Vofl REESE LIBRAR"

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:E APODID^

A specimen of the
Spitzbergen variety of
Apus glacialis(x 10), to
show the general ar-
rangement of the limbs
and their gradual dimi-
nution from front to
back. The white masses
on each side near the
brood pouches are par-
tially extruded eggs,

NATURE SERIES

THE APODID/E

A MORPHOLOGICAL STUDY

BY

HENRY MEYNERS BERNARD

M.A. CANTAB.

WITH SEVENTY-ONE ILLUSTRATIONS

U o n ft o n

MAC MILL AN AND CO.

AND NEW YORK
1892

The Right of Translation and Reproduction is Reserved

BiOLO<

lUl." • "

RICHARD CLAY AND SONS, LIMITED,
LONDON AND BUNGAY.

TO THE MEMORY OF

DR. ALFRED WALTER

TOO SOON LOST TO SCIENCE

AND

TO THE MANY FRIENDS WHO LOVED
HIM AS A MAX

TABLE OF CONTENTS

PAGE

INTRODUCTION xi

PART I

§ I. OBJECT AND LINE OF ARGUMENT i

§ II. THE OUTER BODY FORM . . . • 1 1

§ III. THE APPENDAGES 26

§ IV. THE MUSCULATURE 52

§ V. THE NERVOUS SYSTEM 67

§ VI. THE SENSORY ORGANS 84

§ VII. THE ALIMENTARY CANAI 112

} VIII. THE CIRCULATORY SYSTEM 117

§ IX. THE EXCRETORY AND OTHER GLANDS 124

§ X. REPRODUCTION 140

§ XI. DEVELOPMENT — THE XAUPLIUS 148

TABLE OF CONTENTS

PART II

• PAGE

§ XII. RELATION OF APUS TO THE OTHER CRUSTACEA—
LIMULUS 173

§ XIII. THE TRILOBITES 209

§ XIV. THE EURYPTERID.*: 237

§ XV. THE NEW CLASSIFICATION PROPOSED 252

§ XVI. PERIPATUS AND THE TRACHEATA 279

APPENDICES 291

LITERATURE 313

INTRODUCTION

AMONG the numerous interesting zoological
specimens brought back by Professor Kiikenthal
from East Spitzbergen on his return from the Bremen
Expedition (iSSp),1 was a small species of Apus —
presumably Lepidurus glacialis (Kroyer). Professor
Kiikenthal very kindly handed these specimens to
me for examination. By the kindness of the Rev.
Canon Norman, I also received Greenland specimens
of Lepidurus glacialis (Kroyer), and from Professor
Leche, of Stockholm, specimens of the Lepidurus
glacialis found by Professor Nathorst in West Spitz-
bergen ; Professor Mobius, the director of the Berlin
Natural History Museum, generously sent me speci-

1 Dr. Alfred Walter, to whom this book is dedicated, \vas Professor
Kiikenthal's companion during this expedition. He died shortly after
his return.

xii INTRODUCTION

mens of Apus cancriformis, and Professor Biedermann,
of Jena, very kindly obtained for me specimens of
Apus cancriformis, Lepidurus productus, and Branchi-
pus stagnalis from Prague, especially preserved for
histological purposes. An examination of the Spitz-
bergen specimen led to the conclusion that it
was a small variety of Lepidurus glacialis, which
I propose to call Lepidurus Spitzbergensis ; the
grounds for this determination are given fully in
Appendix I.

In studying the anatomy of the Spitzbergen
specimens, and in comparing it with that of the other
members of the family kindly placed at my disposal
by the gentlemen above named, I was gradually led
to cast my notes into the form in which they are now
published.

It has long been recognised that the Phyllopoda
possess many markedly Annelidan characteristics, and
that they are therefore, of all living Crustacea, nearest
in affinity to the primitive Crustacean. In my study
of the Apodidae I was so much struck by the resem-
blance between the organisation of Apus and that of
a carnivorous Annelid, that I finally decided to work
entirely along this line. I resolved, by a closer study

INTRODUCTION xiii

of each organ and system of organs, to find out as fat-
as possible whether this resemblance was a case of
homology or analogy, and, if the former, to endeavour
to trace the causes which led to the transformation of
the carnivorous Annelid into the Crustacean.

Shrewd conjectures have been made as to the
possible derivation of the Crustacea from Annelids,
but I am not aware that this point has ever before
been worked out in detail, and I should hardly have
ventured to undertake such a task had not my study
of Apus forced it upon me.

My original intention x of preparing a comparative
anatomy of the Apodidae thus gave way before the
more ambitious attempt to use Apus as a key to
solve the hitherto unsolved problems as to the origin
of the Crustacea, and the true affinities between the
various groups.

This resolution, however, was not formed at once.
The book is written in the order in which the subject
was worked out.

The first part, which deduces Apus from a carnivo-
rous Annelid, was all I at first intended to publish.
Having never made a special study of Limulus nor

1 Announced in a letter to Nature, reprinted in Appendix V.

INTRODUCTION

of the Trilobites, I hesitated to discuss their relation
to Apus, and my knowledge of the Crustacea was
not sufficient to justify my attempting to form a
genealogical tree of the whole class. I intended to
content myself with an endeavour to show that in
the Apodidae the process and method of the trans-
formation of carnivorous Annelids into Crustacea
was still visible in almost every organ and system of
organs. The unavoidable conclusion from this would
be, that Apus must be — for some groups at least —
the original form.

Here I thought to leave the matter to be followed
up by zoologists whose knowledge and experience of
the special groups were greater than my own. My
curiosity, however, was too great, and after the first
part of this manuscript was practically in its present
form, I decided to sec, by a study of Limulus and
the Trilobites, whether Apus was to be looked upon
as the original form of the modern Crustacea only, or
whether it could pass for the original of the whole
class, including these archaic forms. This investiga-
tion led to the writing of the second part, which was
thus an afterthought — an afterthought which, how-
ever, gives to the book whatever value it possesses.

INTRODUCTION

The attempted proof in the first part that Apus is
an original Crustacean easily derivable from an
Annelid, however interesting in itself, must have
remained little more than a curious morphological
study. The appeal made in the second part to
palaeozoic Crustacea must, however, be decisive as to
whether our claims for Apus as one of the original
forms can be definitely established.

In commencing Part II., it was no small encourage-
ment to find that most of the earlier zoologists, as if
by instinct, classed Apus with the Xiphosuridae and
the Trilobites. This provisional classification had
not, however, held its own, and it was necessary to
examine the reasons why it had not done so, and to
see if it was not after all justified by the facts. My
investigations led me to the conclusion that if Apus is
deducible from a carnivorous Annelid in the manner
described in the first part, there is no possible escape
from accepting a similar derivation for the Giganto-
straca, as Haeckel has called these ancient forms. I
found that, strange as it may at first seem, the
very differences between Apus and these ancient
Crustaceans yielded almost more striking proofs of
their having had the same origin and of their close

xvi INTRODUCTION

relationship, than did the many resemblances which
have been long recognised as existing between them.
The second part thus proves what the first part only
rendered probable.

In such an investigation as this a writer is always
open to the charge of having interpreted the facts as
he wished to interpret them. I cannot of course deny
that the speculation was of such absorbing interest
that I was not indifferent to the conclusion, and that I
therefore naturally seized upon the facts most favour-
able for the establishment of my argument ; but at
the same time I am not conscious of having ignored
difficulties. If, nevertheless, I have unconsciously
distorted the facts in order to establish my con-
clusions, I comfort myself by the reflection that those
conclusions are of such great zoological importance
that they cannot long pass unchallenged.
, I may perhaps mention the fact that whereas in the
first part I have relied almost entirely upon my own
researches into the anatomy of the Apodidae and of
the carnivorous Annelids, in the second part I have
had to draw many of the facts used in the arguments
from the works of others.

My sincere thanks are due to Professor Ernst

INTRODUCTION

Haeckel, in whose laboratory the researches on which
the following essay is based were carried out, for the
friendly interest he took in them ; and also to Professor
Kiikenthal for his cordial sympathy and encourage-
ment to proceed in a speculation which claims to
solve so intricate a problem as the origin of the

Crustacea.

H. M. 13.

Streatham, 1892.

PART I

THE APODID^E

PART I

SECTION I

OBJECT AND LINE OF ARGUMENT

THE Apodidae have been known and studied for
the last one hundred and fifty years. They have
always attracted considerable attention, not only on
account of their great size in comparison with other
fresh-water Entomostraca, but also on account of their
strange and sudden appearance in pools and ditches
which owe their water entirely to the rainfall. This
also is not all : their morphology has been a perpetual
puzzle to zoologists, and they have been classed by
some with archaic forms such as the Trilobites and
Limulus, while by others they have been considered
as highly specialised recent forms.

This essay claims by a new explanation of the
morphology of the Crustacea, to set this latter point

*

THE APODID^: PART I

at rest, and to show that Apus must not only be
ranked by the side of the Trilobites as one of the
primitive Crustacean forms, but that it is itself a true
link between the living Crustacea and the Annelida.
By careful examination of the organisation of Apus,
and a comparison of it with that of a carnivorous
Annelid, it is possible to show, as will be done in the
following pages, that Apus is perhaps the most perfect
" missing link " which zoology so far possesses, perfect,
not only because its morphology is easily deducible
from that of a carnivorous Annelid, but also because
the mechanical causes of the transformation are
apparent. The Apodidse will in fact be found to
afford us the first complete illustration of the rise of
one large animal class out of another by the simple and
natural adaptation on the part of one single species
of the latter to a new manner of life. Close investiga-
tion shows the Apodidse to be both morphologically
and biologically an almost ideal transition form.

More or less satisfactory transition forms be-
tween most of the great animal classes are now
known, but none has till now been discovered
between the Annelida and the Crustacea. The
object of this book is to satisfy this want, not by
the discovery of a new animal, but by a new ex-
planation of one long known and often described.

The established transition forms between the other
classes of the animal kingdom still leave much to
be desired. Between the Protozoa and Metazoa
the transition forms are either claimed by botanists,
or else, however probable, are somewhat hypothetical.

SECT. I OBJECT AND LINE OF ARGUMENT 3

Between the Coelenterata and the Platodes we have
rival links. When reading the arguments in favour
of the claims of those specialised Ctenophora, the
Cceloplana and Ctenoplana, we feel convinced ; but,
on the other hand, when we study for ourselves
under the microscope such a simple Rhabdoccele as
Microstomum lineare, especially during its changes
of shape when moving about under a cover glass,
our former conviction fades away, and we see in it
a specialised larval form of a Ccelenterate. Between
the Platodes and the Annelids the gap seems small,
but we cannot bridge it over until we decide whether
the segmentation of the Annelids is a kind of axial
strobilation, or the natural mechanical selection of
internal symmetry. Between the Annelids and the
Molluscs we have the claims of Solenogaster to attend
to ; but this animal is unfortunately so rare, that
it will be long before we can hope to have any very
thorough knowledge of its morphology. Between the
Annelids and the Tracheata we have Peripatus ; this
highly interesting animal, has a special claim on our
attention, as the Tracheata form with the Crustacea the
great class known as the Arthropoda. We shall find
that our explanation of the rise of the Crustacea
supplies us also with a very probable clue as to the
origin of the Tracheata. The Echinodermata and
the Tunicata hover almost entirely in the air. And,
lastly, we have the giant trunk of the Vertebrata, the
roots of which are being eagerly sought in different
directions. The claims of Amphioxus and of^the
Ascidian larva are confidently put forward by the

B 2

THE APODID^ PART I

majority of our leading zoologists, but there are diffi-
culties not yet explained which make many restless,
and lead them to search in other directions.

