, flowers, grains, etc. The swiftest horses, for
instance, are selected to breed together; then the fleetest offspring
of these, time after time, until horses are produced whose speed far
surpasses that of the originally selected pair from whence they were
derived. Darwin has taught us that what man does on a small scale, in
a comparatively short time, Nature has been doing on a vast scale for
long ages and has thus given rise, from simple forms, to the infinite
variety and complexity of animal and plant life, as we behold it on the
globe to-day. The selection by man of useful variations in domesticated
creatures being appropriately called Man’s Selection, or Artificial
Selection; the vastly greater selection by nature of animals and plants
with useful variations and on an infinitely grander scale, through
inconceivably long ages, is most fittingly called =Natural Selection=.
The struggles—of animals, for instance—that necessarily lead to the
survival of the fittest, are intensified and made exceedingly acute and
severe by the fact that _all animals tend to increase in a geometrical
ratio, and by the further fact that the food and place for animals are
limited_. In other words, the population of the animals in a given area
tends greatly to outrun the means of subsistence. And since animals are
constantly varying in many directions and are as plastic in the hands of
Nature as clay under the chisel of the modeler, those that possess any
useful variations, whether congenital or acquired, that give them any
advantages in this great battle of life, will most likely come out of the
struggle as victors.
[Illustration: FIG. 16.—Wild Boar contrasted with a modern Domesticated
Pig. Reproduced from Romanes’ “Darwin, and After Darwin.” By courtesy of
The Open Court Publishing Company.]
=Multiplication of Animals.= Even in the slow-breeding elephant, the
offspring tend to increase threefold in each generation. Some animals
tend to increase twenty or thirty fold in each generation, while still
others tend to increase a thousand fold or even ten thousand fold. If all
the offspring of the elephant lived, in eight hundred years there would
be over nineteen million elephants alive. If the eight million eggs which
the roe of certain fishes, such as the cod or the eel, contains, were to
develop into adult forms, the ocean would quickly become a solid mass.
The aphis or plant louse is so very prolific that it has been estimated
that the tenth brood of one female alone would contain more ponderable
matter than all the population of China,—estimating this population at
five hundred millions. Yet, in spite of this tendency on the part of
animals and plants to increase in numbers at such a stupendous rate,
it is found that, in any given area, the conditions of which are not
changing, the number of the animals and plants remains fairly constant.
This is because of the fact that, _along with the stupendously large
birth rate, there is an equally stupendous death rate_. This high death
rate is to a large extent indiscriminate, for it involves those that are
physically fit to live, as well as those that are unfit to live. At the
edge of a coral-reef, free-swimming, active embryos are found in immense
numbers. After a while some of these settle at too great a depth in the
water or on the muddy bottom, and die; others get into a more suitable
position and live. Again, whole nests of bees are destroyed by the
badger; tongue loads of ants are engulfed at one gulp by the ant-bear;
hundreds of thousands of fry are destroyed by the Greenland whale at
one swallow. In all these cases, the destruction is indiscriminate,—the
good, bad, and indifferent are alike decimated, but in spite of this
wholesale and indiscriminate destruction, keeping down, as it does, the
stupendous birth rate of living creatures, _more animals and plants are
born than are required to keep up the normal number of individuals that
can be supported in a given area. Among these creatures there arises a
struggle for existence, a struggle for food and place, and those that
are the best fitted to live come out of the contest as conquerors._ In
this struggle those creatures that emerge as victors, on account of
having been best adapted to their conditions of life, may be spoken of
as having been selected by Nature; or they may be spoken of as chosen by
Natural Selection. In this great struggle, over which Natural Selection
presides as some _inexorable, ever-watchful, sharp-eyed task-master_,
the victory is to the cunning instead of the stupid; the race is to the
swift instead of the slow; and the battle is to the strong. The wolves
of keenest scent, the tigers of more supple spring and sharper sight,
secure their prey and thrive, while the weaker members fail to get their
food and starve. During migration the birds that are strongest on the
wing reach the land whither they are flying, while the weaker perish
on their course. Thus Natural Selection acts in two ways, _eliminating
the unfit_ and _selecting the fit_, and there are, besides, two special
modifications of Natural Selection which are called =Sexual Selection=
and =Insect Selection=.
The following analysis of Natural Selection may be useful to the reader,
viz.:
{ Physical and Climatic Causes.
{ Elimination { Enemies.
{ { Competition.
{
Natural Selection { { Natural Selection Proper.
{ {
{ { { By Preferential
{ { { Mating.
{ Selection { Sexual Selection. {
{ { By Battle.
{ Insect Selection.
=Elimination of the Unfit.= The elimination of the unfit takes place
through the agency of physical and climatic causes and also through
enemies, and competition of members of the same species. Elimination
through the action of surrounding physical and climatic conditions is
shown by the following facts: if certain tropical animals be transferred
to sub-Arctic or even temperate regions they are unable to adapt
themselves to the requirements of the new climatic conditions, and die
sooner or later; many animals are killed if the fresh-water lake in
which they live be invaded by the waters of the ocean; fishes which
live at great depths in the sea and are, therefore, subject to great
pressure, are killed when they are brought to the surface, on account
of the expansion of the gases in their tissues; if the water where
corals are living becomes too fresh, too muddy, or too cold, they will
die. The change of climate to a much colder temperature at the close
of the Jurassic Ages was probably the cause of the extinction of the
huge reptiles that took place at that time. In the winter of 1854-5,
four-fifths of the birds in Darwin’s grounds perished on account of the
severity of the cold.
As to elimination by enemies, it is well known to naturalists that
throughout nature battle within battle is continually recurring with
varying success. When weaker animals are preyed upon by stronger ones,
and self-defense is useless, the bulky and slow animals are eliminated,
while the swift and agile ones escape; the stupid are destroyed, while
the cunning often survive. As to elimination by competition, the
stronger animals kill the weaker ones, and then quarrel and fight with
one another over the prey, the strongest, etc., getting the food. While
weaker animals are being preyed upon by various enemies, and are thus
eliminated, these enemies are also competing with one another for the
prey. At the same time that the stupid and slow creatures are being
destroyed by their captors, thus leaving the more cunning and agile
animals in possession of their habitat, the stupider and less active
captors are gradually eliminated by competition, through failing to
capture their more agile and cunning prey.
The agency of Natural Selection in bringing about the innumerable
adaptations of animals and plants, and, therefore, causing transmutations
of living creatures, can most interestingly and instructively be
illustrated by a study of the _coloration of animals and plants_.
As a correct idea of the mode of operation of Natural Selection, and the
pronounced results attained thereby, is very desirable, it will well
repay the reader to study _in extenso_ the relations of _color-patterns_
to _environment_. The whole subject is nothing but repeated
illustrations of one principle, viz.: Natural Selection. This fact and
the interest of the subject will justify the numerous details.
=The Coloration of Animals and Environment.= The colors of animals often
harmonize most wonderfully with their surroundings. Thus green is a
common color of animals in the evergreen forests of the tropics; white is
the prevailing color in the arctic regions; and a yellowish hue in desert
places. In the evergreen forests of tropical America, whole groups of
birds are found whose fundamental color is green; there the parrots and
fruit-eating pigeons are commonly green; the bee-eaters, leaf-thrushes,
and many other birds, have so much green in their color as to add greatly
to their concealment in the dense green foliage. In the desert places,
the lion, the desert antelope and the camel harmonize with the color of
the rocks and sand among which they live. In the Arctic regions the polar
bear, the Greenland falcon, and the American polar hare are white. We
have further the dusky hue of creatures that haunt the night, such as
mice, moles, and bats; and the gorgeous tints of fishes that swim among
the coral reefs. These local color adaptations of animals are of great
use to them, either enabling them to escape the notice of their enemies
or to come upon their prey with the least risk of being detected. Certain
groups of animals have a local color-adaptation, and may be noticed under
the heading of _Protective Coloration_; others have acquired a wonderful
resemblance to surrounding inanimate objects, such as leaves, twigs,
bird-droppings, flowers, etc., and may be described under the heading of
Protective Imitation of particular objects, or _Protective Resemblance_;
closely allied to the latter are those resemblances to surrounding
objects which are not so much for the purpose of protecting the animals
from enemies as for attracting their prey, and these resemblances,
therefore, will be described under the title of _Alluring Coloration_.
Other animals have such color patterns as to be very conspicuous in their
surroundings; these animals are usually very poisonous or possess other
deleterious qualities that cause beasts of prey to avoid them; hence
these animals may be said to possess _Warning Coloration_. Many animals
that are very desirable food for carnivorous creatures have acquired,
in past ages, a remarkable resemblance to these dangerous animals with
warning coloration, and are treated of under the title of _Mimicry_.
There are still other groups of animals whose peculiar coloration enables
the member of a herd or flock which may have become separated from the
herd to readily recognize its companions at a distance as friends, and
thus distinguish them from enemies. The color patterns of these animals
may be classed under the heading of _Recognition Marks_.
=Protective Coloration, or Local Color Adaptation.= In forest-haunting
animals of large size, such as forest-cats, and forest-deer, rounded
spots are frequently noticed. Animals like the tiger, that spend a great
deal of their time among high grasses and reeds, are striped vertically.
The combined artistic effects of these spots and stripes, in connection
with the lights and shades of the forest and the reeds, are such as quite
effectually to conceal the animals from view: for the black stripes of
the tiger, for instance, correspond with the black shadows of the reeds
or grasses; and his yellow stripes with the yellow of the reeds. In like
manner the rounded spots of the forest-deer harmonize with the spotty
shadows of the leaves in the forest. An experienced tiger hunter has
stated that in following up a wounded tiger the natives saw the animal
at a distance of about twenty meters, under a tree among the reeds, and
pointed out the animal to him; but the color effects of the stripes of
the tiger so harmonized with the artistic effect of the light and shade
of the reeds as to effectually prevent his seeing the animal for a minute
or so. The zebra, which is such a conspicuous animal in our zoölogical
gardens, on its native soil and in a bright star-light night, may be
so close to one as to be heard breathing, and yet cannot be seen, so
completely are its color patterns in harmony with its native habitat.
Marine organisms that float on the surface of the water are beautifully
tinged with blue in harmony with the color of the ocean as it appears to
their enemies above, the birds; while, looked at from below, they are
white, thus harmonizing with the white clouds and the foam as seen by
enemies from below.
There are many animals that are very conspicuous when removed from their
native haunts, yet when in their proper environment are invisible or
detected with the greatest difficulty. Such a large animal as the giraffe
is effectually concealed by its form and color when standing among the
broken and dead trees that exist on the edges of the thickets where it
may be seeking its food. The odd shape of the head, with its horns that
resemble broken branches, and the blotchy spots on the skin, so harmonize
with its surroundings that even the keen eyes of the natives sometimes
mistake giraffes for trees and trees for giraffes.
There is a bat (_Kerivoula picta_) found in the island of Formosa that
has a very conspicuous black and orange color. The body of this bat is
of an orange color, and its wings are black and orange-yellow. When
resting it suspends itself, head downwards, from the branches of an
evergreen tree. During all the year some part of the foliage of this tree
is undergoing decay, so that many of the leaves assume tints of orange
and black. When the bat is suspended among such decaying leaves its
colors and those of the leaves so harmonize that the animal is perfectly
concealed, and thus eludes its enemies.
The sunbirds of Africa are very conspicuous when out of their natural
environment, being brilliant and gorgeously colored. These birds find
their main food supply among plants that have very conspicuous flowers;
the aloe-blossoms, especially, which they frequent, are brilliantly
colored. The colors of these birds so completely harmonize with the gay
colors of the blossoms that even the keen eye of the hawk is unable to
detect them. One species of these birds, the black sunbird, is never
absent from a forest tree known as the Kaffir boom. This tree has not a
single green leaf on it, but consists of a great mass of purplish-black
and scarlet blossoms. A dozen of the black sunbirds may be feeding in
this tree, and their notes may be heard among its branches all the day,
yet their color adaptation to their environment is so complete that they
are seen with the greatest difficulty or are entirely invisible.
Those birds whose colors have varied in such a way that they have not
harmonized with their surroundings,—that have not become adapted to
their environment,—are seen by the hawks and are exterminated; but those
whose colors were more favorably arranged and more in harmony with their
surroundings have been most often undetected by their enemies, the hawks,
and have lived and transmitted their useful color patterns by heredity
to their offspring. This process going on, age after age, has developed
such perfect adaptations of the sunbirds to their environment as the
naturalist observes at present.
These examples are illustrations of the fact that those creatures that
are most in harmony with their surroundings are the ones that live and
procreate their kind. They are cases of the survival of those animals
that are best adapted to their environment,—the survival of the fittest;
the selection by nature of favored creatures; in short, Natural Selection.
Among our native birds, the woodcock and snipe have such tints and
markings as strikingly to harmonize them with the dead marshy vegetation
which constitutes their native haunts. The ptarmigan in winter has a
light coloration in harmony with the environment of snow, while in
its summer plumage it is tinted and mottled in harmony with the color
effects of the lichen-covered stones among which it spends a great deal
of its time. Young unfledged plovers are spotted in such a way as very
accurately to resemble the beach pebbles among which they remain for
protection.
The white-headed fruit pigeon (_Ptilopus cinctus_) is a very conspicuous
bird when taken from its native haunts. It has a pure white neck and
head, black back, yellow belly, and well-marked, deeply-curved black band
across the breast, and black wings. It is a handsome as well as very
conspicuous bird. It frequents trees that are a species of Eucalyptus in
the island of Timor, and which have very open foliage and yellowish or
whitish bark. The pigeons may be sitting motionless on exposed branches
of the tree during the glaring heat of the day. The yellow and white bark
of the tree, the deep blue sky seen through the openings of the leaves,
with the intense tropical sunlight casting black shadows of one branch
upon another, make color effects that harmonize so completely with the
color patterns of the pigeons that they are entirely invisible.[8] Here
again the complete color adaptation of the pigeons to their environment
has been accomplished through the agency of Natural Selection,—Nature
selecting those pigeons most frequently that varied most in the direction
of useful color patterns, by concealing them from their enemies, the
hawks.
There are some birds that are not protectively colored, which therefore
do not possess a local color adaptation. The common raven, which inhabits
the Arctic regions, is black instead of being white. The raven is found
as far north as any known bird or mammal. It is a powerful and fearless
bird, and needs no protective coloring; and since it feeds on dead
animals it needs no concealing coloration to enable it to secure its food.
The bright and conspicuous coloration of many birds’ eggs have often
been looked upon as a difficulty on the theory of adaptive coloration.
But Wallace thinks that a careful consideration of the subject in all
its bearings shows that in a great number of cases these colorations
are instances of protective coloration. He further thinks that when we
cannot see the meaning of the particular colors, we may suppose that in
some ancestral forms they have been protective, and, being harmless,
have persisted under changed conditions which rendered the protection
needless. He states, in illustrating the protective coloration of
eggs, that “the beautiful blue or greenish eggs of the hedge-sparrow,
the song-thrush, the blackbird, and the lesser redpole, seem at first
sight especially calculated to attract attention; but it is doubtful
whether they are really so conspicuous when seen at a little distance
among their surroundings. For the nests of these birds are either in
evergreens, as holly or ivy, or surrounded by the delicate green tints
of the early spring vegetation, and may thus harmonize very well with
the colors around them. The great majority of the eggs of our smaller
birds are so spotted or streaked with brown or black on variously tinted
grounds, that when lying in the shadow of the nests and surrounded by the
many colors and tints of bark and moss, of purple buds and tender green
or yellow foliage, with all the complex glittering lights and mottled
shades produced among these by the spring sunshine and by the sparkling
raindrops, they must have a quite different aspect from that which they
possess when we observe them away from their natural surroundings.” We
have here, probably, according to Wallace, a similar case, of general
protective harmony, to that of the green caterpillars with beautiful
white or purple bands and spots, which, though gaudily conspicuous when
seen alone, become practically invisible among the complex lights and
shadows of the foliage they feed upon.
Eggs that are not protectively colored are usually white or of some
uniform pale color, and are concealed by the birds in covered nests or in
holes in trees or in the ground. Many birds lay white eggs in open nests,
but even here the devices for concealing them are very effective and
interesting. In some cases, such as the partridge, the goat-sucker, and
others, the birds have the habit of sitting close and almost continuously
on the eggs. These birds are protectively colored. Ducks, pheasants,
and other birds have the habit of covering their white eggs with dead
leaves or other material when they leave the nest. There are some large
and powerful birds that lay conspicuous white eggs in open nests. Such
are the cormorants, herons, storks, pelicans, and others. But they guard
their nests carefully, and are able to drive away any enemies.
It is thus seen that there are many devices by which birds’ eggs are
protected, and these have all been developed through the agency of
Natural Selection. For example, those exposed eggs of timid birds that
varied most in the direction of protectively colored ones were the
ones that were most likely to be overlooked by egg-eating creatures,
and those that were least protectively colored, and therefore the most
conspicuous, were the ones most likely to be destroyed. The protectively
colored eggs being thus, in the nature of things, the ones that have
suffered less destruction from enemies, have developed into birds that
have transmitted, by heredity, their characteristics to succeeding
generations of birds’ eggs. And thus in the course of time have many of
the colorations of eggs been developed from simple to elaborate patterns.
Many snakes, frogs, butterflies, caterpillars, and so on, are colored
green in harmony with the foliage of the trees among which they live. By
being colored green these animals are more or less effectually hidden
from their enemies, such as carnivorous birds. If they were not concealed
by protective coloration they would quickly be exterminated by hungry and
greedy enemies. The green frogs are greatly protected by their colors
from their enemies, the green snakes. If they were not so colored, they
would quickly be exterminated by these snakes. Even protectively colored
as they are, large numbers of them are caught by the snakes, which are
also protectively colored and keener witted. It may be stated that the
protectively colored frogs are hunting for their food, the protectively
colored insects; but, at the same time, they try to avoid their
protectively colored enemies, the snakes, though they are often caught;
the protectively colored snakes are hunting for their food, the frogs,
but, in their green surroundings, endeavor to avoid their enemies, the
reptile-eating birds, though often unsuccessful. And so on, from day to
day, the tragedies of the animal world are enacted.
It may be well to pause a moment here and ask why it is that animals are
not completely exterminated which are thus perpetually being persecuted
and preyed upon by stronger foes. Why, for instance, are any frogs left
in a green locality, when they are so eagerly sought for by their
enemies, the snakes? What are the factors that permit the average or
normal number of them to exist in the given locality? The most important
of them have already been referred to separately, but it will be useful
to recapitulate, briefly, a few of them now. One of the most important
of them is the tendency of the frogs to increase in geometrical ratio.
In the breeding season great numbers of young are brought into the
world, thus reinforcing the depleted ranks of the adults. Another factor
is the instinctive effort of each frog to avoid its enemies, and their
protective coloration greatly facilitates their concealment and escape.
Another factor is the circumstance that the snakes also have enemies.
They must be wary and careful in hunting prey lest they be unduly
exterminated by their foes, the snake-eating animals. Then another factor
is the alternating day and night, by which the warfare, offensive and
defensive, must be periodically checked. Then again, the hibernating
season, winter, checks or rather suspends the life-destroying crusades.
It may thus be recognized that many complex factors occur to explain the
circumstance that a given locality generally contains the normal number
of individuals that constitute the species of frogs.
The complex factors at work in these relations and inter-relations of
animals and plants are so nicely balanced that with a comparatively
stable environment the number of species of the different classes of
animals may remain more or less constant. But suppose one of these
factors is profoundly diminished, such, for instance, as a great
diminution in the number of the snake-eating birds. Then the snakes,
not being much persecuted by their enemies, will be in a most favorable
environment; they will increase greatly in numbers; they will be eagerly
on the look-out for their food, the frogs; the frogs, being thus
unusually persecuted and hard-pressed, will diminish greatly in numbers.
Now, this will be a golden opportunity for the insects; their environment
becomes much more favorable. The snakes do not bother the insects, but
destroy their enemies, the frogs. Therefore, the insects increase greatly
in numbers and profoundly affect the vegetation upon which they live
and thrive. The injury to the vegetation may thus seriously affect an
altogether different class of animals—the grass-eating or foliage-eating
animals, like the ruminants.
It may thus be understood how profoundly complex are the relations and
correlations of living creatures, and how a disturbance of some of the
links in these living chains may very extensively affect the other links.
The different creatures referred to above are probably the simplest
illustrations that can be given of protectively colored animals. For
here we have green animals adapted to a green environment—the green
foliage of trees or grass. This adaptation has been brought about by
Natural Selection. Suppose, for instance, that the habitat of the
_non-protectively_ colored _ancestors_ of the frogs was for any reason
unduly crowded by their enemies, the snakes. The frogs would be hard
pressed and much persecuted. Suppose that among the young of these
frogs there were some that varied in the direction of grass-green or
leaf-green colors, while others did not so vary, or possibly varied in
the direction of even conspicuous colors, such as black or white. It is
evident that the latter groups would be easily detected by the snakes and
destroyed, while the green group would frequently escape notice and would
thrive and procreate their kind. This process being repeated generation
after generation, there would come a time when the given habitat would
contain none but frogs with a protective coloration of green. All of
these protective colorations of snakes, butterflies, and caterpillars,
as well as of frogs, are adaptations. They are illustrations of the
survival of the fittest; the survival of those best adapted to their
environment—in short, Natural Selection.