In this state of affairs it will be a clear gain and
encouragement if we can connect the Annelida and
the Crustacea in the way described in these pages, in
which we show how a typical carnivoroifs Annelid
(presumably a Nereid, though probably not so
specialised as any modern member of that family)
can, by a simple and natural adaptation to a new
manner of life, be established as the ground type
of Apus. We mean a great deal by this expression
"ground type," much more than any mere general
resemblance of organisation ; we mean that every
single organ of Apus, where it does not resemble
that of its Annelid ancestor, is capable of being
deduced from some organ in the latter, and, further,
that the causes of the transformation are not far to
seek. These are large claims ; the following pages
will show whether they are justifiable.

Before entering into the morphological and
anatomical details upon which our deduction of the
Apodidae from a carnivorous Annelid is based, it
will make the task of the reader lighter if we here set
out the line of argument.

Many carnivorous Annelids have, as is well known,
a protrusible pharynx, armed with teeth, which is shot
out for the seizing of prey. We assume that the
Annelid from which Apus is derived, adopted a
habit of browsing, which rendered this protrusible

SECT. I OBJECT AND LINE OF ARGUMENT 5

pharynx unnecessary, so that it degenerated. The
Annelids afford us such a wonderful variety of
forms adapted to almost every possible manner
of life, that this assumption presents no difficulty.
Cambrian and Silurian formations have revealed to
the palaeontologist abundant evidence that early
Chaetopods crawled about along the bottom of the
seas of those times. That one of these should become
specialised for feeding in the manner supposed, is
not too much to ask.

The use of the pharynx just described is, as far as
we can see, a clumsy method of obtaining food. The
loss of it, and the adoption of a browsing method of
feeding, might well be a gain. The further develop-
ment of this habit would lead to a bending round of
the head sufficient to enable the animal to use its
anterior parapodia for pushing prey into its mouth.
In time the bend of the head would become fixed, and
the parapodia modified as jaws and maxillae. The
parapodia of at least a certain number of anterior
trunk segments would certainly also serve to rake
food together into the middle line and forward it
towards the mouth. From this very simple and natural
modification of a Chsetopodan Annelid, we believe that
all the Crustacea, living or extinct, can be deduced.
To establish this, is the object of this little book, which
we have called " The Apodidae," since it was during our
study of these Phyllopods that we first caught sight of
the Annelid, so effectually disguised under its Crusta-
cean dress. Although this disguise is so complete as to
have eluded all former research, yet when once under-

THE APODID^: PARTI

stood, it is found to be very superficial. We shall be
able to show, in the following pages, that the Apodidae
agree in almost every detail of their organisation with
such an Annelid, and that any disagreement is chiefly
due to further specialisation in adaptation to the
new manner of life described.

Commencing with the head, we shall show how the
morphology of the typical Crustacean head is easily
explained by the bending round of the five anterior
segments of such an Annelid for the purpose of
browsing.

The trunk of Apus will be shown to be a true link
between the many-segmented Annelids, and the
Crustacea with their small and almost constant
number of segments. The rise of the shield will be
briefly mentioned, a fuller account of it being reserved
till we compare Apus with the Trilobites.

The gradual transformation of the Annelidan
cuticle into the exoskeleton of the Crustacea, to which
many of the changes in the inner organisation of the
latter are to be referred, will be found well illustrated
by the Apodidse.

The Annelidan parapodia (with their dorsal and
ventral branches) will be shown to be capable of
developing every form of Crustacean limb, the reasons
for the suppression of one part and the development
of another being generally fairly evident, Apus again
supplying the clue.

Coming to the inner organisation, we shall take in
turn, the musculature, the nervous system, the sensory
organs, the alimentary canal, the circulatory system,

SECT. I OBJECT AND LINE OF ARGUMENT 7

the excretory and other glands, and, lastly, the repro-
ductive organs. We shall either point out the
resemblances in each case between these organs and
those of our Annelid, or else show how they can be
deduced from Annelidan organs. It will be found
that while some of the modifications of Annelidan into
Crustacean organs are easy to follow, the explanation
of others has to be sought, and may thus appear to be,
in some cases, far-fetched.

And here we must remind our readers that it is
enough for our argument if we can show that such a
deduction is possible. It is not essential to our theory
that we should show exactly Jiow the inner transform-
ations actually took place. Our explanations may
themselves be incorrect, but the validity of our argu-
ment can only be seriously weakened by showing that
a set of organs in Apus could not possibly have been
derived from any organs in the Annelida ; or that the
improbability of such a transformation is so great that
no experienced morphologist would accept it.

We shall conclude the first part of this essay by an
appeal to the Nauplius, to see whether it bears out our
theory that Apus is the original form of the majority
of the modern Crustacea ; or, in other words, whether
Apus can itself claim to be the proto-Nauplius of
zoologists. We shall endeavour to describe the exact
morphology of the Nauplius considered as the Apus
larva or the Apus-stage in the development of the
other Crustacea.

This will conclude Part I., which we hope will have
shown that, so far as such claims can be based purely

THE APODID^: PARTI

upon morphological, anatomical, and biological reason-
ing, the Apodidae deserve to take the place we assign
them as an almost ideal transition form between the
Annelida and the Crustacea. Here, as stated in the
Preface, we thought to leave the matter as an interest-
ing suggestion. Fortunately, however, we have the
means of testing the accuracy of our conclusions..

Admitting, on the one hand, that the confirmatory
evidence as to the truth of our theory given by the
Nauplius need by no means be conclusive, we maintain,
on the other hand, that the answer which we receive
to our appeal to palaeontology and to such archaic
living forms as Limulus must be decisive. Thus we
enter upon the second part of our essay in order to
obtain a final " yea " or " nay " as to whether our theory
is, as a whole, but a morphological tour de force, or a
fairly close guess at the truth.

We commence with Limulus, and show that if Apus
is to be derived from an Annelid with the first five
segments bent round ventrally, Limulus must have
had a similar origin.

In the second section we venture into the dangerous
realm of the Trilobite's. The mystery which surrounds
these primitive Crustacea is so great, that every
announcement of a new discovery bearing upon their
morphology meets with more or less scepticism.
Nevertheless, we believe that we can prove that our
derivation of Apus from a bent Annelid reveals the
Trilobites also in their true light, as so many attempts
of browsing Crustacean-Annelids to adapt themselves
to their surroundings — attempts which, in the long

SECT. I OBJECT AND LINE OF ARGUMENT 9

run, proved unsuccessful, for reasons which we shall
try to point out.

After briefly discussing the Eurypteridae, we shall
give an outline sketch of a new classification of the
Crustacea based upon our theory, showing that
while only one group of modern Crustacea admits of
derivation from the Trilobites, all the rest, except
Lirnulus, can be deduced from the Apodidae. We
shall see reasons- for believing that it was the develop-
ment of the shield, either as bivalve shell, or as a large
fold of the tergum of the fifth segment, which led to
success in the struggle for existence.

We should here say something as to the preservation
of the Apodidae through so many geological ages.
This is explained by the manner of life of the animals.
They usually appear in ditches and pools dependent
on the rainfall. In such waters they naturally come
little into competition with other animals. The dry
seasons are bridged over by the eggs being preserved
in the mud. In this strange but perfectly natural
way, Apus has, from the earliest times, been so com-
pletely isolated that its preservation presents no
difficulty. Its presence in every part of the globe,
with almost always the same manner of life, is a sign
of its great antiquity.

The fact that no true fossil Apodidae are found,
among the rich yield of Crustacean remains of the
Silurian strata, admits of simple explanation. We say
no true Apodidae, for we shall find that such forms as
Hymenocaris and Ceratiocaris, though perhaps some-
what more specialised, were probably very closely

io THE APODID^ PARTI

related to the Apodidae. Both these points will be
discussed more in detail in Part II.

Finally, in a short concluding section we shall show
that the method of differentiation which turned the
Annelid into the Crustacean throws a flood of light on
the origin of the Tracheata, and on some of the
morphological differences which separate these two
divisions of the Arthropoda.

Several new points in the anatomy' of Apus will be
described and illustrated. Where these do not bear
directly upon the subject, they will be given in full in
an appendix, so as not to interfere with the course of
the argument.

SECTION II
THE OUTER BODY FORM

THE HEAD

THE comparative anatomy of the Crustacea has
long ago established the fact that the Crustacean head
must originally have been composed of five fused
segments of an annulate body. Our derivation of
Apus from a browsing Annelid explains the method
of this fusion, that it did not take place along the
longitudinal axis of the body, but by a doubling of
this number of segments upon themselves. This
term " doubling " or " doublature " has already been
applied to the under sides of the forehead of such
animals as Apus, Limulus, and the Trilobites, but
apparently meaning nothing more than the doubling
of the forehead, which has both a dorsal and a ventral
surface. In reality, however, this " doubling" is the
true description of the Crustacean head as shown in
Figs, i and 2 ; these should be further compared with
Fig. 46, p. 212, which represents a longitudinal section
of a Trilobite, where the doubling is very clear.

12

THE

PART 1

Owing to the bending on itself of the cylindrical
Annelidan body, the original head must have been
anteriorly almost completely hemispherical (Figs, i and
46). This form of the original Crustacean-Annelid head

< 6

3— -J

9-'

FIG. i. — Diagram showing the first six segments of a carnivorous Annelid ; the first
five being bent round ; /, the prostomium with two pairs of eyes and a median
cirrus, i, the ist segment carrying a pair of antennae, its under edge projecting
backwards as the lower lip(/). 2, the 2nd segment with a pair of antennal para-
podia. 3, the 3rd segment with rudimentary dorsal parapodia, the ventral
parapodia developing into mandibles. 4, the 4th segment with a pair of maxilla;,
the dorsal parapodium slightly less degenerated. 5, the 5th and last head
segment, the dorsal parapodium with large aciculum and gland. 6, the 6th
segment (ist free segment) with large dorsal parapodium carrying gill (g) and
sensory cirrus (c).

was, howeverrvery clearly modified. In most Trilobites
traces of it are still visible in the glabella (Fig. 47, p. 2 1 3).
In Apus, the disguise is very complete, the whole head

SECT. II

THE OUTER BODY FORM

being broad and flat. This form is due to a ridge run-
ning round the anterior surface as prolongation of the
lateral edges of the shell fold. This ridge is of consider-
able interest, as it appears in almost every Trilobite.
We shall later find reason to believe that it was a

2

--c

FIG. 2. — Diagram of head and first trunk segment of Apus, for comparison with
Fig. i — the lettering the same. In addition : s, shell gland (the acicular gland
of Fig. i drawn into the shell fold). The distances between the limbs are much
exaggerated in order to show their forms more clearly.

primary differentiation of the new " head," i.e. the new
Crustacean head, composed of five Annelidan segments.
Its origin and modifications will be discussed in
another place. This form of head is no doubt useful
for swimming and perhaps for burrowing in the mud.