[Illustration: PLATE II.—_Kallima paralekta._ A butterfly of Sumatra
illustrating the work of Natural Selection: C, butterfly with expanded
wings (dorsal surface) which are conspicuously colored through Sexual
Selection; B, same butterfly with wings closed (ventral surface) and
presenting a close resemblance to a dead leaf, A, through the agency of
Natural Selection. B, illustrates the Protective Resemblance of an animal
to an inanimate object.]
=Protective Imitation of Particular Objects.= Insects often exhibit a
very great amount of detailed resemblance to the leaves, flowers, and
twigs of plants among which they live. Those that live on grass are
striped longitudinally, while those that feed on ordinary leaves have
an oblique striation. There is a larva of a Georgia butterfly (_Sphinx
fuciformis_) which feeds on a plant having small blue flowers and linear,
grass-like leaves. This larva has a blue head and a green body striated
like the leaves. The resemblance of the insect to its environment is very
striking. There is another species that feeds on a plant with small red
flowers situated in the axils of the leaves, and this larva has a row of
seven red spots of unequal size, which corresponds quite closely with
the size and color of the flowers. There is a caterpillar in Borneo that
resembles a piece of moss with two exquisite pink-white seed-capsules.
Its general hue is greenish, with two little pink spots on its upper
surface, and it is covered with hair. Its movements are very slow, and
when eating it withdraws its head beneath a mobile fleshy hood that it
possesses, so that its motions in feeding are not noticeable. When living
in its native haunts it is all but impossible to detect it, so completely
does it resemble the surrounding moss. Other insects resemble green or
dead leaves in all their varieties of form and color, and to show what
a great protection such resemblance affords to insects in concealing
them from view the following observation of a naturalist in Nicaragua
may be related. In that country there are armies of foraging ants that
devour every insect they can catch. Among a multitude of these ants he
observed a locust that looked very much like a green leaf. The ants, many
of them, were continually running over the body and legs of the insect
without detecting its character. In many parts of the world there are
many butterflies (_Kallima_, for example) the under surfaces of whose
wings very closely resemble dead leaves. They frequent dry forests and
are rapid flyers. They are rather large, and the upper surfaces of their
wings are quite showy, having bluish and orange colors (Plate II). It is
their habit always to settle on some twig where there are decaying or
dead leaves. In doing so it folds its wings together over the back, thus
concealing the gay upper surfaces and presenting the protectively colored
under surfaces. The resemblance to a dead leaf is much more striking from
the fact that the short tails of the hind wings just touch the branch on
which the insect rests and look very much like the stalk of a leaf. From
this stalk a dark curved line extends to the elongated tip of the upper
wings, thus imitating the midrib of a leaf. On both sides of this midrib
are oblique lines that are partly markings and partly nervures, which
give the appearance of a leaf with its veining. The head and antennæ
fit in such a way between the closed upper wings as not to interfere
with that irregular outline which is characteristic of the withered and
dry leaves. Often the closed wings are covered with small black dots
gathered into circular groups that exactly resemble the minute fungi
found on decaying leaves, and it is sometimes difficult to believe that
the insects themselves are not attacked by a fungus. Wallace states that
this wonderful imitation is most complete, and that in Sumatra he has
often seen a butterfly enter a bush and then disappear as if by magic. He
states that once he was so fortunate as to see the exact spot on which
the insect settled, but even then lost sight of it for some time, and was
able to discover it close to his eyes only after persistent and careful
search.
The curious and interesting leaf-insects of Java are veined and colored
in such a way that, with the leaf-like expansions from various parts
of the body, not one person in a dozen can detect them when they are
resting upon their food-plants right under one’s eyes. Other insects
resemble pieces of stick (Plate III), with all the minute details of
branches and knots. An eminent naturalist has stated that after being a
practical entomologist for thirty years, he was deceived by one of these
stick insects and took out his pruning knife to cut from a plum-tree
what seemed to him to be a projecting spur. This spur proved to be the
caterpillar of a geometer-moth, about two inches in length. He placed
a portion of the plum-tree on a table, and showed it to several members
of his family, designating a space of several inches in which the
caterpillar was to be found, but none of them could detect the insect
until it was pointed out to them. These protective resemblances of living
creatures to inanimate objects are beautiful illustrations of adaptation
to environment through Natural Selection.
Beautiful illustrations of protective resemblance to particular objects
are furnished by leaf-hoppers (insects) in Central America. They resemble
the thorny and prickly growths of the plants on which they presumably
live. Some of them also resemble gall growths on the plants (Plate IV).
[Illustration: PLATE IV.—Central American Leaf Hoppers resembling the
prickly and thorny growths of plants on which they presumably live.
Certain of them also represent gall growths on the plants. Protective
Resemblance. [Figures collected by Dr. L. O. Howard, from various plates
published in the Biologia Centrali-Americana.]]
=Alluring Coloration.= Besides those insects which secure protection from
enemies by their resemblance to the inert objects among which they live,
there are others whose adaptive resemblance, and therefore concealment,
is for the purpose of securing their food,—for alluring their prey. A
most interesting case of alluring coloration is that furnished by a
wingless insect of India, the mantis. Its color and form are such as to
closely resemble such a fantastic flower as the pink orchis. The insect
rests motionless among the bright green foliage, being very conspicuous
on account of its pink color, and looking so much like a flower that it
allures and captures the butterflies which settle upon it.
There is a species of spider (_Thomisus citreus_) of a creamy-white color
whose abdomen completely resembles in color and contour the unopened buds
of the flowers among which it rests. It has been seen to capture flies
that were attracted to the flowers. There is another species of spider
that looks exactly like the excreta of birds, and through this alluring
resemblance captures certain butterflies. A naturalist has related how,
in pursuing a butterfly through a jungle in Java, he was stopped by a
dense growth of bushes. Here he observed a leaf with a bird’s dropping
upon it, and sitting on this dropping was a beautiful butterfly.
Surprised at such a usually dainty and pretty butterfly seeking such
inappropriate food, he carefully approached to study the actions of the
insect. The insect permitted him to get so close that he seized it by
the wings, and to his astonishment a part of the body remained behind as
if the bird’s dropping was very adhesive. He touched the dropping to see
if it really was sticky, and found that his eyes had been deceived and
that what he took for the excreta of a bird was a most artfully colored
spider, lying on its back with its feet crossed and depressed closely to
the body. The spider had been firmly holding the butterfly.
[Illustration: PLATE III.—Caterpillar, B, of a Geometer Moth
(_Prochœrodes transverrata_) on the stem of a plant (_Ailanthus_), A.
Illustrating Protective Resemblance.]
A very pretty illustration of alluring coloration is furnished by the
common spider (_Misumena vatia_), which is often found spread out upon
the yellow heart of an ox-eyed daisy and in like position upon Coreopsis
(Plate V). It so closely resembles the flower upon which it lurks that
the ordinary observer might well fail to notice its presence. The
coloration facilitates the taking of prey and protects the creature from
the assaults of enemies (Rev. H. C. McCook). It will be observed from the
illustration that the flower is mostly yellow with some red; the same
is true of the spider. This case is a very instructive illustration of
alluring coloration by which the insect has become a living bait for
entrapping butterflies and other insects. Nature has brought about this
complete color adaptation, from the variations of myriads of spiders.
For those ancestral spiders that varied most in the direction of color
adaptation to their environment were less often shunned by shy insects
which could serve as prey. The spiders with useful color variations
would thus most likely secure an abundance of food, and, thus living,
transmit to many of their offspring their useful variations; while those
with inharmonious, and therefore harmful, variations would not be so
able to deceive their food insects, and would thrive poorly or starve
altogether. This is another illustration of the survival of the fittest,
the selection by nature of those best adapted to or in harmony with the
environment; in short, it is Natural Selection.
Wallace, from whom many of these illustrations are taken, says that to
many persons it will seem impossible that such beautiful and detailed
adaptations and resemblances—and these are only samples of thousands
that occur in all parts of the world—can have been brought about by the
preservation of fortuitous useful variations. Yet this will not seem so
surprising, continues Wallace, if we keep in mind the facts of the rapid
multiplication of animals, the severe struggle for existence, and the
constant variability of these and all other organisms; and, further,
that we must remember that these delicate adjustments are the result of
a process, Natural Selection, which has been going on for millions of
years, and that we now see the small percentage of successes among the
myriads of failures. “From the very first appearance of insects, for
instance, and their various kinds of enemies, the need for protection
arose and was usually most easily met by modifications of color. Hence
we may be sure that the earliest leaf-eating insects acquired a green
color as one of the necessities of their existence; and, as the species
became modified and specialized, those feeding on particular species
of plants would rapidly acquire the peculiar tints and markings best
adapted to conceal them upon those plants. Then, every little variation
that, once in a hundred years perhaps, led to the preservation of some
insect which was thereby rather better concealed than its fellows, would
form the starting-point of a further development, leading ultimately to
that perfection of imitation in details which now astonishes us.” So it
is with the beautiful color adaptation of birds, mammals, lizards, and
other animals. There is a lizard (_Phrynocephalus mystaceus_) inhabiting
certain sandy districts in Asia, whose body is protectively colored and
some of whose mouth-parts have alluring coloration and form. The general
surface harmonizes with the sand in which it is found, while the skin
at each angle of its mouth is of red color and so folded as to closely
resemble a little red flower which grows in the sand. The lizard, being
thus in harmony with its surroundings, resembling the sand and the
flowers, is hidden from its enemies, the reptile-eating creatures. But at
the same time insects, being attracted by what they take to be flowers,
approach the lizards and are thus captured, being allured to their
destruction.
[Illustration: PLATE V.—A spider (_Misumena vatia_) lurking for prey on
the center of a flower (_Coreopsis_). Illustrating especially alluring
Coloration (for attracting prey), but also Protective Resemblance
(against enemies). Reproduced from “American Spiders.” By courtesy of
Rev. Henry C. McCook.]
=Warning Coloration.= Many animals possess color patterns that render
them very _conspicuous_ in their environment. It is a very interesting
fact that most of these creatures are the possessors of some deadly
weapons, as poison-fangs or stings, or that they are very disagreeable
and unpalatable food for other animals. Warning colors are most abundant
and best developed among insects. A family of butterflies (Heliconidæ),
in tropical South America, possesses very pronounced and conspicuous
color patterns, so that they are easily seen in their native haunts. Many
of them have deep blue-black with vivid red, white, and yellow spots and
bands, totally unlike those butterflies in the same locality that are
protectively colored and palatable. Their bodies have juices that exhale
a powerful odor. If one kill them by pinching the body, a liquid exudes
that stains the fingers yellow and leaves an odor on them that can be
removed only by repeated washing of the hands. There is a great deal
of evidence to show that this odor is very offensive to insect-loving
animals. Protectively-colored butterflies fly with great rapidity and are
very wary and seek concealment; while the butterflies with conspicuous
colors fly slowly, and do not conceal themselves, as if conscious that
they have no enemies.
Many caterpillars have gay and conspicuous colors and do not conceal
themselves. Bates noticed one in South America four inches long, striped
across the body with yellow and black bands, and with bright red head,
tail, and legs. It could be seen by any one who passed by, even at a
distance of many meters. All of these conspicuous and brightly-colored
caterpillars are unpalatable, and are refused as food by insect-eating
creatures.
Grasshoppers and locusts generally possess green protective tints and
are very palatable, but in tropical regions there are many species most
gaudily decorated with blue, red, and black colors. They are inedible and
are invariably rejected as food by lizards and birds.
A spider whose bite is exceedingly poisonous is found in Queensland. Its
bite will kill a dog, and produces serious illness in man, with agonizing
pain. It is black with a bright red patch on the middle of its body. This
warning coloration is so conspicuous that even the spider-hunting wasp
avoids it.
In all parts of the world frogs are usually protectively colored with
browns or greens; and little tree-frogs are either curiously mottled to
imitate dead leaves or bark, or they are green like the leaves they rest
upon. These protectively colored frogs are always eagerly sought after
by snakes and other enemies. But there are some frogs that are very
conspicuously colored and that hop about with impunity, being avoided by
the snakes and birds of prey. Such is a little frog in Nicaragua which
with its “scarlet vest and stockings of blue” is very conspicuous in its
native haunts.[9] Such, also, is a small toad found in South America,
which is colored a bright vermilion and an intense black, which crawls
about in the sunshine over the sands of arid places. Both of these
animals are altogether avoided by the frog-eating creatures, because they
have disagreeable properties that make them inedible.
In tropical America the very poisonous snake Elaps (Plate VI) is found
abundantly. Its style of coloration is very conspicuous and one that
does not occur in any other group of snakes, consisting alternately of
rings of red, black, and yellow, or red and black of varying width and
arranged in different patterns. Snake-eating birds and mammals have
learned, through hereditary experiences, to avoid these snakes with gay
livery because they are poisonous and therefore dangerous. In Plate VII
the conspicuously-colored black and yellow salamander is an animal with
warning coloration. It is inedible and avoided by carnivorous birds.
These warning colorations have been evolved through the Natural Selection
of fortuitous useful color variations in the ancestors during the
Geologic Ages.
[Illustration: PLATE VI.—Illustrating Warning Coloration (_Elaps_) and
especially Mimicry (_Erythrolamprus_). Elaps is a very poisonous reptile
and Erythrolamprus is harmless. Reproduced (and modified) from Romanes’s
“Darwin and after Darwin.” By courtesy of the Open Court Publishing
Company.]
=Mimicry.= Protective resemblance of a harmless animal to another
of a different species that is harmful is known as mimicry. Mimicry
is bound up with and altogether dependent upon warning coloration.
Some beetles are protected by having integuments, etc., of very great
hardness. Several genera of weevils are in this way saved from attack
by insect-loving birds. These weevils are often closely imitated in
appearance by softer and more eatable species of different genera from
the weevils. Wasps and bees are often mimicked by insects of other orders.
Insectivorous birds are very active in hunting out the edible beetles
(Longicornia), and everywhere in tropical regions these beetles so
closely resemble other insects which are avoided by the birds that the
longicorns are very frequently avoided and thus protected.
In tropical America many butterflies (Heliconidæ) are found that possess
warning coloration. They possess an offensive taste and odor which
almost entirely exempt them from the attacks of insect-eating animals.
The insectivorous birds have learned, by transmitted experiences
(heredity) to avoid the Heliconidæ. It is an interesting fact that in
the same locality with these distasteful butterflies are other species
that are very palatable to insectivorous creatures; but they so closely
resemble the non-edible species that the birds pass them by, not
recognizing their character.
There are some cases of mimicry among birds. There is a genus of large
honey-suckers known as friar birds found in the Malay Archipelago. They
are noisy and powerful birds which go in small flocks. They have sharp
beaks which are long and curved, and also powerful grasping claws. They
are perfectly able to defend themselves, often driving away such birds
of prey as hawks and crows when they approach them too closely. In the
same environment are weak and timid birds known as orioles, which trust
chiefly to their retiring habits and concealment for protection. The
orioles, although an entirely distinct species from the friar birds, very
closely resemble the latter. In each of the great islands of the Malayan
Archipelago there is a distinct species of friar birds, and always in the
same locality is a species of oriole that exactly mimics it. The separate
species often look so thoroughly alike that competent naturalists, prior
to a very close examination, have considered them as belonging to the
same species.[10]
The most remarkable cases of mimicry are those in which poisonous snakes
are mimicked by harmless ones. There is an egg-eating snake in South
Africa that possesses neither teeth nor fangs and is not poisonous. It
very closely resembles the poisonous Berg adder. When alarmed it still
more closely resembles the adder by the habit of flattening its head and
darting forward as if to strike an enemy, hissing at the same time.
[Illustration: PLATE VII.—An Amphibian (_Salamandra maculosa_)
illustrating Warning Coloration.]
In tropical America, in the localities where the poisonous genus Elaps
is found so abundantly, are several genera of harmless snakes of other
families, some species of which so closely resemble or mimic the
poisonous species that they are distinguished from them with difficulty.
The peculiar color patterns of the poisonous snakes serve as warning
colors to snake-eating mammals and birds. The mimicking snakes by flying
these danger-flags are protected.
In Plate VI Elaps is the venomous snake, and illustrates warning
coloration; Erythrolamprus is the edible, non-venomous reptile that has
acquired, through Natural Selection, a protective resemblance to Elaps,
and illustrates mimicry. At the first glance these two snakes look
very much alike; but a closer inspection will show that the detailed
color patterns differ in the two cases. Elaps, though a very poisonous
snake, has the reputation of not being venomous. This error has probably
originated from the fact that it has a gentle disposition and mild
temper, and also from the fact that no doubt it is frequently mistaken
for the mimicking non-venomous species.
Both warning coloration and mimicry are interesting illustrations of
adaptation to environment through Natural Selection; for the myriads
of ancestral forms were continually giving slight variations in color
patterns, some of which were useful to the creatures and others harmful.
Those that harmonized mostly with the environment gave their possessors
an advantage in the fierce struggle for life,—the struggle for food and
place and safety; and procreating their kind age after age, led to the
perfection of mimicry as we behold it to-day.
=Coloration as Recognition Marks.= In gregarious animals, whether
herbivora or carnivora, and whether mammals or birds, a ready recognition
of their own kind at a distance, in the dim twilight, or during rapid
motion, is of the greatest use, and probably often leads to the
preservation of life. Gregarious animals will not usually permit a
stranger in their midst. So long as these animals keep together they are
generally safe from enemies; but a single animal straggling off by itself
may become an easy prey to enemies. In such cases it is of the highest
importance to an animal that it should have every facility for quickly
discovering its companions at any distance within the field of vision.
Also to the young and inexperienced of each herd some means of easy
recognition is of vital importance. Recognition marks also enable the
sexes to identify their kind readily. The necessity for easy recognition
probably is at the basis of the bilateral symmetry in the coloration of
animals.[11] In the struggle for existence those gregarious creatures
that have the best color patterns for recognition marks are the most
likely to get the scattered members of a herd together with the greatest
rapidity, and thus to save them from their enemies. The surviving
members, transmitting their useful variations in recognition marks
to their offspring, are thus able in the course of ages to bring into
existence pronounced color patterns. Thus Natural Selection can account
for much of the coloration in animals known as recognition marks.
One or two illustrations of recognition marks in creatures will be
sufficient. A rabbit when alarmed and fleeing to its burrow displays a
conspicuous, upturned white tail. The rest of the body is protectively
colored. This conspicuous white surface of the upturned tail is a signal
flag of danger. The rabbit mostly feeds during moonlight nights, or
soon after sunset. The white upturned tails of alarmed rabbits serve as
signals and guides to the feeble and young, and also to those at a more
remote distance. Thus a number of rabbits, each following the one or two
in front, are all able in the quickest manner to reach a place of safety.
The spring-bok has a white patch on the face and one on the sides. It
also has a curiously well-marked white stripe above the tail. When
the animal is at rest this last-mentioned white stripe is very nearly
concealed by a fold of skin, but when it is in motion it comes into
full view, like the upturned tail of the rabbit, and serves as a guide
to friends. There are some animals inhabiting the Arctic regions
that are not white,—they are not protectively colored. Such is the
musk-sheep. This animal, though living in Arctic regions, is yet brown
and conspicuous. Its safety depends upon its association in small herds.
It is gregarious. Therefore it is of much more importance to this animal
that it should be able to quickly recognize its companions at a distance
than that it should be protectively colored and so concealed from its
enemies. So long as they keep together in herds they are abundantly able
to protect themselves. This is an exception to the rule of local color
adaptation that proves the rule.
=Sexual Selection.= Among most backboned animals it is the rule that both
sexes should be alike in color. This is especially true among the fishes,
reptiles, and mammals. But in birds the diversity of sexual coloring is
very frequent. It is among this class, therefore, that Sexual Selection
can best be studied. One of the most fundamental characteristics of
birds is the greater conspicuousness of coloration in the males. In the
tropical regions especially are found the most striking examples of
divergence in sexual color patterns. In humming birds, the pheasants,
the peacocks, chatterers, tanagers, and birds of paradise, the females
are exceptionally dull-colored and plain, while the male birds are
gorgeously colored and conspicuously attractive. The male birds of
paradise, for instance, are not only brilliantly colored, but also have
remarkable gorgets, plumes, and crests; whereas the female paradise birds
are without these decorations and as plain as our thrushes in their
ornamental plumage (Plate VIII).
The splendor of plumage which characterizes the male pheasants is
entirely wanting in the females. The intense crimsons and pure whites,
the gorgeous purples and blues of the male chatterers contrast strikingly
with the dull browns or olive greens of the females. The sober hues of
the females have been accentuated by Natural Selection.[12] When the
females were brooding on the eggs in their nests, those of them that had
varied in the direction of conspicuousness would most readily be detected
by their enemies, the hawks, and would be exterminated; but those that
had quiet and dull ornamentation would most frequently escape discovery,
and would pass successfully through the brooding season; thus living and
transmitting their color patterns to posterity,—Natural Selection ever
eliminating the conspicuous and preserving the sober-hued,—in the course
of time the dull ornamentation of the females would become more and more
pronounced.
[Illustration: PLATE VIII.—Male and Female Paradise Birds (_Paradisea
minor_). Illustrating the effects of Sexual Selection. The upper figure
(male bird) is much more beautiful than the lower (female bird).]