I4 THE APODID^: PART I

Excepting in the number of limbs, all external traces
of its having been composed of five segments arc
obliterated. Internally, however, there are abundant
indications of its origin from the bending of the
Annelidan segments.

The mouth lies ventrally and faces posteriorly, a
fact which, taken in connection with the sharp bend
in the oesophagus, to be described later, is enough of
itself to suggest the original doubling of the segments.
This ventral position of the mouth is an important
characteristic of the whole class of the Crustacea,
which has not received the attention it deserves.

Projecting ventrally and posteriorly over the mouth
is a large upper lip, corresponding with the prostomium
of the Annelid ancestor. This upper lip is thus a
primitive feature among the Crustacea ; it occurs in a
more or less pronounced form in most Nauplii, and
persisted as a very highly developed organ in the
Trilobites, but in the modern Crustacea it is
generally more or less rudimentary

The under edge of the Annelidan mouth would also
naturally project backwards as a sharp fold (see Fig. i,/).
Such a fold in Apus, however, would form an obstacle
to the pushing of food forward towards and into the
mouth by means of the maxillae and ventral parapodia
of the anterior trunk segments ; hence we find it
modified into two lateral projections, the middle part
of the fold being merely indicated by a low ridge,
which is not sufficient to form a barrier across the
ventral surface. These two projections have been
handed on to the higher Crustacea as the paragnatha.

SECT, ii THE OUTER BODY FORM 15

In the larva of Euphausia they develop early as two
limb-like projections posterior to the mandibles,
and strongly resemble those of Apus.

In Apus, only the mandibles work between the
labrum on the one hand and the under lip on the
other. In Limulus, however, where the under lip also
consists of two projections, the mouth is so stretched
in the longitudinal direction that the masticatory
ridges vi five pairs of limbs work as jaws between
them and the labrum. That these labial projections
in Apus and Limulus are really homologous with one
another, and with the under lip of our original
Annelid, will be seen to follow as a necessary con-
sequence of our explanation of their morphology.1

The shield is of great size, and stretches back (as a
fold of the fifth segment) over the greater part of the
body. Laterally it covers and protects the limbs.
Posteriorly, it is armed with thorns, and has a keel
along the dorsal middle line due no doubt to the
central thorn which it supports. The carrying of
these thorns, which are now so slightly developed,
may have originally been the chief function of the
shield in its early stages ; Fig. 48, page 215, in con-
nection with which the origin of the shield will be
discussed more in detail, illustrates what we imagine
to have been the first step in the formation of the
dorsal shield. This function has, however, long given
place to that of protective covering of the whole dorsal
surface and (laterally) of the gills. The lateral edges
of the shield are prolonged into the ridge which, run-

1 fy~- PP- 39> 4°> and X94> a^so Fig. 43> P- l&&-

16 THE APODIDyE PART i

ning round the front of the head, makes the latter
broad and flat, obliterates all external marks of seg-
mentation, and effectually disguises its origin out of
five Annelidan segments.

The coils of the shell glands form one of the most
notable marks of the shield (see Frontispiece). Their
origin, position, and structure will be discussed in
the section on the excretory and other glands.

THE BODY PROPER.

On removing the shield we find a long vermiform
annulate body, Fig. 3. In the anterior part of the trunk
region the rings correspond in number with the limbs
or parapodia ; as we approach the posterior region*
however, the limbs are much more numerous than
the rings. We find two, three, four, or as many as
six rudimentary limbs on one ring. The last five
rings have no limbs at all. This whole phenomenon,
which has hitherto puzzled morphologists, may be
explained as follows.

The great length of the original Annelid being
of no use to the Crustacean-Annelid, the hinder
part of the body remains in the latter at an un-
developed or larval stage. The rule in the develop-
ment of Annelid larvae is that the successive
segments form in front of the anal segment, and
differentiate from before backward, those furthest
from the anal segment being the most developed.
In Apus, we find in front of the anal segment

SECT. II

THE OUTER BODY FORM

five segments with no limbs developed, with no
ventral ganglia, and with no organs except the
most necessary, viz., the intestinal tube and the
musculature. Then follows a row of rudimentary
segments, each with a minute pair of limbs and a pair
of ganglia, which increase in size and development
from behind forward. The rudimentary segments
which have become fixed in the adult Apus do

bp

FIG. 3. — Lepidurus Spitzbergensis, from nature. The left half of the shield removed
to show the vermiform body. The first 14 trunk segments carry a pair of limbs
each, the following 10 "rings" carry between them ca. 28 rudimentary limbs,
and therefore correspond to 28 segments. The last 5 trunk segments (excluding
the anal segment) are limbless, bp, brood pouch formed by the nth pair of
trunk limbs.

not correspond with the rings of the body ; only
gradually as they recede from the limbless segments,
and thus are more fully developed, do the segments
correspond with the rings. The fusing of several
rudimentary segments to form one body ring, i.e. a
muscular segment, presents no difficulty. The Myria-
poda afford us several examples of the fusing of true
segments to form body rings.

The study of the development of Apus has shown

18 THE APODID/E PART i

that it grows directly out of the Nauplius by the
gradual differentiation of new segments in front of
the anal segment, with no metamorphosis worth
mentioning. This fact has led to its being compared
with an overgrown Nauplius. According to our view,
indeed, the Nauplius is only the young Apus, or
Apus-stage in other Crustaceans. This steady develop-
ment of Apus from its larva, as an Annelid develops
from the Trochophora, falls in with our explanation
of its morphology. Apus, however, differs from its
Annelid ancestors in that it reaches its adult shape
before its inherited number of segments are fully
developed. This fixation of the hinder part of the
body at a larval stage can be easily accounted for by
the process of natural selection, compactness being a
decided advantage to an active free-swimming animal.
The great number of segments, developed and
rudimentary, in the Apodidse is a matter of
considerable importance in estimating their true
position. In all the other specialised Crustacean
groups the number of segments is constant, i.e.
constant for each group. In deriving such animals
with a small but constant number of segments from
Annelids with a large and varying number of seg-
ments, the ideal transition form would be an animal
with a medium number of segments, which is not
quite constant and is visibly diminishing. Both these
points are specially clear in the Apodidae. We find
that all descriptions of Apus cancriformis give a
varying number of limbs, which can hardly be due
merely to the difficulty of counting them, but more

SECT. II THE OUTER BODY FORM 19

probably to the fact that the number of rudimentary
limbs actually does vary. And even if it should be
proved that the same species always possesses the
same number of segments developed and rudimentary,
the different species of Apus and Lepidurus are
marked by decided differences . in the number of
segments. Whereas the more specialised Crustacea
(the Malacostraca) have either the constant number
of twenty or twenty-one segments, the number in the
Apodidae varies between thirty-five and sixty-five.
In the Entomostraca the number varies, but never
reaches even the lowest number in the Apodidae.
That the number in the Apodidae is visibly decreasing
follows from our explanation of their morphology.
The fact that the posterior segments remain fixed,
in a larval and undeveloped condition, shows that
they are gradually being dispensed with. On this
ground alone, then, the Apodidae deserve to occupy
the place, half way between the Annelids and the
Crustacea, which we claim for them.

Many of the segments, as already seen, are so
rudimentary as to be useless, i.e. as movable segments,
so that three, four, or even six combine to form one
body ring. In the Trilobites we shall find that the
posterior rudimentary segments, which were for the
same reason immovable upon one another, form
together, in many genera at least, a solid tail plate,
the pygidium (cf. Fig. 49, p. 220).

It is a characteristic of the Crustacea that no limbs
develop on the anal segment. In the Apodidae, this
segment is already fully developed ; they are the

C 2

20 THE APODID^: PART i

segments anterior to it which are rudimentary, and
which disappear in the development of the higher
Crustacea.

The anal segment is provided with two long
cercopoda or cirri, projecting posteriorly and slightly
ventrally, and two rudiments, probably of similar
appendages, on the posterior dorsal surface of the
segment. These four together correspond with the
four anal cirri found in some carnivorous Annelids
(cf. pp. 85 and 274). The two cirri are stiffened for the
greater part of their length by a thickened cuticle
covered with setae, and showing slight rings of thinner
skin. The tips of the cirri are quite thin-skinned, and
seem to function as tactile papillae.

The posterior dorsal surface of the anal segment
is sometimes prolonged into a variously shaped caudal
plate or lamella,1 which we shall find to be the
homologue of the caudal spine of the Xiphosuridae.

THE CUTICLE AND EXOSKELETON.

The generally thin and flexible Annelidan cuticle of
the Apodidae shows local thickenings which may
well be recognised as the commencement of the
Crustacean exoskeleton. A closer study of these
reveals to us the principles of the original formation

1 Apodidas having this characteristic have been classed by Leach as a
separate genus, Lepidnrus. But Dr. Alfred Walter, to whose memory
this essay is dedicated, discovered a form in a desert well in Trans-
caspian Russia, A pus Hackdii (Walter), which makes it doubtful whether
this division can be sustained. (Bulletin de la Societe Imperiale des
Naturalistes de Moscou, 1887.)

SECT, ii THE OUTER BODY FORM 21

of such an exoskeleton. Perhaps the best way to make
the subject clear is to discuss and illustrate these
principles.

(1) First and chiefly, the cuticle is thickened for
the protection of exposed parts. We find the cuticle
of the dorsal surface of the head, which, on the bending
round of the anterior segments, was left entirely un-
protected, and of the upper surface of the shield,
considerably thickened ; also that of the exposed
segments, i.e. of those segments which are not covered
by the dorsal shield. Underneath the shield, on the
contrary, the skin of the body is very thin and flexible,
though towards the posterior edge of the shield it
begins to thicken. We find the same principle in
Limulus and the Trilobites, where, under the protec-
tion of the thick shield, the cuticle of the ventral and
lateral parts remained soft and flexible.

We also find certain parts of the body thickened
for protection against other parts. Thus the outer
edges of the under lips are thickened for protection
against the working of the powerful jaws and of the
first maxillae, between which two pairs of limbs they
are placed.

(2) We find local thickeni-ngs to counteract the pulls
of the muscles, and this in two ways, (a.) There are
thickened areas such as the ventral and lateral parts
of the mandibles, to resist the almost rectangular pulls
of the mandibular muscles. In the higher Crustacea
such thickenings of the cuticle go hand in hand
with the concentration and physiological perfection
of the muscle bands, which, instead of being attached

22

THE APODID.E

PART I

to large irregular areas of a soft cuticle, as in the
Annelids and in the trunk segments of Apus, are
attached to definite firm points. (#.) There are thick-
ened strips to resist the longitudinal pulls of muscles,
as along the shafts of the limbs, where, but for such

777

FIG. 4. — Anterior (concave) aspect of a trunk limb of a large specimen of Apus
cancriformis. ^The shading shows the commencement of the thickening of the
cuticle, the white parts being thin and transparent. The musculature at the base
of the limb is slightly indicated. Lettering the same as in Fig. 6, p. 32.

bands, the cuticle of the limb would be drawn into
folds (Fig. 4).