But another factor has been at work in accentuating the marked
differences in the ornamentation of the sexes. This factor is Sexual
Selection.[13] In the brooding season there is an intense rivalry
among the males for the possession of the females. Among them the art
of courting has become, indeed, one of the fine arts. The male birds,
like the males of almost all animals, have stronger passions than
the females, and with rarest exceptions are much more eager than the
females. In courtship they display their adornments and accomplishments
most zealously before the females; they strut around them in most eager
courtship and pompous vanity, displaying the utmost rivalry. The males
charm the females in various ways, such as dancing, or performing
fantastic antics either in the air or on the ground; and then again by
most melodious song. After man, the female birds appear to be the most
æsthetic of all animals; therefore, those male birds that are the most
pleasing songsters, or the most attractive in their dances and fantastic
performances, are the ones selected by the females for mating. In this
selection the female birds have paid the minutest attention to fleeting
fashions in strut and dance, in form and color,—the progeny of those
males that have been selected by the females for mating, transmitting
the inherited peculiarities of the parents, have tended more and more to
stamp as fixtures these fleeting fashions, and in this way the males have
become endowed with all sorts of decorations and accomplishments.[14]
Thus have been produced in them the many forms of topknots, wattles,
combs, plumes, and feathers elongated and springing gracefully from many
portions of the body; also the naked skin of the head and the beak,
frequently colored gorgeously. The feathers, through this means, are
often most beautifully tinted in charming patterns. As bearing upon this
theory of Sexual Selection, it can be observed that birds pay the closest
attention to the songs of each other. A bullfinch, for instance, had
been taught to pipe a German waltz, and in doing so was a most excellent
performer. He was placed in a room where there were kept some eighteen
canaries and linnets, and immediately commenced producing his melodies.
The birds all ranged themselves on the sides of their cages nearest
the performer, listening to his singing with the greatest interest.
Undoubtedly this singing is most often a matter of courtship; the female
finch selects that one out of a hundred males whose notes charm her the
most; the female canary always chooses the best singer. The soft cooing
of pigeons and of turtledoves is a matter of courtship. In the breeding
season there is the most intense rivalry between the males in singing; a
bird will sometimes sing until he drops down almost dead.
That female birds exercise choice in mating was believed by Audubon.
He describes how a woodpecker hen was followed by six gay suitors who
continued to perform strange antics until a marked preference was shown
for one of them. A study of the Australian bower-birds illustrates both
the courting antics of the males and the exercise of choice by the
females. These birds build bowers which are sometimes quite large. That
of one species is raised on a thick platform of sticks and is nearly four
feet in length and eighteen inches in height. These bowers are built
on the ground, and are for the sole purpose of courtship, since the
nests are formed in the trees. They are highly decorated with leaves,
berries, feathers, shells, and kindred objects. Both sexes assist in the
erection of the bower, although the male bird is the principal worker.
The bower-constructing instinct is so strong that it is practiced even
in confinement. A naturalist in describing the habits of some satin
bower-birds kept in an aviary says that at times the male will chase the
female all over the aviary, then go to the bower, pick up a gay feather
or a large leaf, utter a curious kind of note, set all his feathers
erect, run round the bower, and become so excited that his eyes appear
to start from his head. He continues opening first one wing and then the
other, uttering a low, whistling note, and, like the domestic cock, seems
to be picking up something from the ground, until at last the female goes
gently toward him and the wooing is completed.
Instances of love dances may be taken from all classes of the animal
kingdom. “Mr. Peckham has described a very interesting love-dance by a
certain species of spider (_Saitis pulex_). He placed a male in a box
with a female. As soon as the former saw the latter, about twelve inches
away, he became excited and at once moved towards her; when some four
inches from her, he stood still, and then began the most remarkable
performances that an amorous male could offer to an admiring female.
She eyed him eagerly, changing her position from time to time so that
he might always be in view. He, raising his whole body on one side by
straightening out the legs, and lowering it on the other by folding the
first two pairs of legs up and under, leaned so far over as to be in
danger of losing his balance, which he maintained only by sidling rapidly
towards the lowered side. The palpus, too, on this side, was turned back
to correspond to the direction of the legs nearest it. He moved in a
semicircle of about two inches, and then instantly reversed the position
of the legs, and circled in the opposite direction, gradually approaching
nearer and nearer the female. Now she dashes towards him, while he,
raising his first pair of legs, extends them upward and forward as if to
hold her off, but withal slowly retreats. Again and again he circles from
side to side, she gazing towards him in a softer mood, evidently admiring
the grace of his antics. This is repeated until are counted one hundred
and eleven circles made by the ardent little male. Now he approaches
nearer and nearer, and when almost within reach whirls madly around and
around her, she joining and whirling with him in a giddy maze. Again he
falls back and resumes his semicircular motions, with his body tilted
over; she, all excitement, lowers her head and raises her body, so that
it is almost vertical; both draw nearer; she moves slowly under him, he
crawling over her head, and the mating is accomplished.”[15]
In addition to that form of Sexual Selection where the female chooses a
mate from among a number of competing males, and which may be designated
_preferential mating_, there is another form of selection in which the
males fight with one another for the mastery and the possession of the
females. Among the higher mammals it is a very general fact that the
males fight together for the possession of the females. This leads,
especially in polygamous animals, to the better armed or stronger
males becoming the parents of the next generation, which inherits the
peculiarities of the parents. Thus the offensive weapons and the vigor
of the males are continually increased, resulting in the antlers of the
stag, the tusks of the boar, the fighting instinct and spurs of the
gamecock, and the horns and strength of the bull. Even mammals that are
not specially armed fight to the death for the possession of the females,
such as beavers, moles, squirrels, and hares. Almost all male birds are
especially pugnacious during the breeding season. Battles have been
observed in such different groups as ducks, finches, woodpeckers, humming
birds, and waders. Among fishes deadly battles occur between the males
of sticklebacks. Also the males of salmons engage in deadly contests;
among reptiles fighting occurs among the male tortoises, crocodiles, and
lizards. Spiders and many butterflies often fight for the females. Thus
Sexual Selection through the _law of battle_ occurs widely throughout
the animal kingdom. This form of Natural Selection greatly increases the
vigor and fighting power of male animals; for, in every case, the weaker
males are either driven away, killed or wounded, and the field is left to
the most vigorous for procreating their kind.
The male stickleback is a little fish that builds its nest among the
weeds, weaving the material together by a secretion from its kidneys.
It is a very passionate little animal, and is exceedingly pugnacious in
relation to its male rivals. The battles of the males are often very
desperate. The combatants fasten tight to each other for a time, tumbling
over and over again, until they appear to be completely exhausted. The
males of the rough-tailed stickleback, when fighting, swim round and
round one another, endeavoring to pierce each other with their raised
lateral spines. In fighting they are perfect little furies, and their
bite is very severe. Their lateral spines are used with such fatal effect
during a battle that a male has been observed to rip open his opponent,
so that the latter sank to the bottom in a dying condition. The females
are very peaceful. When they come out of their hiding-place and view the
nest that the male has made he is mad with delight.
The male salmon is as pugnacious as the little stickleback. Two males
have been known to virtually battle with each other all day long. In
breeding ponds the males can be seen constantly fighting and tearing one
another on the spawning beds, and so many are injured in consequence
that they may be seen swimming near the banks in a state of exhaustion,
many of them apparently dying.
Among birds the law of battle holds as well as the law of preferential
mating. During the breeding season they are exceedingly pugnacious. The
humming birds, the smallest of any, are among the most quarrelsome. Two
males rarely meet without a fierce fight on the wing.
Darwin gives the following illustration of the invincible courage and
fighting instinct of the gamecock. One of these birds had both of its
legs broken in a cockpit, and its owner made a wager that if his legs
could be supported with splints and bandages so that he could stand
upright he would keep on fighting. This was accomplished, and the bird
fought on with dauntless courage until he received a death stroke.
It is probable that even with the most pugnacious species of birds
the pairing does not depend alone on the courage and strength of the
male, for these males are usually decorated with various ornaments.
Furthermore, these decorations during the breeding season often become
more brilliant, and are eagerly displayed before the females. Darwin
states that twenty or more males of the _Tetrao cupido_ (species of
grouse) will assemble at a particular spot, keeping up a tremendous
chattering and strutting about. At the first response from a female the
males take to fighting furiously, and the weaker are vanquished. Both the
victors and the vanquished pay court to the female, so that the latter
must make a choice or the battle is renewed. Here we have the combined
action of selection through battle and by preferential mating.
Among mammals the males win the females much more through fighting than
through the display of charms. In the breeding season the most timid
male animals, which are not even supplied with any special weapons for
fighting, engage in the most desperate conflicts. Two male hares have
been seen to fight until one was killed. Those male mammals which are
provided with special weapons for fighting enter into the fiercest and
most deadly conflicts. The wild male elephant, during the period of love,
is one of the fiercest fighters in the world. Lions engage in terrible
battles, and a young lion dare not approach an old one.
Male seals fight most desperately during the breeding season, using
both their claws and teeth. The conquerors appropriate the females and
transmit their qualities to their offspring.
These are all cases of the survival of the fittest; of the survival of
the males best adapted to the exigencies of their surroundings—in short,
additional illustrations of Natural Selection.
=Insect Selection.= Flowers usually consist of several parts, such as
the stem, the calyx composed of green sepals, the corolla formed of
attractively colored petals, the stamens, the pistil, and finally of
certain nectar-forming organs situated at the base of the last-named. The
upper portions of the stamens are known as antheridia. The free extremity
of each pistil is called a stigma, the intermediate stalk is the style,
and the base is the ovary. The flowers constitute the sexual parts of the
plant. The stamens are the male elements and the pistils are the female
structures. In some species of plants the flowers possess the stamens
but not the pistils,—they are male flowers; in others the flowers have
pistils but not stamens,—they are female flowers. In still other plants
the flowers possess both the male and the female structures, and are
therefore bisexual or hermaphroditic flowers. The ovary of a pistil has a
number of cells in it called ovules (female germ cells); the antheridia
on the stamens have cells in them called pollen (male germ cells). When a
pollen cell is carried by any means to the stigma of a pistil, it sends
down through the latter a tubular prolongation by which the nucleus and
protoplasm of the pollen cell unite with the nucleus and protoplasm of
the ovule; so that we now have a fertilized egg,—the germ of a new plant.
This fertilized ovule by repeated cell multiplication can grow into an
adult plant.
There are two agencies by which the pollen is carried to the pistil;
first, by the wind, and, secondly, by insects (or, occasionally, by
humming birds). Flowers that are fertilized through the agency of insects
are the most beautiful in existence, displaying all the varied hues and
gorgeous patterns that are found in the organic world. On the other hand,
the flowers which are fertilized through the agency of the wind are
incomparably less beautiful than those of insect production, and they do
not secrete sweet juices or nectar. The earliest flowers in geologic time
consisted only of those essential portions, the stamens and pistils, and
had no colored whorl of petals within another colored whorl of sepals.
The poorly developed nectaries secreted only small quantities of honey.