(3) There are thickenings for the formation of claws
and teeth at the tips or edges of limbs, and of thorns

SECT. II

THE OUTER BODY FORM

on exposed angles, edges, or surfaces, as, for instance,
round the posterior segments (see frontispiece).

(4) There are thickenings for the formation of
rudimentary articulations, especially of the limbs upon
the body and of the joints of the limbs on each other.
Figs. 4 and 5 give two views of one and the same leg
of a large specimen of Apus cancriformis. In these

FIG. 5. — Posterior (convex) aspect of the same leg, the skin being nearly all thin and
semi-transparent, but showing rudimentary hinges. A few muscles are indi-
cated. Lettering the same as in Fig. 6. On the endites are seen the denticulate
seta^ referred to on p. 46.

the thickenings forming rudimentary hinges are very
instructive, the rest of the cuticle of the leg, with
the exception of the thickened strips and areas for
the counteraction and attachment of muscles, being
thin and flexible. It would be an interesting mechanical
problem to try to discover why the bent concave side

24 THE APODID.C PART i

of the limb should develop most strongly the
exoskeleton, and the convex side the hinges.

We thus find in Apus the Annelidan cuticle
changing into the exoskeleton of the Crustacea ; the
principles of the change being for the most part
easily deciphered.

The importance of this gradual thickening of the
cuticle for the whole organisation can hardly be over-
estimated. As one of the special characteristics of the
Crustacea, useful at all stages, it is naturally very
early developed, the youngest larva having a cuticle
too thick to allow of gradual regular growth. This
leads to the habit of moulting, which was doubtless
very gradually acquired. The earliest thickenings
probably peeled off separately in flakes, as the areas
which they covered increased in size. Such half-
loosened flakes in all parts of the body would, however,
materially hinder the animal in the struggle for
existence, and natural selection would soon bring
about a shortening of the process, those animals being
most successful who were, during life, least encumbered
by loosening flakes, i.e. who threw them off altogether.

But still more important consequences of the stiff
cuticle are to be traced in the inner organisation.
Some of the greatest differences between the anatomy
of Apus and that of an Annelid can be traced directly
to its development.

The hairs with which the cuticle is covered will be
described in the section on the sensory organs.

We thus find in their outer organisation that the

SECT, ii THE OUTER BODY FORM 25

Apodid^e are but slightly modified Annelids, the
widening of the head being due simply to a fold of
the skin ; and the cuticle being for the most part thin
like that of the Annelids, showing, however, localised
thickenings in which we can recognise the commence-
ment of the Crustacean exoskeleton.

The hinder part of the body — the trunk — has long
been recognised as worm-like, but we here see that
the front or head part is also essentially Annelidan,
especially in its possession of a prostomium or upper
lip. The head of Apus differs from that of our
Annelid only in the development of the shield and
of the ridge-like fold which gives the head its great
breadth.

The Annelid character of the Apodidae, thus visible
in the form of the body (i.e. of both head and trunk),
will be even more clearly seen when we come to
consider the appendages.

SECTION III
THE APPENDAGES

THE appendages of the ApodicLnc have been much
discussed, and many attempts have been made to
homologise them with the limbs of other Crustacea.
From our point of view, we must look to the para-
podia of our bent Annelid for the true understanding
of these limbs. It is, indeed, generally acknowledged
that the Crustacean limb arose from the Annelidan
parapodium. But the way this took place has not
been worked out. The limbs of Apus, however,
supply us with a clue. From these Phyllopodan
limbs we can work both backwards to the Annelidan
parapodium and forwards to the typical Crustacean
biramose limb.

Continuing the detailed comparison of Apus with a
bent Annelid, already begun in the first section, we
have now to show that it is possible to deduce the
limbs of the former from the parapodia of the latter,
and that the modifications which transformed the one

SECT, in THE APPENDAGES 27

into the other are due to adaptation to the browsing
manner of life.

Our deduction of Apus from an adult carnivorous
Annelid, which gradually adopted the habit of bending
round its head, and of using its parapodia for capturing
its prey and pushing it into its mouth, gives us at once
the general direction along which we should expect
modification to take place. First of all we should
expect the parapodia along the whole length of the
body to be bent round towards the ventral middle
line. Further, the dorsal and ventral branches of
these parapodia would be somewhat differently de-
veloped in adaptation to the various needs of the new
manner of life. The ventral parapodia on the three
posterior head segments would be differentiated into
jaws and maxillae, while on the anterior trunk seg-
ments they would serve to rake food into the middle
line and forward it towards the mouth. The uses to
which the dorsal parapodia could be put are not so
apparent. Since, however, the habit of browsing
necessitates a certain amount of locomotion, we may
safely conclude that they would be utilised for this
purpose. The original Annelid in its Annelidan
days no doubt moved in the typical manner of
Annelids by the alternate extension and contraction
of the body. As, however, the body of our Crustacean
Annelid shortened and began to develop an exo-
skeleton, some other method of locomotion would
become necessary. The dorsal parapodia would thus
naturally be brought into requisition. The same
efforts which brought the ventral parapodia round

28 THE APODIDiE PARTI

towards the ventral middle line might be expected to
bring the dorsal parapodia as well, at least far enough
round to allow them to assist in locomotion. There
is, further, no need to limit the functions of the dorsal
parapodia simply to locomotion, — they may at the
same time assist in capturing food. Starting from
the assumption that it was the habit of browsing
'which first led to the transformation of the Annelid
into the Crustacean, the above is, in outline, the way
in which we should expect the Annelidan parapodia
to be gradually developed into Crustacean limbs.

It is not possible in this place to bring forward at
once all the arguments which, we think, show that this
sketch of the rise of the Crustacean limbs out of
Annelidan parapodia is a fairly correct account of
what actually took place. One reason is, however, here
in place while discussing the limbs as a whole. It is
only in such primitive Crustacean forms as the
Apodidae and the Trilobites that we find the ventral
parapodia retained and functioning as jaws along the
greater part of the body, as we assumed for our
original Crustacean-Annelid ; the dorsal parapodia of
the same segments functioning, in the Trilobites,
purely as locomotory organs, in Apus both for
locomotion and for capturing food. In the higher
Crustacea we find a pronounced division of labour,
viz., the perfection of the ventral parapodia round
the mouth for mastication, and of the dorsal para-
podia in the rest of the body, either anteriorly for
seizing food, like the chelate limbs of the Decapoda,
or posteriorly for locomotion, like the ambulatory legs

SECT, in THE APPENDAGES 29

and swimmerets of the same animals, the ventral
parapodia on these limbs disappearing entirely.

We shall further find that this division of labour
in the modern Crustacea was not discovered by
Nature all at once. Many different combinations of
the ventral parapodia as jaws, with dorsal parapodia
as auxiliary appendages, held their own for long
'periods. In the long run, however, the typical
Crustacean formula for mandibles and maxillae has,
except in Limulus and in the Ostracods, which have
different masticatory formulae, alone survived. These
efforts of Nature to select the best arrangement of
ventral and dorsal parapodia for the transformation
of a carnivorous Annelid into an armoured Crustacean
will be found tabulated in Part II. p. 250.

Before describing the limbs of Apus in detail,
we must call attention to several important points,
which tend to support their claim to have originated
from Annelidan parapodia in the way described.

(i) The limbs of Apus are little more than highly
developed integumental folds with only rudiments
of articulations, either between the different joints
of the shaft and its appendages, or between the
shaft and the body (Figs. 4 and 5). This absence
of developed articulations has already been pointed
out by Lankester and others, but its true signifi-
cance does not seem to have been noticed. It
is true that in many small thin-skinned typical
Crustaceans the articulations seem to be slightly
developed, but in these the whole exoskeleton has
been reduced ; this does not affect the significance

30 THE APODID^: PARTI

of their absence in Apus, where we find many
thickenings of the cuticle which we regard as an
exoskeleton — not in the act of disappearing — but in
that of appearing for the first time.

The fact that the limbs arc little more than folds
of the integument, like Annelidan parapodia, is fully
borne out by the examination of their musculature,
which will be described in detail later. (Sect. IV.)

(2) The course of a line traced through the bases
of the limbs of Apus agrees well with that of a
similar line drawn through the parapodia of our
imaginary bent Crustacean-Annelid. Commencing
at the anterior antennae (see Figs. I and 2) at the
side of the prostomium or upper lip, this line passes
in both cases vertically upwards and (for reasons
to be given later, see p. 212) slightly outwards;
passing through the 2nd antennae, it bends round
to run backwards, trending, however, gradually
towards the ventral surface. The close agreement
between the courses of these two lines is, morpho-
logically, a fact of great significance. The more
ventral trend of the line in the posterior end of
Apus was to be expected as a necessary adaptation
to the Crustacean manner of life, i.e., to the use
of the appendages as limbs whose functions were
primitively all directed towards the middle line.
The position of the antennae is especially interest-
ing. In the Annelid, these antennae were originally
metastomial, but have become prostomial by the
bending of the segments on one another. The
assumption of the rise of the Crustacean head from

SECT, in THE APPENDAGES 31

five bent segments thus offers a clear solution of
the difficult morphological problem involved in the
prostomial position of these appendages.

(3) As to the great number of the limbs, remind-
ing us of the row of parapodia on each side of the
Polychaetan Annelids, we need only refer to what
was said in the previous section as to the great
number of the segments (pp. 16-18).

(4) The marked difference between the head- and
the trunk-limbs will be presently discussed in detail,
and the differences shown to be exactly what our
theory demands. In the meantime we find a gradual
change in the trunk limbs as we go from front to back.
Though the Phyllopodan type is preserved through-
out, the anterior limbs (except the first, which is
specialised) are highly developed seizing limbs, the
posterior are simplified as rowing plates. There
can be no doubt that the more rudimentary limbs,
though necessarily repeating the type of those
previously developed in front of them, partake more
of the character of an integumental fold, like an
Annelidan parapodium, than do the anterior limbs
with their developed shafts, claws, &c. (cf. Figs. 4, 5,
and 10).

(5) On placing a typical Phyllopodan limb by the
side of a typical Annelidan parapodium, the homo-
logies of some of the parts are very clear. As,
however, the establishment of the homologies in detail
is not so easy, we shall, in this place, have to content
ourselves with merely stating our conclusions. The
reasons which led to these conclusions, apart from

THE APODID^E

PART I

those already given in the foregoing pages, will be
gradually gathered as we proceed, for we shall find it
necessary to return to the subject again and again in
the course of the following discussion.

A comparison of a limb of Apus with an Anne-
lidan parapodium such as is shown in Fig. 6 A,1 is
sufficient to enable us to homologisc the shaft and its

FIG. 6. — Diagram to compare an Annelid parapodium (A) 1 with a limb of Apus (/?)•
£•, gill; c\, sensory cirrus of the dorsal parapodium ; c-^ ditto of the ventral para-
podium ; v, ventral parapodium ; e, sensory endites. In all the figures the same
letters are used for the homologous parts.

appendages with the dorsal parapodium, the claw
being the true tip.

The gnathobase is the ventral parapodium. The
position of the gill is the same in both, and the
flabellum of Apus is clearly homologous with the

1 Figure A agrees fairly well with that given by Ehlers as the para-
podium of a young Nereis ; we have added the gill, and the sensory
cirrus of the ventral parapodium. See Taf. xxi. 3 in "Die Borsten-
witnner," Leipzig, 1864-68.

SECT, in THE APPENDAGES 33

sensory cirrus of the dorsal parapodium of the
Annelids.

The sensory cirrus of the ventral parapodium (r.7)
entirely disappears in Apus, but is retained in Limulus
(see Fig. 44, p. 192). Both the dorsal and ventral para-
podia in Apus carry highly developed setae, as in the
Polychaeta, and again, in both, the gill is entirely free
from setae, which would hinder the free exchange
of the respiratory medium. It may be noted that
many Polychaeta have appendages on their parapodia
quite as complicated as those on the limbs of Apus
known as the endites (B. e.\ whose origin will be
discussed later.

Passing from the Phyllopodan to the typical Crus-
tacean limb, we assume that the flabellum is the ex-
opodite, the shaft of the limb is the endopodite, and
the ventral parapodium is the masticatory ridge.

I. The first pair of Antenna (Fig. 7 A) — This limb
has retained its original position at the side of the
Annelidan prostomium or upper lip of the Apodidae.
It has already been pointed out that the bending of
the head has changed its position from behind the
mouth to in front of it.

The form of this limb needs no special comment ;
the bend in it is not a true joint, nor is it provided
with any muscles except a few which run into the bulb
on which it stands. Its setae are modified into sensory
hairs, homologous with the olfactory hairs of the
higher Crustacea.

Morphologically, the first antenna must be regarded

D

34

THE APODID^

PART 1

'

as the sensory cirrus of the parapodium of the first
segment, the parapodium itself having disappeared, or
possibly being represented by the bulb on which the
antenna stands ; this latter homology is, however, very
doubtful. Though the parapodium itself has dis-
appeared, the gland of its aciculum is probably still
present, and functions as a salivary gland. Fig. 29
p. 114, shows the position and form of this gland. A
comparison of this figure with Fig. I will make it

?d

FIG. 7. — A. First antenna (L Sfit*&er*tHStt)vnth hooked sensory hairs, homolo-
gous with the olfactory hairs of the higher Crustacea, c, the sensory cirrus ; d,
the bulb on which it stands, perhaps the remains of the dorsal parapodium on
which it stood. B. Second antenna (Z.. Spitzbergcnsis) showing the rudiments
of the endopodite, i.e. of the distal end of the dorsal parapodium which had
been highly developed in the Nauplius, but is degenerated in the adult ; c, the
sensory cirrus.

clear that this gland can be so homologised ;— the
point will, however, be discussed in detail in connec-
tion with the description of the gland itself. From
Fig. i it will be seen that the acicular gland of the
(vanished) parapodium of the first segment could
easily open within the mouth, on the under lip.

SECT, in THE APPENDAGES 35

II. Tlie second pair of Antenna (Fig. 7 B) — The
second antenna is so much reduced in the Apodidaj
that its absence has often been considered characteristic
of the family. In no specimen examined by us has it
been wanting. Its position has already been described
(p. 30) as agreeing exactly with that of the correspond-
ing antennal parapodium of the Annelids. It has, like
the first antenna, been brought in front of the mouth
by the bending of the head. Although it is very much
degenerated, it shows three divisions, with the slight
rudiment of a branch at the end of the second, which
is the only trace of its former relatively greater develop-
ment as a branched swimming limb in the Nauplius.
We deduce the limb from the antennal parapodium of
the second segment of the Annelida (cf. Fig. 7 B with
Figs. I, 2). Its great reduction in the Apodidae is no
doubt due to its being caught, as it were, in the angle
of the bend, and further shut in under the shield. In
Limulus, owing to the greater space under the shield,
it is freer to develop into a chelate foot (see Fig. 43,
p. 1 88). In Branchipus also, in which the shield has
disappeared, it undergoes no such reduction.

The second antenna, like the first, stands on a small
bulb which may perhaps be homologous with that
of the first antenna, but certainly in this case cannot
represent the remains of the parapodium. According
to our homologies for this limb (see Fig. 7 V) the
dorsal parapodium is still present, and forms its
proximal half, ending in the minute rudimentary
branch shown in the figure. In sections of the basal
bulb we found a deep indentation, which led us to

D 2

THE APODID^: PART I

suspect the presence of a rudimentary antennal gland.
No such gland, however, could be found, although the
indentation may mark the spot where one formerly
opened. Whether this indentation, which was very
distinct in some specimens, really represents the
remains of an opening of an antennal gland or not,
we are still able to assume that such a gland,
homologous with the acicular gland of the dorsal
parapodium, once existed, and has reappeared in
the higher Crustacea as the antennal gland. We
have, in the Crustacean head three glands derived
from three setiparous glands, viz. the gland of the
first antenna (?) developed into a salivary gland (in
Apus), the gland of the second antenna into the well-
known antennal gland, and the gland of the second
maxilla into the shell gland. The homologous
setiparous glands of the mandibles and first maxillae
have quite disappeared, like the parapodia to which
they belonged. To these points, however, we shall
again refer.

We have given in the Figure (7 B) our explanation
of the parts of the second antenna. The correctness
of this explanation naturally depends on a right
understanding of the same limb in the Nauplius. This
matter will therefore be further discussed in the section
dealing with the development of Apus. We may here
anticipate our conclusion by saying that, in accordance
with the homologies given on p. 32, and indicated in
the lettering of Fig. 7 B., the biramose limb of the
Nauplius consists of the dorsal parapodium, the distal
portion of which forms the endopodite, the sensory

SECT. Ill

THE APPENDAGES

37

cirrus forming the cxopodite. As the larva grows,
the endopoditc gradually degenerates, leaving the
sensory cirrus to form the distal end of the limb
which is thus a sensory organ (compare also Figs.
34 and 35).

III. 77/6' Mandibles. — These are the first limbs which
admit of undoubted comparison with parapodia. \Yc
find, however, that while the dorsal branch has entirely

B

FIG. 8.— A, diagram of mandible; s, shell-fold. At d the dorsal parapodium has
disappeared, but is indicated by the musculature. 7', ventral parapodium form-
ing the mandible itself, the musculature coming from the sternal plate ($/). £,
diagram of ist maxilla. At d, a larger rudiment of the dorsal parapodium is
retained than in the mandible.

disappeared, the ventral has grown enormously in all
directions to form the powerful masticatory limbs
which are such a striking feature in the Apodidae.
That these mandibles arc homologous with the gnatho-
bases of the trunk limbs, and therefore, according to
our view, with the ventral parapodia of the Annelida,
may be seen at once by comparing their muscles with

38 THE APODID^R PART I

those of the similar parts of other limbs (see Figs. 8
and 14, p. 59). Muscles also are found which are the
remains of the muscles which once ran into the now
completely degenerated dorsal parapodia (see section
on Musculature, p. 52). The redevelopment in
varying degrees of the dorsal parapodium in the
mandibles of some of the higher Crustacea, to
form the mandibular feelers, or palps (or perhaps
feeler- or palp-carriers) is a matter of considerable
interest.

We see in these large fleshy mandibles of Apus an
undoubtedly primitive characteristic. They form a
perfect morphological transition between a limb like
the parapodium of the Annelida, and the hard special-
ised jaw of the Crustacea. The "teeth" are only
hard protuberances of the cuticle. The setae are very
small and grouped in tufts round the teeth. This
limb has no hinge on which it works ; it orobably
moves round its upper dorsal end as axis.

The homologous limb in Limulus, i.e., the third, is
not so specialised as it is in Apus ; its masticatory
process is not more developed than that of the second
antennae, or of the two pairs of maxillae, and of the
first trunk limb. In Apus, only the mandibles work
between the upper and lower lips, but in Limulus all
the masticatory ridges of five pairs of limbs do so.
In Pterygotus (see Fig. 55, p. 239), the third limb
seems to have ceased to function as a jaw, and the
powerful swimming limb, the sixth or (morphologi-
cally) the first trunk limb, has developed strong masti-
catory processes, which seem to function as the chief

SECT, in THE APPENDAGES 39

mandibles.1 It is difficult to see on what principle
this occurs ; the advantages gained by using the
ventral parapodia of the most powerful limbs as jaws,
would seem to be more than counterbalanced by the
disadvantage of combining locomotory with masti-
catory functions ; perhaps we might assume that in
this case the dorsal and ventral parapodia became
separately articulated with the body, so as to secure
all the advantages of the division of labour. From
what we know both of Crustacean and of Annelidan
morphology, there is no difficulty in the assumption
of the separate articulation with the body of two
parts of the same limb or parapodium.

TJie Under Lips. — These, though not limbs, are best
described here, as they have hitherto always been mis-
taken for limbs or parts of limbs. They owe their
origin, as has already been described, to the change of
position of the mouth, the under edge of which must
naturally project backwards (as shown in Fig. i, p. 12).
This under lip, which was originally a straight ridge,
has been cut out in the middle, in order not to form a
barrier which would hinder the pushing of food into
the mouth by the maxillae. We thus find, instead of
the pronounced ridge right across the posterior edge
of the mouth, two limb-like projections, the inner
edges of which gradually slope down towards the
middle line, the remains of the ridge being easy to
follow from side to side in a series of sagittal sections.

1 The different attempts of the primitive Crustacea to find the best
combination of limbs to function as jaws will be found in a table,
p. 250,

40 THE APODID^l PART i

It is clear that this division of the under lip must have
been a very primitive feature in the Crustacean-
Annelid. It was absolutely necessary if the ventral
parapodia (posterior to the under lip) were to be used
for pushing food forwards into the mouth, which
habit led the way, according to our theory, in the
differentiations which transformed the Annelid into
the Crustacean. This early division of the under lip
accounts for its appearance as two ventral projections
in Limulus, where such a form can have no special
significance. The mouth in Limulus is a long median
slit, and instead of only the mandibles and two
maxillae being used as jaws, the ventral parapodia of
the four posterior cephalic and the first trunk limb
perform the masticatory functions.

We find a divided upper lip in some Trilobites (sec
Fig. 49, p. 220), which offers a curious parallel to the
divided under lip of A pus and Limulus, but must
naturally be due to other causes (sec however note
p. 241).

IV. The First Maxilla. — Unlike the mandibles,
the first maxilla has retained a small rudiment of
the dorsal parapodium in the form of a fold. That
this is, in fact, the reduced dorsal branch is clear
from a comparison of the musculature with that of the
other limbs (see Fig. 8 B). In the higher Crustacea,
this limb also may redevelop its dorsal branch as
maxillary feeler. The maxillae work behind the
lower lip. The limb itself requires no special descrip-
tion, its form can be seen from the figure.

In Limulus the dorsal parapodium is well developed

SECT, in THE APPENDAGES 41

in addition to the masticatory ridge or ventral para-
podium. In the Eurypteridae, the ventral masticatory
part seems to have almost disappeared in Pterygotus,
(see Fig. 55, p. 239) but to be well developed in
Eurypterus (Fig. 56, p. 245).