The food-seeking insects visited these primitive flowers for the pollen
and nectar, even as they do now. The nectaries in the plants were so
situated that the insects could not get at the honey without rubbing off
some of the pollen and carrying it to other plants which it visited for
honey. In this way the nectar-seeking insects would often carry pollen
from the flower of one plant to the stigma of another plant’s flower,
and thus secure cross-fertilization. Undoubtedly often the stigma of a
plant’s flower was fertilized by pollen from the same flower; but it is
known that seeds produced by cross-fertilization are much more vigorous
and hardy than those produced by self-fertilization in the same flower;
therefore, those plants that varied most in the direction best adapted
for securing cross-fertilization would have decided advantages in the
struggle for existence. Their fertilized eggs (seeds) would be most
vigorous, and would most readily secure nutriment and withstand adverse
circumstances. The variations of the primitive flowers most calculated
to secure cross-fertilization would be those that made the flower more
conspicuous by the appearance of bright colorations and delightful odors,
such as would attract the attention and visits of nectar-seeking insects.
These variations being transmitted, by heredity, to the descendants and
accentuated as the ages passed would ultimately lead to all the wonderful
adaptations of flower and insect structure to one another that are found
in nature.
[Illustration: PLATE IX.—Two Primrose flowers (_Primula elatior_). A,
with a long style; B, with a short one. Vertical sections have been made
through the flowers.]
Natural Selection has acted in a double manner here, preserving those
flowers that had the most delightful odors or the most conspicuous and
enticing colorations and patterns; and at the same time selecting those
insects that varied most in the direction of keenness of scent, acuteness
of vision, and color perceptions. It will be instructive to briefly
describe the fertilization of the primrose and an orchis by bees. The
_primrose_ (Plate IX) has two sorts of flowers that are never found on
the same plant; one has stamens high up the corolla tube and a short
style with its stigma below the anthers; the other has the stamens far
down the tube and a long style with the stigma above the anthers. In
both cases the nectar is at the very bottom of the corolla tube. When a
bee visits a short-styled flower, it extends its proboscis to the bottom
of the tube and in withdrawing it brings away some pollen cells at its
base. If it should next visit another short-styled primrose, it would not
fertilize it because the pollen on the base of the proboscis could not
reach the stigma; it would only gather still more pollen. But when it
visits a long-styled primrose, it will necessarily deposit some pollen
cells on the stigma, inasmuch as that is at the commencement of the
corolla tube. If the bee should first visit a long-styled form of flower,
it will carry off pollen on the tip of its proboscis, and when it visits
a short-styled primrose will deposit the pollen on the stigma.
In the _orchis_ the stigma is placed below the anther. The stigma is
in the front part of the flower and at the base of the lip, the latter
being prolonged into a long tube full of nectar. The stigmatic surface
is composed of very viscid matter. A bee when seeking the nectar pushes
its head against the anther, and in so doing detaches the two sticky
glands to which the club-shaped masses of pollen cells are attached
(Plate X). It carries these away on the front part of its head. So long
as the masses of pollen cells remain erect on the bee’s head, they cannot
reach the stigma of any other orchis that it visits. As the sticky glands
on the head contract, the pollen masses incline forward and become
horizontal, so that they touch the next sticky stigma that is visited.
The greater stickiness of the stigma detaches the pollen masses from the
bee’s head, so that the flower is fertilized. It takes so long for the
pollen masses on the bee’s head to assume the horizontal position that
the insect has visited all the flowers on one orchis and then visits
another plant. By this time the pollen masses are in the proper position
(horizontal) to fertilize the flowers of another plant. In this way
cross-fertilization is secured and the vigor of the plant maintained.
It may now perhaps be appreciated how intimate are the relations between
the form and habits of insects and the structure and coloration of
flowers. Our standards of beauty have largely been created for us through
Insect and Sexual Selection. If insects had not been developed on the
earth, plants would not be ornamented by beautiful flowers, but would
show only such poor and inconspicuous ones as are found on grasses, on
oak, ash, and fir trees. Grant Allen has beautifully written that, “while
man has tilled only a few level plains, a few great river-valleys, a few
peninsular mountain slopes, leaving the vast mass of earth untouched by
his hand, the insect has spread itself over every land in a thousand
shapes, and has made the whole flowering creation subservient to his
daily wants. His buttercup, his dandelion, and his meadowsweet grow
thick in every English field. His thyme clothes the hillside; his heather
purples the bleak grey moorlands. High up among the Alpine heights his
gentian spreads its lakes of blue; amid the snows of the Himalayas his
rhododendrons gleam with crimson light. Even the wayside pond yields
him the white crowfoot and the arrowhead, while the broad expanses of
Brazilian streams are beautified by his gorgeous water lilies. The insect
has turned the whole surface of the earth into a boundless flower-garden,
which supplies him from year to year with pollen or honey, and itself in
turn gains perpetuation by the baits that it offers to his allurement.”
[Illustration: PLATE X.—Illustrating the fertilization of an orchis by an
insect. A, represents a section of the flower and shows a bee standing on
the flower’s lip with its head touching the sticky portion of the pollen
masses; C, shows the pollen masses stuck to the bee’s head and erect; B,
shows the pollen masses horizontal, the proper position to leave them
detached from the bee by the sticky stigma.]
Natural Selection is the great agency that accounts not only for the
color patterns and forms of living creatures, but also for the great
majority, if not all, of the useful characteristics of organic creatures,
including their internal organization.
One more illustration of the power of Natural Selection may be given,
outside of the subject of color patterns. It may be interesting to
readers who are not familiar with the elements of zoölogy to know that
whales are not fishes but mammals. They belong to the same class that
man does. The embryology of a whale reveals that it is descended from
ancestors that were _land_ mammals, and that these mammals had a scanty
covering of hair, teeth of different shape, broad tails like beavers,
short fore and hind legs, and well developed sense organs. The olfactory
organ was especially well developed. It is probable that these ancestors
of the whale lived in marshy districts and were omnivorous, eating both
plant and animal food. They sought their food in both shallow water and
in swamps. As the conditions of life became more and more unpropitious
on land, they were slowly modified through the ages under the action of
Natural Selection into creatures somewhat like dolphins. At first they
lived in fresh water, but finally they found their way into the sea and
became the rulers of the ocean, from which the giant sea reptiles of
earlier epochs had vanished. Hence are explained the adaptive changes
of structure: the fore-limbs were modified into flippers enclosed in a
fin-like sac, but retaining the bones corresponding to like structures
in other _mammals_, as in the arm of man, the wing of the bat, and the
fore-leg of the horse. Traces of the hind legs may be detected in a few
species; the tail, which acted as a powerful swimming organ, became
divided into two lobes; the head became fish-like in shape; the seven
bones of the neck, common to most mammals, grew together; the skin became
hairless; and the teeth, which appear in the young of the true whale, but
are never cut, gave place to hanging fringes of whalebone, in the meshes
of which the animal entangles the minute organisms it feeds upon.[16]
ISOLATION OF VARIETIES IN NATURE.
The following analysis of Isolation will be useful:
{ Preferential Mating.
{ Sexual. {
Isolation { { Cross-Sterility.
{ Geographical.
It has been stated on an earlier page that the commingling of diverse
hereditary units accomplished through the fertilization of an ovum by a
spermatozooid is the source of many variations in the offspring. In this
fertilized ovum the complexity of chemical substance, and, therefore,
the complexity of inheritance, gives instability to the embryo, and
thus produces variations in the offspring. In this embryo there is
a struggle among the hereditary units,—a struggle among the various
qualities inherited from both sides,—and a survival of the fittest, a
veritable intra-cellular Natural Selection. It is a well-established
law in biology that the union of germ cells of very closely related
individuals, that is, of consanguineous individuals (in and in breeding)
leads to less vigorous and variable offspring, and the parents are
less prolific; while the commingling of diverse heritages by the union
of germ cells from individuals belonging to strong but different
varieties leads to vigorous and quite variable offspring. The union of
such individuals is also most prolific. On the other hand the union of
individuals belonging to very diverse varieties becomes less and less
prolific until the cross-sterility of species is reached, although there
are many exceptions to this rule of cross-sterility. The individuals
of a species living in a state of nature are constantly varying. With
every generation trivial variations take place in all directions and
of all kinds. But these variations are all funded in the common stock,
for the varieties freely mingle among one another and cross-breeding
is constant not only between them, but with the parent stock. Of the
variations that are constantly taking place some are advantageous to
the creatures, some are disadvantageous, while many are neutral, being
neither useful nor harmful. Natural Selection is ever alert, selecting
the advantageous variations and eliminating the disadvantageous ones.
The neutral variations are not touched by Natural Selection; among these
intercrossing of varieties probably affords Nature an opportunity to
make almost endless combinations, some of which might be useful to the
animals, and others harmful, and in either case would come under the
influence of Natural Selection. The commingling of diverse heritages
due to the union of more or less pronounced varieties of the species in
nature not only leads to a funding of varietal characteristics, but also
increases the instability of the offspring, augmenting their plasticity,
so that more numerous and diverse variations take place. According to
Romanes the reproductive organs are among the most variable in the body.
Of the numerous variations taking place in the individuals of a species
under nature, some, therefore, affect the reproductive organs in such a
way that certain of these individuals are cross-fertile with one another,
but _cross-sterile with other varieties and with the parent stock_. This
interesting and very important kind of variation is known to occur in
some individuals of the human species. It is well known that a man and
woman have been cross-sterile with one another, being unable to have
children; yet when separating and mating with others they have both been
cross-fertile, families being reared by both of them.
Variations are commencing species; isolated variations diverge more and
more into distinct species. This fact, then, of the occurrence in nature
of variations in some of the individuals of a species by which they are
cross-fertile with one another, but cross-sterile with other varieties
and with the parental stock, shows that Nature has a most effectual means
by which varieties may be isolated from one another,—just as effective
means as man, the breeder of varieties of domestic animals, possesses
in isolating these domestic varieties by physical barriers, as fences,
etc. Cross-sterility, therefore, in nature, is a most effective sexual
barrier. The special form of it under consideration is what Romanes has
called Physiological Selection. The varieties that are isolated by this
sexual barrier have got to run the risks of in and in breeding, which
Darwin has shown occur in domesticated animals, but which Wallace thinks
are much less in a state of nature.
Another very important mode of Nature for isolating varieties is that
which arises out of the instinctive preferences of animals. There is a
tendency for like to breed with like where varieties are formed. The pale
and dark colored herds of fallow deer in the Forest of Dean have never
been known to interbreed. In the Falkland Islands all the cattle are
known to have descended from the same stock. Here there are differently
colored herds of cattle, and those cattle of the same color will
interbreed with each other, but not with individuals outside their own
color-caste, as Morgan expresses it. When two flocks of heath sheep and
merino sheep are mixed together, they do not interbreed. This isolation
of varieties by instinctive preferences for those individuals with
like color patterns may be spoken of as _preferential mating_ through
_recognition marks_. A very obvious mode of isolation in nature is by
geographical barriers (including migration). In treating of environment
we learned that during the geologic ages of the globe, the physical
geography and climate have repeatedly changed. A very cold temperature, a
mountain chain, a body of water, a stretch of desert land, may completely
prevent interbreeding, on either side of the barrier, between the
individuals of a species.