V. The Second Maxilla. — This much-discussed
limb has retained rather a larger rudiment of the
dorsal parapodium than the first maxilla ; in it, in
fact, the dorsal is the more important of the two
branches, the ventral being rendered almost useless
by the powerful first maxilla, between which and
the ventral parapodium of the first trunk limb it
is squeezed (see Fig. 2 and Frontispiece). The
dorsal parapodium is reduced to a stump without
appendages, but is interesting on account of the
aperture of the shell gland at its tip ; we are thus
able, as already mentioned, to homologise the shell '
gland with the setiparous sac of the original An-
nelidan parapodium (see section on Excretion, also
Fig. I, p. 12). The opening of the shell gland on
this limb shows it to be the second maxilla, and not
a maxillipede, as some authors maintain. An ex-
amination of the musculature shows further that this
limb belongs to the fifth segment, a long thin muscle
band descending into it from in front of the opening
of the shield, which is a fold of the dorsal integu-
ment of this segment. This muscle band is doubt-
less homologous with the bands which, in each
segment, run from the dorsal integument into its
limb (see Figs. 14, p. 59, and 15, p. 60).

The five limbs just described are the typical five

42 THE APODID^: PART I

pairs of limbs of the Crustacean head, so marked off
for all time by the bending round of the five anterior
segments of the original Annelid, and the growth of
the shield. The three posterior limbs develop the
ventral branches of the original Annclidan parapodia
as masticatory apparatus round the mouth ; the dorsal
branches are more or less completely degenerated,
reappearing, when the antennae adopt a frontal
position, as palps, or palp-carriers. This arrange-
ment of masticatory ridges may have had some-
thing to do with the preservation of the modern
Crustacea, while the older groups with other, and
probably less advantageous, combinations, such as
those developed by the Trilobites, have died out.

We find here also the origin of the rule that, among
the Crustacea, the dorsal parapodia are less developed
on the mandibles than on the first maxillae, and less
on the first than on the second maxillae. The typical
Crustacean mandibular palp consists of the dorsal
parapodium carrying its sensory cirrus, both apparently
being sensory organs.

The limbs liable to most modification are, naturally,
the first two, i.e. the two pairs of antennae. In Apus
we find the first pair retaining perhaps the original
size, the second pair, however, degenerating. In
Limulus both pairs are well developed as chelate
seizing limbs, the second even redeveloping its ventral
parapodium as a masticatory ridge. In the Euryp-
teridae, other characteristic changes will be noticed,
and where possible will be traced to changes in their
manner of life. We may here perhaps suggest the

SECT, in THE APPENDAGES 43

rule, that the more the forehead is pressed against the
ground, the antennae pointing backwards, the more
likely would the latter be to degenerate, as in Apus,
and in the Trilobites (?), or by losing the sensory
function to become modified as in Limulus. On the
other hand, the more the animal adopted the swimming
manner of life, the more the antennae would tend to
point forwards as sensory organs, and would then
undergo further development as such. This we see in
some Eurypteridae, Branchipus, and Nebalia.

The Apodidoe thus furnish us with a complete
explanation of the way in which the typical Crustacean
head is composed of five fused Annelidan segments
with their parapodia, and why the limbs of the head
differ from those of the trunk throughout the whole
class. The former (leaving out the antennae) develop
chiefly the ventral parapodia for mastication, the
latter the dorsal for locomotion and for other functions
which will be discussed in the following pages. In
Apus we find a transition between the two, the last
head segment having both dorsal and ventral para-
podia about equally developed.

The important modifications, however, which the
cephalic limbs may present, will be referred to again
when discussing Limulus, the Trilobites and the
Eurypteridae, and are tabulated p. 250.

44 THE APODID^: PART

THE LIMBS OF THE TRUNK.

Passing from the second maxillae to the first trunk
limb, we are struck by a sudden contrast, the former
being much reduced, while the latter is highly de-
veloped, indeed the most highly developed limb of
the whole body. This point is of more than ordinary
interest, as we shall find that it throws considerable
light on the homologies of the limbs in the Xipho-
suridse, the Eurypteridas, and the Trilobites, in which
animals, from what we learn from Apus, we are able
to assume that the first large locomotory limbs
must be homologous with the sixth pair, i.e. with the
first trunk limbs. The explanation of the great
differentiation of the first pair of trunk limbs, in all
these primitive Crustacea, is no doubt to be found
in the fact that the sixth segment was really the
first free segment, i.e. the first segment not used up
in any way in the bend which forms the head. Its
parapodia were thus free to develop as limbs for
locomotion or for some other function unconnected
with the mouth (sec Fig. i). The development of the
anterior trunk limbs into maxillipedes in the higher
Crustacea, has long been considered to be a secondary
modification. In the chief point which constitutes
them maxillipedes, i.e. in the retention of the ventral
parapodia as masticatory ridges, they are however the
more primitive form of limb. Those Crustaceans, on
the contrary, in which the first trunk limbs have lost
all traces of the ventral parapodia, and are purely loco-

SECT. Ill THE APPENDAGES 45

motory, are the more highly specialised. The max-
illipedes are secondary modifications only in the
degeneration of their dorsal parapodia which, as the
limbs of the first free segment, typically developed
into large locomotory limbs.

In Apus, the gnathobases of the anterior trunk
limbs are retained, and are doubtless functional. In
Limulus those of the first pair even work in front of
the under lips ; the same is true of the Trilobites, and
of the Eurypteridae, in some of the latter they have
taken on almost the whole function of mastication.

The extraordinary number of the limbs, and their
gradual simplification from before backward, i.e., from
complicated Crustacean limbs to parapodium-like
swimming plates, has been already discussed and
described.

The following points are also of interest. Unlike
the limbs of the head, in which, for the purpose of
pushing food into the mouth, the ventral parapodia
were developed at the expense of the dorsal, the
trunk limbs develop the dorsal at the expense of the
ventral parapodia.

Apus shows its primitive character in that the
ventral parapodia do not disappear, but are retained
as functional appendages to the limbs for pushing
food towards the mouth. In the higher Crustacea,
however, they have, as a rule, entirely disappeared,
except in a few anterior trunk limbs.

The reason of the greater development of the
dorsal than of the ventral parapodia in the trunk
limbs of Apus may be conjectured ; the greater sweep

46 THE APODID,£ PARTI

was needed both for locomotion and for the purpose of
grasping and bringing prey towards the middle line.
This bending round of the dorsal parapodium towards
the ventral middle line naturally leads to its greater
development, at least in the anterior limbs, which
function as described ; in the posterior limbs, which
have only to assist in swimming, the dorsal branch is
not so highly developed, forming with the ventral the
swimming plate (Fig. 10).

The anterior limbs are the most highly differentiated
from the original parapodia, they are the most
Crustacean. On the principle of the division of
labour, we find the first limb almost entirely specialised
as a sensory limb. Its appendages, omitting the
ventral parapodium, formerly considered as an
appendage of the limb, are, on the inner side, feelers
(called endites) with tufts of sensory hairs on slight
protuberances, which are regularly arranged alter-
nately on the two sides (see Fig. 9). The feelers vary
in length in different species, and are of importance
in classification. The corresponding appendages on
the other limbs are leaf-shaped plates with highly
developed denticulate setae : on their flat surfaces, and
tufts of sensory hairs along their edges. Besides
helping in swimming, these sensory endites have
been specially developed to assist in the capture of
prey. The action of sweeping together prey into the
middle line by means of the long dorsal parapodia,
requires a fine sense of touch on the under edges of
the limbs. The denticulate hairs at the sides of the

1 See Fig. 5.

SECT. Ill

THE APPENDAGES

47

endites may have been developed in order to prevent
prey from getting away between the limbs, the
endites being provided with special muscles to move
them. The prey, which the dorsal parapodia succeed
in raking into the middle line, is probably held,

g- -

d'

FIG. 9.— First trunk limb (L. Spitzbergensis). Lettering as in former figures.
e, sensory endites.

and perhaps killed, by the ventral parapodia, which
are provided with sharp thorn-like setae, as well as
with long sensory hairs (Figs. 9 and 10). We can
further judge from their shape and from the direction of

48

THE

PART I

their muscles (see Fig. 12, p. 55) that what the
gnathobases held between them would be rapidly
forwarded towards the mouth. The disappearance of
the gnathobases, excepting on the most anterior
trunk limbs — the maxillipedes — in the majority of
modern Crustacea, is explained by a change in the
manner of life. As the trunk limbs became more and

r""n.

FIG. 10. — The last rudimentary trunk limb (L. productus), \ mm. in length. Lettering
as before. It is seen to be a typical Phyllopodan limb.

more purely locomotory organs, such as ambulatory
legs, and as other methods of capturing prey were
acquired, the ventral parapodia would degenerate, not
only from having lost their function, but as positive
hindrances to the free movement of the limbs.

The most distal appendage, on the first limb of
Apus, is a probably functionless scale (Fig. 9), and in
the following limbs it is a toothed claw (Fig. 4), while

SECT, in THE APPENDAGES 49

in the posterior rowing limbs it is a flat piece forming
part of the swimming plate (Fig. 10). What we here
call the most distal appendage is, however, according
to the homologies described on p. 32, in reality the tip
of the dorsal parapodium.

The exites, or appendages on the back of the limb,
are always two in number, the distal being a
swimming plate (flabellum, homologous with the
sensory cirrus of the Annelidan parapodium), much
reduced on the first limb but well developed on all
the rest, and the proximal a gill which corresponds
in position with the gill on the dorsal parapodium of
the Annelid.

The limbs themselves have been so often described
that it is not necessary here to do more than refer to
the drawings (Fig. 9, 4, 5, 10).

Returning now to the homologies here set forth
between the Crustacean limbs and the Annelidan
parapodia, we find that they differ considerably from
those generally put forward. Although as far as I
know the point has not been worked out in detail, it
is assumed that the exopodite is homologous with the
dorsal, and the endopodite with the ventral para-
podium. We have here, however, seen that in the
typical biramose trunk limb the ventral parapodium
has disappeared, perhaps being used up in the forma-
tion of the basal joint, and that the limb proper is
formed of the dorsal parapodium, the distal end of
which forms the endopodite, while its sensory cirrus
forms the exopodite.

Such a result might be expected on theoretical

E

50 THE APODID^ PART I

grounds ; an integumental fold, developing outwards
into a limb in order to fulfil various functions, would
require to be provided with sensory organs. Its
efficiency could in fact only be secured by its being
provided with a fine sense of touch. It is true that in
Apus the ventral edge of the dorsal parapodium has
itself developed organs of touch, viz., the endites.
The tip of the parapodium might also have developed
its own sensory organs. Economy, however, would
certainly lead to the use of whatever sensory organs
happened to be already present.

This homology of the flabellum of Apus with the
sensory cirrus of the Annelidan parapodium, finds
some support from a study of its finer structure. Its
fringe of sensory hairs, its numerous ganglion cells and
conspicuous nerve fibres, show it at once to be a
sensory organ. Its flattened shape presents no diffi-
culty, since it is clearly thus modified to help the
animal in swimming. Packard thinks that it takes
a special share in respiration, but this we do not
believe, for the gills, in their inner structure and in
their freedom from hairs, show that they are specialised
for that purpose.