It should now be understood that Nature, like man, produces divergences
from the parental stock, and isolates them by various effective
agencies. Natural Selection ever carefully watches over these processes,
eliminating the unfit variations and selecting the useful ones.
Variation, Heredity, Environment, Isolation, and Natural Selection,
having been acting and reacting through the ages, have produced, from a
common parental stock, all the innumerable divergent and adaptive forms
of living creatures that can be traced through the geologic strata to
those on the surface of the earth to-day; have produced, therefore, _man_
also as the inflorescence of the topmost branch of the tree of life.
SECTION VI.
EVOLUTION OF MAN.
EVOLUTION OF MAN.
The detailed study of the development (ontogenetic and phylogenetic)
of man is so vastly intricate and extensive a subject that it will be
impossible in a work of this character to do more than refer to it in
brief outline. One of the best and simplest methods to approach the study
of such a subject is to acquire some idea of the _development of the
frog_.
The fertilized frog’s egg, which is the starting point in the life of
a new frog, is deposited in water and hatched by the warmth of spring.
After fertilization the egg or cell divides into two cells, these two
into four, the four into eight, the eight into sixteen, and so on till a
large number of small cells, associated together, is formed (Fig. 10).
These cell phases represent different stages in the life of the growing
frog. It is thus seen that the frog, in its earliest stages, consists of
nothing but many small cells. These cells, through the mysterious powers
of heredity, are going to differentiate as development proceeds into the
various organs (groups of various kinds of cells) that form the tadpole
and finally the frog. For further stages in the development of the egg
(oösperm) towards the tadpole, works upon Comparative Embryology should
be consulted.
At length the tadpole is hatched from the egg, and then soon swims about
in the water (Fig. 17).
The creature that comes from the egg looks nothing whatever like a frog.
It has no limbs whatever, and consists mainly of a bulky head and tail.
This is the tadpole stage in the development of the frog. It can exist
only in water, breathing air therefrom by means of gills. Like the fish,
it has a two-chambered heart. At this stage it has no lungs, and the
gills consist of an external (Fig. 17, a) and an internal pair. The mouth
is small, with only horny toothless jaws, with no tongue. The creature
is herbivorous, living on decaying vegetable matter. The vertebræ of the
spinal column are bi-concave, as in fishes. The tadpole is essentially a
fish, and would be so classed if it did not develop further. An evolving
fish does not go beyond this stage. But the developing frog does go
beyond this stage to a higher one. As its evolution proceeds through the
multiplication and differentiation of the cells that form its body, limbs
begin to bud out, first posteriorly (Fig. 17, b) and then anteriorly
(Fig. 17, c). The lungs now begin to develop, and the external gills
dwindle more and more until they soon disappear, the internal ones
persisting for a while longer. The tongue, at this stage, also makes its
appearance. The creature now can breathe both air and water. This is
the permanent condition of many adult amphibians belonging to a lower
order than the mature frog, such as the _siren_, _menobranchus_, etc.
The siren in developing also passes through the fish stage, but does not
get beyond the siren stage. But the evolving frog does go beyond this
stage, for with the growth of the legs the tail dwindles slowly by its
gradual absorption (Fig. 17, d). The internal gills now disappear through
absorption, and the lungs develop more thoroughly. Great changes take
place in the blood-vascular system, the fish-like, two-chambered heart
evolving into the three-chambered, amphibian heart. In spite of its
dwindling, the tail is still a very conspicuous organ. In this phase of
its development the frog can breathe only air, and must frequently come
to the surface of the water for that purpose, and soon leaves the water
altogether. Now this stage of the creature’s development corresponds to
the permanent adult condition of another order of amphibians, which is
higher than that to which the siren belongs but lower than the order of
the adult frog. This intermediate order has such creatures in it as the
_triton_. The triton in developing passes through the fish and siren
stages, but does not get higher than the triton stage. But the evolving
frog goes even higher than this triton-like condition. Its tail is more
and more absorbed until it finally disappears, and then the young but
perfect frog appears (Fig. 17, e). During this period the teeth develop
and the creature becomes carnivorous, feeding on insects. It is thus seen
that the developing frog passes by small gradations from one class (the
fish class) to an altogether different and higher class (the amphibian
class). When it has evolved to this higher class, it then passes from
the lower order (“siren” order) to a higher one (“triton” order), and
then to the highest order (“frog” order). The bi-concave vertebræ of the
fish-like tadpole have now developed into vertebræ with the cup-and-ball
joints of the higher amphibian. It is the same with all the complex
organs of the adult frog; they evolve from the much simpler structures of
the tadpole.
[Illustration: FIG. 17.—Tadpoles and Frog; a, tadpole with branching
external gills; b, gills absorbed and hind legs have appeared; c, fore
legs have appeared; d, tail shrunk and legs enlarged; e, perfect, young
frog,—tail entirely disappeared. The figures represent some stages in the
life history of the frog.]
This study of the frog’s evolution from the fertilized egg is profoundly
instructive. It reveals to us, through direct observation, that a
creature varies in its form and structure at succeeding intervals of
time. These variations diverge more and more, so that specific, generic,
and even ordinal and class distinctions are revealed as the development
proceeds. Owen, the distinguished comparative anatomist, in speaking of
the transmutation of one species into another in the course of geologic
history, says, though with a hostile purpose in view, that in the
metamorphoses of the amphibians we seem to have such process carried on
before our eyes to its extremest extent. Not merely is one specific
form changed to another of the same genus; not merely is one generic
modification of an order substituted for another, the transmutation is
not even limited by passing from one order (Urodela) to another (Anura);
it affects a transition from class to class. The fish becomes the frog
(amphibian); the aquatic animal changes to the terrestrial one; the
water-breather becomes the air-breather; an insect diet is substituted
for a vegetable one. And these changes, moreover, proceed gradually,
continuously, and without any interruption of active life. Such is the
language of Owen in reference to these remarkable transmutations of the
developing frog.
The development of the frog is a brief recapitulation, an epitome,
through heredity, of the main transmutations of its ancestral forms
in geologic time. It is not true that the embryonic phases in the
development of a higher form always resemble the adult stages of lower
forms. This may or may not be the case; but what always does occur is
that the embryonic phases of a higher form resemble the corresponding
phases of the lower forms. So far as the frog’s development is
concerned, it is very instructive to know that the order of succession
of its embryonic forms undoubtedly parallels the order of succession
of corresponding forms in past geologic ages. Fishes appeared in the
Upper Silurian rocks with amphibian characteristics. In the succeeding
Carboniferous Ages the fishes still continued under new forms; but also
the lowest forms of amphibians, the most fish-like forms, now appeared.
They were somewhat like the sirens, they were perennibranchs. In the next
succeeding rocks, the Permian and Triassic, higher, triton-like forms
appeared. They were caducibranchs. Finally, in the Tertiary rocks, the
highest forms of amphibians are found, such as the frogs.
In order to understand the relation of Ontogeny to Phylogeny, it must be
carefully borne in mind that the simple and lowly organized creatures on
the globe at the first appearance of life were performing the two great
functions that all living creatures perform, viz.: those of nutrition
and reproduction. These functions imply that organisms were reacting to
environment, and, therefore, undergoing modifications and adaptations;
and at the same time the organisms were giving origin to offspring—they
were reproducing their kind through heredity. As these simple organisms
lived through the ages and became more and more complex by modifications
and adaptations to an ever-changing environment, they still evolved
their kind in reproduction. Every new adaptation gained by the parent
was transmitted by heredity, in the course of time, to the offspring;
every form and structure modified in the parent was modified by heredity
in the offspring; and every structure lost by the parent was finally
lost in the offspring. Just in proportion as the parents, through the
ages, became modified, often becoming more complex by the addition of
adaptation to adaptation, retaining some structures of their ancestors
by heredity (through use) and losing others, eventually, through disuse;
so the offspring of these modifying parents became correspondingly
modified, and acquired by heredity the modified structures and habits
of the parents, while losing other structures in time that the parents
had lost. Just as complex _organisms_ of later ages have been evolved
from the simpler organisms of earlier ages by the addition of adaptation
to adaptation, in an orderly sequence (Phylogeny); so, therefore, the
complex _offspring_, while growing, unfold these inherited adaptations
in the order of their acquisition. This last process is called Ontogeny
or Embryology. Ontogeny is undoubtedly an illustration of the results
of Natural Selection’s activity; for, during the phylogeny of the frog
throughout the incalculable ages of the past its ancestors undoubtedly
assumed innumerable forms and structures which were adaptations to
the times and surroundings. But with the advancing time and changing
environment, some of the old forms and structures continued useful and
were retained, while others became useless and were eliminated by Natural
Selection. In addition to the old useful structures that were retained
changing environment often modified some of the retained structures and
added still other adaptations to these. And so on, throughout the ages,
in building up a frog, through geologic embryos, geologic “infants,”
geologic “children,” and finally geologic adult frogs, Natural Selection
has retained during ontogeny many useful structures in the order of
their first appearance, and eliminated innumerable others that became
useless. The ontogeny of the frog, which has been built up by its
phylogeny, reveals the useful structures that have been retained, and in
the order of their appearance; often showing structures that have been
lost in the parent, but are not yet quite lost in the embryo, while it
fails to show innumerable useless structures that have been lost in the
past. This is the reason why we say that the ontogeny of a frog is a
brief outline recapitulation of the main points in the phylogeny of the
frog, with even some main points occasionally omitted altogether. The
geologic ancestors of the frog were the scaffoldings by which it climbed
from simple creatures up to its present complex organization; just as
the embryological phases at present are the scaffoldings by which a
simple, unicellular, fertilized ovum climbs up through heredity to the
huge complexity of the multicellular adult frog. What is true of the
development of the frog, ontogenetically and phylogenetically, is also
true of all living creatures, and is therefore true of =man=.
Man, in his individual development, commences life as a small,
microscopic cell—the fertilized ovum—which is only one-fifth of a
millimeter in size. His first stage resembles an encysted protozoan
animal. As cell-multiplication proceeds he soon gets into the morula
stage, which resembles a colony of undifferentiated protozoans. He soon
evolves into a stage which may be compared to a colony of protozoans some
of the members of which have undergone differentiation. Then comes the
gastrula stage, which is distinctly suggestive of a low metazoan, and
in which the developing germ assumes fundamental anatomical qualities
such as characterize lowly animals like polyps. Then, by gradual
transmutations, the vertebrate characteristics appear; but it could not
be said at this stage of development, if one did not know, whether one
is observing a fish, an amphibian, a reptile, or a mammal. Finally, the
developing man passes through his fish and reptile phases and reaches the
mammal stage. But as yet it cannot be said to which order the animal
belongs. The evolution of the individual continuing, he finally assumes
those anatomical characteristics that stamp him as belonging to the order
of man.