Further, the consequent homology of the exopodite
of the typical Crustacean limb with the sensory cirrus
of the dorsal parapodium of the Annelida, receives
considerable support from a study of any series of
Crustacean limbs, such, for instance, as those given in
Lang's Text-book of Comparative Anatomy. It is im-
possible to avoid the impression that as a rule (with
no doubt many exceptions) the exopodite is a sensory

SECT, in THE APPENDAGES 51

organ attached to the limb. We shall see further
reasons for this homology in the section on the
Nauplius.

In later sections we shall see, further, that although
the ambulatory limb of the Decapoda has been
derived from the Phyllopodan limbs of their Lepto-
stracan ancestors, yet ambulatory limbs may develop
straight from Annelidan parapodia, as we assume to
have been the case in the Trilobites. The method of
differentiation is the same in both cases, but the fact
that the Decapodan limb first passed through a
Phyllopodan stage has made a slight difference in
the result (see section on the Trilobites).

This brief discussion on the limbs of Apus and of
the Crustacea is by no means exhaustive. We have
purposely limited ourselves chiefly to our main point,
viz., how the limbs of Apus have been derived from
the parapodia of an Annelid. In so doing we have
naturally had our attention called to several homolo-
gies which may not at this stage appear altogether
satisfactory, but which will be found to hang together
with our whole argument.

E 2

SECTION IV
THE MUSCULATURE

THE musculature of the Apodidae is so essentially
Annelidan in its arrangement, showing only such
differentiations as we should expect would arise from
the modification of the body already described, *>.,
the bending of the head, that were there no other
resemblance between the Apodidae and the Annelida,
it alone would be almost sufficient to establish their
relationship.

It may perhaps be interesting to mention that it
was the Annelidan character of the musculature of
Apus which first attracted our attention, and led to
the discovery of the other homologies recorded in this
volume.

Anteriorly, where the body has been apparently
most modified, we should naturally expect that the
Annelidan character of the musculature would be
least recognisable. ' This, however, is hardly the case,
for just as the head of the Apodidae can be traced to
the anterior segments of an Annelid fixed in the

SECT, iv THE MUSCULATURE 53

bent position, the only striking alteration being its
development of the sharp ridge round the front as a
continuation of the lateral edges of the dorsal fold,
so the musculature can easily be traced back to that
of a typical Annelid transformed, first by the bend-
ing of the body, and secondly by the development
of the exoskeleton.

We shall first describe the musculature in a car-
nivorous Annelid, and see what transformations it
would undergo owing to the bending of the five
anterior segments. Fig. 1 1 is a transverse section
of such an Annelid. A rather weakly developed
circular muscle layer is found immediately under the
hypodermis, and under this runs a strongly developed
longitudinal muscle layer, the two forming together
the dermo-muscular tube. The development of para-
podia leads to important modifications, such as the
grouping of the longitudinal muscles into four strong
bands, two dorsal and two ventral, each being a chain
of segmentally arranged muscular bundles marked off
by the transverse dissepiments. The circular muscles
are also modified, running out laterally, both dorsally
and ventrally, into the parapodia.

At the posterior end of the body where the
parapodia are less developed, we might expect that
the muscle bands would gradually spread out to
form a more and more complete clermo-muscular
tube, the dorsal bands eventually uniting with the
ventral in the last segments.

It is not difficult to describe the changes which would
naturally take place in this musculature by the fixing

THE APODID^E

PART I

of the anterior segments in the bent position. The
dorsal bands in the five anterior segments would be
much stretched in order to bend round over the
intestine to be attached near the prostomium. The
ventral bands in the same region would, on the con-
trary, be much shortened, the bend behind the lip
being very sharp. The muscle bands here would

FIG. ii. — Transverse section through the trunk of a carnivorous Annelid, diagram-
matic (from Lang's Text-book of Comparative Anatomy), g, gill ; d, dorsal ;
v, ventral parappdium ; c-[, cirrus of the dorsal parapodium ; c», ditto of the
ventral parapodium ; cm, circular musculature ; lin, longitudinal musculature ;
tin, transverse musculature ; ac, aciculum ; n, nephridium ; e, developing eggs,
some of which are floating free in the body cavity.

be rendered almost useless, in fact, would be a
hindrance, and would therefore degenerate, not,
however, without leaving some traces. While the
muscular elements disappeared, the sinewy elements
would persist as points of attachment for those

SECT. TV THE MUSCULATURE 55

muscles which are still functional, i.e. those which
diverge in the transverse plane.

Let us now compare this sketch with the muscula-
ture of the Apodidae (Figs. 12 and 13). The dorsal
longitudinal bands in the head region, after springing
across the opening into the interior of the shell fold
of the fifth segment, are attached to the forehead by
numberless fine fibres of connective tissue, so that it

FIG. 12. — Section through a specimen of Apus cancriformls, partly diagrammatic, to
show the longitudinal musculature. Anteriorly the dorsal bands are stretched
round the bend of the head, the ventral bands of the five segments being clumped
into a sinewy mass, the sternal plate. Posteriorly the two unite round the body
to form a dermo-muscular tube. <////, dorsal ; v»i. ventral muscle bands ; s, shell ;
e, eye ; /, under lip ; mi, in*, ist and 2nd maxillae ; ?/, ventral parapodia of trunk
limbs ; 6 ditto of ist trunk limb. The five original anterior segments of the
Annelid indicated by dotted lines.

is not at first apparent that the two strong bands
which start a little in front of them from very
scattered points of insertion on each side of the eyes,
running down over the oesophagus to be attached in
front of the prostomium, are really a continuation of
the dorsal bands. That this is the case, however, is
clear from Fig. 1 3. We see here not only the dorsal
bands themselves lengthened to pass round the curve

THE APODID^E

PART I

formed by the bending of the head, but the attach-
ments of the muscles also spread out over a large
surface. Further, partly from having no segments to
move, and partly on account of the growth of the
frontal ridge, they apparently pass through one or two
segments without being attached at all.

dm

FIG. 13. — Diagram to show the musculature of the head. Lettering as in Fig. 12.
ind, mandibles ; z, intestine.

The ventral muscle bands of the Apodidae, on the
other hand, on reaching the sharp bend of the head
near the lip, are all clumped together into an ap-
parently shapeless sinewy mass ; the muscle bands

SECT, iv THE MUSCULATURE 57

themselves have clearly disappeared ; their sinewy
connections, however, being more resistant, and still
functional as attachments for the mandibular, maxillar,
and cesophageal muscles, &c., have been retained,
massed together as described in our imaginary
Annelid. This sinewy mass is known as the sternal
plate or entosternite. It is clear that, if the origin we
have ascribed to it is correct, it becomes a morpho-
logical characteristic of great value, and, whenever
met with among the Crustacea in the same position,
i.e., within the angle of the bent intestine, must be
referred back to the ventral muscle bands of a bent
Annelid. We shall see in the second part how im-
portant this point is in establishing the relationship
between Apus and Limulus. The Arachnida are,
we believe, the only other animals with an entoster-
nite. The origin and significance of it in this case
will be discussed in another section.

Again, turning to the posterior end of the body,
we find, as we expected, that as we pass from front
to back the longitudinal bands gradually widen out,
as the limbs are less and less developed, until, in
the limbless segments, they unite to form a simple
dermo-muscular tube.

Thus, in the arrangement of its longitudinal
musculature, Apus is a typical carnivorous Annelid
with its five anterior segments bent round in adapta-
tion to the browsing manner of life.

On turning now to the circular musculature, we find
a more complete differentiation. The circular muscle
layer in the carnivorous Annelids is, as a rule.

58 THE APODID^: PARTI

much more weakly developed than the longitudinal.
This is also the case in the Apodidae, where it is
almost entirely confined to the muscular bands which
run into the limbs, especially to those from the dorsal
surface. In the limbless part of the body, where the
longitudinal muscles form a complete clermo-muscular
tube, the circular muscle layer has entirely dis-
appeared. The commencement of the formation of
an exoskeleton renders it useless. We shall return
to this subject in discussing the musculature of the
limbs.

The muscles attached on each side, just above the
ventral cord, to the membrane which encloses the
intestine and genital glands, and forms the intestinal
sinus, may perhaps best here be mentioned as in part
having arisen from the circular musculature. We
shall return to these also when we discuss the cir-
culation and the origin of the above-mentioned
membrane.

Two especially interesting groups of muscles, of
unmistakably Annelidan origin, deserve particular
attention. These are the rows of dorso-ventral
muscles (Fig. 14, dv.) which pass between the intes-
tine and the genital glands in almost exact corre-
spondence with the longitudinal muscle dissepiments
so common among the Annelida (cf. Fig. 1 1, tni}. In
the Apodidae, these rows are composed of a kind of
lattice work of muscle bundles with definite points
of attachment, ventrally to the sinewy partitions of
the ventral muscle bands and thus indirectly to the
body wall, and dorsally to the segmental con-

SECT. IV

THE MUSCULATURE

59

strictions in the integument. In each segment we
find two bands crossing each other diagonally, in
addition to those placed between the segments. These
strikingly Annelidan dorso-ventral muscle bands of
Apus are not, as far as we know, preserved in any
of the higher Crustacea, being rendered useless by
the development of the exoskeleton. Clear traces of

Til

FIG. 14. — Transverse section through Apus cancnformis, to show the distribution of
the musculature. ft, heart ; rf»/, dorsal ; vnt, ventral muscle bands ; c»r. circu-
lar musculature (as shown in Fig. 15, A)\ dv, dorso-ventral musculature (cf.
Fig. n, tut) ; tit, membrane enclosing the intestinal and genital sinus ; /, intes-
tine ; g, genital glands ; e, eggs ; a and b have reference to Fig. 15.

them are, however, found in Limulus, where their
points of attachment have drawn in the outer integu-
ment to form the entapophyses. Their use in Apus
will be discussed in the sections on circulation
and reproduction. They are not developed in the
posterior rudimentary segments.

The muscles of the limbs present complications

6o

THE APODIDyE

PART I

which might have been expected, when we take into
account the transformations which have developed
the latter out of parapodia. It will not be necessary
to describe the muscles of the more distal parts of
the limbs ; we must confine ourselves to those which
move the limbs on the body, and endeavour to show

FIG. 15. —Tangential sections through three segments posing between the longi-
tudinal muscle bands and the lateral body wall, diagrammatic, the plane of
the section passes through the points a and b in Fig. 14, A to show the
circular musculature ', B to show the arrangement of the longitudinal muscu-
lature ; C to show the crossing of the longitudinal muscle bands.

how they have arisen as simple modifications of the
original dermo-muscular tube.