The theory of evolution, then, teaches that this development of man in
the course of a few short months, like the development of the frog, is a
very condensed and abbreviated epitome of the evolution of mankind from
primitive protozoans during the incalculable ages of the past.
Drummond has prettily written that “the developing human embryo is
like a subtle phantasmagoria, a living theater in which a weird
transformation scene is being enacted and in which countless strange and
uncouth characters take part. Some of these characters are well known
to science, some are strangers. As the embryo unfolds, one by one these
animal-actors come upon the stage, file past in phantom-like procession,
throw off their drapery, and dissolve away into something else. Yet,
as they vanish, each leaves behind a vital portion of itself, some
original and characteristic memorial, something itself has made or won,
that perhaps it alone could make or win,—a bone, a muscle, a ganglion,
or a tooth,—to be the inheritance of the race. And it is only after
nearly all have played their part and dedicated their gift that a human
form, mysteriously compounded of all that has gone before, begins to be
discerned as the resultant.”
As has been stated in the introductory part of this book, if all the
animals that have ever lived on the globe should be represented by a tree
those existing on the earth to-day would be indicated by the topmost
twigs and leaves, while the extinct forms would be represented by the
trunk and main branches. Just as the leaves, twigs, branches, and trunk
of the tree have a common origin, viz., the seed that developed into the
tree, so all the different species of animals of the present and the past
are the trunk, branches, twigs, and leaves of the “tree of life,” and
have had a common origin from a primitive protozoan cell (see Diagram of
Development, Fig. 18). Therefore all creatures, living and past, have a
more or less blood relationship.
The Diagram of Development will indicate in a very general way the
possible track taken by a man as he evolved,—grew higher and higher as
the central, straight trunk of the expanding tree of life,—during the
geologic ages; and finally appeared as the inflorescence of the topmost
branch of this central trunk. It is seen from this scheme that the
tree of life commenced in a primitive cell. Without entering into any
discussion of the various theories of evolution and epigenesis, we may
say that the primitive protozoan contained potentially all the animal
forms (each being a cell or group of cells) that have existed on the
globe, just as the fertilized egg contains potentially all the tissues
and organs (groups of cells) of the adult man.
As the tree of animal life unfolded and expanded—like a germinating
seed—from the primitive protozoan, certain of the descendants evolved
along the straight and central branch, through the primitive colonial
protozoans, on through primitive vermes, and still on through primitive
fishes (elasmobranchii), amphibians, reptiles, and on through primitive
ornithodelphia (monotremes), and didelphia (marsupials) to a primitive
order of monodelphia, viz.: primitive primates. The evolution of man
continued through primitive anthropoidea to primitive anthropopithecus.
At this point we meet with the common ancestors of the _higher anthropoid
apes_ (chimpanzee and gorilla) and _man_.
[Illustration: FIG. 18.—Diagram of Development: Portion of the “Tree of
Life,” showing approximately the relative places of the great groups of
animals. The _Central Trunk_ and _Primary Branches_ represent _Primitive_
(geologic) forms; the _Terminal Twigs_ represent _Modern_ forms.]
At each stage of the evolution some of the descendants of the animals of
this stage diverged obliquely, modifying the characters they possessed at
this stage in a direction that varied more and more from those characters
that led on to man. So that all along the central trunk of the tree of
animal life collateral branches were given off. The collateral branches
given off at each upward stage of evolution represent animals higher in
the scale than those that departed from the central trunk lower down.
To illustrate what has occurred at each stage in the evolution of man,
pause for a moment to consider that phase of progress represented by the
primitive reptilia. If we study the anatomy of the specialized reptiles,
birds, and monotremes of the present, we will find that they all have
many characters in common. These characters are _reptilian_. Each class
has its own distinctive specialized peculiarities in addition to its
common reptilian characters. The study of the fossils of the rocks shows
that in the Jurassic and Cretaceous ages animals existed that were
undoubtedly reptiles, but had also very distinct bird characters; also
reptiles existed that had distinct monotreme characters. These reptiles
came from those of earlier times that were still more generalized. As the
ages passed, some of the generalized reptiles (primitive reptiles) lost
more and more the reptilian features and gradually assumed more and more
distinct bird characters, until finally the highly specialized modern
birds (“glorified reptiles”) were evolved as a branch from primitive
reptiles. The specialized reptiles of modern times likewise came from
the primitive reptiles. In like manner those primitive reptiles that had
mammalian (monotreme) characters, by getting into a suitable environment,
gradually lost more and more their reptilian characters and assumed
with increasing accentuation the characters of primitive monotremes—the
lowest of the mammalian class. But observe particularly that the earliest
introduced monotremes were not the specialized monotremes on the globe
to-day, but generalized, primitive monotremes. These gave origin to the
specialized modern monotremes, and also to the generalized primitive
marsupials. The evolution of man continued through the primitive
marsupials to primitive anthropoidea.
Here we meet with the common, generalized ancestors of man and the
monkeys. These creatures contained, potentially at least, _anthropoid_
as well as _pithecoid_ characters. From them were derived the primitive
New World monkeys (primitive Platyrrhines) and the primitive Old
World monkeys (primitive Catarrhines). Some of the descendants of the
primitive Old World monkeys, migrating into an environment which favored
particularly the pithecoid characters, eventually developed into the
tailed monkeys of the Old World (Cercopithecidæ). Others of their
descendants, migrating into a different environment, found conditions
that favored the anthropoid characters especially, and by greater
and greater use of these, with the diminished use of the pithecoid
ones, the characters of the anthropoid apes (primitive Simiidæ and
primitive Simiinæ) became clearer and clearer until, in time, primitive
anthropopithecus appeared,—a tailless anthropoid ape of the Old World.
More than likely this anthropoid ape bore a close resemblance, as Dr.
Theodore Gill long since taught, to the modern chimpanzee. If there were
any differences they could scarcely have been of even a generic value.
This primitive chimpanzee was undoubtedly a quadrupedal, quadrumanous
creature leading an arboreal life. His descendants specialized along
two distinct but closely related lines. Those that continued to live in
trees specialized along the oblique path that led finally to the gorilla
on the one hand and the chimpanzee on the other. Those descendants that
abandoned the trees and lived on the ground used the feet more and more
for purposes of locomotion and less for grasping; while they employed
with increasing frequency the hands for grasping exclusively. Associated
with these adaptations were many other correlated adaptations, such as
the upright posture, an enlarging brain, a change in the character of the
face and of the dentition, etc. As man evolved further and further along
the central trunk of the tree of life, he discarded, through disuse, many
of the characters that are peculiar to the anthropoid apes; and assumed
with increasing emphasis, through use, many of the characters that are
distinctive of man. He passed through the phases of pithecoid man and
pre-palæolithic (primitive) man, until eventually, in palæolithic man,
the visage of humanity is clear and unmistakable.
It is extremely interesting to attempt to form some rough picture of
primitive man. It may help us to do so if we recall what Darwin has said
about the Fuegians, who are among the lowest of savages. He has written
that they are men whose very signs and expressions are less intelligible
to us than those of the domesticated animals—men who do not possess the
instinct of those animals, nor yet appear to boast of human reason, or at
least of arts consequent on that reason.
The Fuegians are much nearer to the ape than to a Shakespeare or Sir
Isaac Newton. In the words of Clodd, primitive man was doubtless much
lower than the lowest Fuegians. “He was a powerful, cunning biped, with
keen sense organs (always sharper, in virtue of constant exercise, in
the savage than in the civilized man, who supplements them by science),
strong instincts, uncontrolled and fitful emotions, small faculty of
wonder, and nascent reasoning power; unable to forecast to-morrow or to
comprehend yesterday, living from hand to mouth on the wild products of
nature, clothed in skin or bark, or daubed with clay, and finding shelter
in trees and caves; ignorant of the simplest arts, save to chip a stone
missile, and perhaps to produce fire; strong in his need of life and
vague sense of right to it and to what he could get, but slowly impelled
by common perils and passions to form ties, loose and haphazard at the
outset, with his kind, the power of combination with them depending on
sounds, signs, and gestures.”
Through the theory of evolution it can readily be understood why the
anatomical characters of the anthropoid apes and of man are so very
closely alike. They have a common origin, and are blood relations—the
one group of animals having specialized from common ancestors in one
direction (obliquely), and the other group having specialized in another
(straight) direction. (Vid. Diagram of Development.)
Man, in his individual development from a fertilized ovum, comes from a
source infinitely lower than the ape. Why, therefore, should he feel such
reluctance to believe that he has passed, during geologic ages, through
the phase of generalized simian ancestors? Is there not much more of hope
in the knowledge that he has risen higher and higher through the æons of
the past than in the belief that he was created an innocent and noble
character and then fell to utter wretchedness through great temptation?
The motto of evolution is Excelsior. For it shows that the human race,
through all the incalculable ages of the past, has risen to higher and
higher levels,—to nobler and nobler phases of being. His progress in
the almost infinite past suggests the hope that he will mount higher
and higher towards perfection during the limitless future. Not only may
we hope that there will be boundless improvement of the human race, but
boundless evolution of each individual human being as well. Evolution’s
motto for each individual may also be Excelsior. And, therefore,
may we say with some assurance of hope that love, while kissing the
pathetic lips of death, need not entertain in vain the splendid hope of
immortality. For if there be no immortality of personal consciousness,
then the evolution of the cosmos, of man, of the highest mind in man,
have no intelligible meaning for us; they are unfathomable enigmas—idiot
stories without meaning.
Man, in specializing along certain lines since separating from the
ancestral simian stock, has displayed more and more that structure of
his skeleton and of the soft parts molded upon it that is best adapted
to the needs of the mind resident within him. His bones are not merely
the jointed framework of an animal, but a framework adapted to that erect
attitude which so befits his intellectual nature. His feet are not the
climbing and grasping feet of the ape, but organs for giving firmness
to the tread and dignity to the bearing of a creature capable of high
thought. The arms and hands are not for strength alone, for these members
are much stronger in many a brute; but they also give greater expression
and power to the thoughts that come from within. The hands possess
such molding of fingers, thumbs, and palms, such delicacy for tactile
impressions, and such capacity for nice adjustments, that they are not
alone used for feeding the mouth and fighting antagonists; but they also
contribute pre-eminently to the desires of a large mind, and are the
efficient servants of its promptings. As Dana well says, “The face, with
its expressive features, is formed so as to respond not solely to the
emotions of pleasure and pain, but to shades of sentiment and interacting
sympathies the most varied, high as heaven and low as earth,—ay, lower,
in debased human nature; the whole being, body, limbs, and head, with
eyes looking, not towards the earth, but beyond an infinite horizon, is a
majestic expression of the divine feature in man and of the infinitude of
his aspirations.”