In the first place, we find that the muscles of the
limbs have for the most part the same character as
those of the trunk, i.e.y they are bands with broad

SECT. IV

THE MUSCULATURE

61

surfaces of attachment. They are, in fact, so far at
least as they are composed of longitudinal muscles,
nothing but the elements of the dermo-muscular tube
drawn out of their position, as may be clearly seen
from Fig. 12, which gives a general view of the whole
musculature. The same fact might also be concluded
from their great number, disorder, and want of con-
centration. When, however, we have to decide which
muscles belong to the circular and which to the
longitudinal layer, the following seems to be the

FIG. 16. — Diagram to explain the courses of the muscle bands in B and C, Fig. 15.
i represents an Annelidan parapodium in its original horizontal position with
three longitudinal muscle bands running into it. 2 represents the same drawn
down to form a limb of Apus.

principle of arrangement. The muscles which enter
the dorsal side of the limb with broad insertions on
the soft integument of the dorsal surface (Fig. 15, A)
are probably elements of the circular muscle layer ;
their position close under the lateral integument
favours this derivation (Fig. 14, cm. ; cf. also Fig. ii).
On the other hand, we find on each side of the limb
a number of muscle bands with more definite points
of insertion ; these are attached dorsally to the sides

62 THE APODIDyE PARTI

of the septa (see Fig. 15, B\ and are probably
longitudinal muscles. The order in which these
latter groups of muscles occur is significant of their
origin, as shown in the diagram (Fig. 16). We find
that those which arise most dorsally run the furthest
into the limbs, this rule being regularly observed.
This order is what we should expect if we assume
that these are parts of the longitudinal musculature
which ran outwards into the parapodium, the dorsal
edge of which was then gradually lengthened, and
the whole turned round the body in the transverse
plane towards the ventral middle line, as shown in
Fig. 1 6. The dorsal muscle bands will naturally be
the most lengthened and reach the furthest down
into the limb ventrally. In the case of longitudinal
muscle bands running outwards into the parapodia,
but traversing them from the anterior to the posterior
wall, the same rule would hold and the bands would
cross one another, as shown in Fig. 15, C.1

The musculature running into the ventral part of
the limb or the ventral parapodium is more easy to
separate into its elements (see Fig. 14). The longi-
tudinal muscles come direct from the ventral muscle
bands, and run sloping backwards, as shown in Fig. 12,
so that the ventral parapodia or gnathobases which
slope away downwards and backwards may be used
for pushing food forwards in the middle line. The

1 We have, however, only once found such muscle bands, and have
since repeatedly looked for them in vain. It is not unlikely that our
observations relating to them were incorrect, being founded on a series
of sections, through which the individual muscle bands had to be
followed,

SECT, iv THE MUSCULATURE 63

circular muscles of the ventral parapodium are as
inferior in development to those of the dorsal para-
podium, as the ventral parapodium itself is inferior
in development to the dorsal. They consist of only
two bands. One passes between the ventral cord and
the ventral muscle band, to be attached proximally
to the ventral membrane of the intestinal sinus, the
other is attached direct to the hypodermis at each side
of the ventral cord (Fig. 14). The former muscle will
be mentioned again in describing the circulation, in
which it perhaps plays a more important part than
it does in connection with the limbs.

It is hardly necessary to describe the musculature
of the trunk limbs more in detail. That of the head
limbs, however, requires special attention, not only on
account of the origin of these limbs almost exclusively
from ventral parapodia, but also because the masti-
catory formula of the Apodidae is, with slight
differences, the same as that found in the majority of
modern Crustacea, viz., one pair of mandibles and
two pairs of maxillae ; although in Apus, the second
maxillae are rudimentary.

Commencing with the mandibles, we there find
an arrangement exactly the opposite of that de-
scribed in connection with the trunk limbs. In
these latter the muscles running into the dorsal
branch are the more highly developed ; in the man-
dibles, however, the dorsal branch is rudimentary,
and the muscles running into the ventral branch
are the most developed. The closing muscles radiate
from the sinewy mass above described, and are

UNIVERSITY

64 THE APODID^E PART I

enormously developed in accordance with the great
development of the limb they have to move.
They evidently correspond with the muscles already
described as running into the ventral parapodia of the
other limbs, that is with those which come from the
longitudinal muscle band. They radiate from the
sternal plate, i.e. from the remains of the ventral muscle
bands of the head segments. At the dorsal extremity
of the mandibles, we find the remains of the circular
muscles which (see Fig. 15, A) were so powerfully
developed in the trunk limbs, in two or three bands
running between the dorsal middle line and the
integument, where the last rudiment of the dorsal
parapodium has disappeared (see Fig. 8, A, dy p. 37).
The longitudinal muscles, attached to the integu-
mental folds between the limbs (see Fig. 15,^), are
strongly represented, and probably serve both for
closing the mandibles and rotating them round their
longitudinal axes.

The same description applies with but slight
modification to the muscles of the first maxillae, but
in this case, those of the ventral parapodium, though
strongly developed in comparison with those of the
ventral parapodia of the trunk limbs, are weak as
compared with those of the mandibles. Again, a
more distinct rudiment of the dorsal parapodium is
retained in the first maxillae than in the mandibles,
and into this rudiment a very long and tolerably
strong band runs, probably homologous with the
circular muscle bands shown in Fig. 15, A. The
powerful muscles which enable the first maxillae to

SECT, iv THE MUSCULATURE 65

function as jaws are also, like the closing muscles of
the mandibles, derived from the sinewy mass, and
run slantingly backwards, across the opening leading
into the under lip.

The muscles of the second maxillae are very slightly
developed as thin slips running into the dorsal and
ventral parapodia ; the former, as already described,
arising from a point close to the opening of the shell
fold.

The rings of muscles round the eyes will be de-
scribed in the section on the sensory organs, and we
shall see that they are developed from the longitu-
dinal musculature, and join the two bands which are
attached to the proximal end of the upper lip. Certain
bands which run from the sternal plate to the open-
ing of the shell fold, to join the dorsal longitudinal
bands, are probably to be referred to the dorso-ventral
longitudinal muscle septa.

We find the expected histological difference between
the musculature of Apus and that of the Annelids,
that of the former being striped, that of the latter
unstriped. Perhaps the primitive character of the
striped muscles of Apus may be seen in that the
muscle cells form a thick irregular layer of nucleated
protoplasm round each bundle of fibres, without any
investing membrane or sarcolemma.

This brief chapter by no means exhausts this
interesting subject ; further study will doubtless
reveal other, and perhaps more conclusive, homologies
between the muscles of Apus and those of a car-
nivorous Annelid. We have here selected only the

F

66 THE APODID^: PART

most obvious ; enough, however, to establish our point
that Apus may have been derived — at least so far as
its musculature is concerned — from such an Annelid
as we have described. We thus find that the
musculature confirms what we learnt from our study
of the outer organisation and of the appendages.

SECTION V

THE NERVOUS SYSTEM

THE nervous system of Apus does not at first sight
seem to support our theory as obviously as does the
musculature. This, however, is the case only at first
sight. A closer study of it, and a comparison of it
with that of an Annelid modified by having its five
anterior segments bent in the way assumed, leave but
little doubt concerning its origin. The central nervous
system of Apus can in fact be shown to be the central
nervous system of a bent Annelid adapted to the
necessities of a new manner of life ; the principal
modification being due to the migration of the eyes on
to the dorso-frontal surface.

Figure 17 shows the general type of the nervous
system of a carnivorous Annelid, such as the ancestor
of Apus may be supposed to have possessed. The
longitudinal commissures may perhaps have been
somewhat wider apart. l We find the brain in the

1 See however p. 80.

F 2

68

THE

PART I

prostomium giving off two pairs of nerves to the
two pairs of eyes, and connected by cesophageal
commissures with the infra-cesophageal ganglion in
the first segment. From this ganglion the nerves to
the first antennae diverge ; they may perhaps have
been united for some distance with the cesopha-
geal commissures. It is even possible that their

FIG. 17. — Diagram of the first five segments of a carnivorous Annelid to sho\y the
arrangement of the nervous system, from above. f>, brain ; £•_>, anterior pair of
eyes on the prostomium ; e\, posterior ditto : aj, first antennae ; a-2, antennal para-
podium of the second segment.

ganglia may have moved forwards along the com-
missures towards the brain, as in many Annelids we
find the antennae moved forwards till they appear to
be projecting from the posterior edge of the prostomium.
The second antennae, belonging to the second segment,
would receive their nerves from the second ventral
ganglion, then would follow the nerves to the para-
podia of the third segment, &c., in order.

SECT. V

THE NERVOUS SYSTEM

69

Just as the sharp bending of the head led to a
condensation of the ventral musculature into the
sinewy mass above described, so it would naturally
lead to a fusing of the anterior ventral ganglia, as
shown in Fig. 18. We should thus expect to find at

2

FIG. 18. — Anterior end of the same, bent as in Fig. i to show the change in the
central nervous system due to the bending of the body.

least the first three or four pairs of ganglia of the
ventral chain fused to form one infra-cesophageal
ganglion ; the outgoing nerves, however, would remain
distinct, except perhaps the first antennal nerve, which,
as we have said, might have been fused for a short way
with the cesophageal commissures, or might even, as

70 THE APODID^E PART i

above stated, come almost direct from the brain. The
other changes, brought about by the bending of the
segments, would be the disappearance of the longi-
tudinal commissures between the four or five fused
ganglia, and perhaps a fusion of at least some of their
transverse commissures. We shall see in the second
part of this book, when we come to compare Limulus
and Apus, that the nervous system of the former,
though showing certain special modifications of its
own, corresponds, to a remarkable degree, with that
of such a bent Annelid, and thus shows even a more
primitive state than that of Apus.

Now let us consider the modification such a central
nervous system would undergo owing to the gradual
migration of the eyes on to the dorsal surface. Figs.
19 and 20 are two diagrams to illustrate the change ;
Fig. 19 supposing the ganglion for the first antenna to
come from the infra-cesophageal ganglion, Fig. 20 sup-
posing this ganglion to have already migrated along
the commissures to near the brain. The brain, follow-
ing the eyes, would divide the original cesophageal
commissures (ce^) longitudinally, thus producing two
cesophageal commissures, one (<2?2) in its original
position, innervating the oesophagus and "the upper lip,
and the other (^3) carrying the brain and the eyes.

This origin of the two cesophageal commissures in
Apus is especially interesting because it explains the
origin of the sympathetic nervous system in the Crustacea.
Reserving, however, this point for the present, we
have to consider the more difficult problem relating to
the position of the antennal nerves, and how they

SECT, v THE NERVOUS SYSTEM 71

would be affected by this splitting of the cesophageal
commissure owing to the travelling backwards 1
of the brain. Taking first the case illustrated in Fig.
19, i.e., assuming that the nerves for the first antennae
branched, in the original Annelid, from the first
ventral ganglia, we tried to answer this question
theoretically. Our answer, however, was not quite
correct. We assumed that the first antennal nerve was
originally united with the cesophageal commissures for
a short distance, and would remain where it was when
the brain dragged away the portion it required for
itself. We were doubtless also misled by the position of
the first antennae of Apus near the prostomium. These
mistakes were very natural. For the second antennae,
however, our answer wras correct. We rightly assumed
that as the brain and cesophageal commissures moved
forwards and upwards, passing through the position
occupied by these antennal nerves, the two might
unite, so that we described the nerves for the second
antennae as branching off from the brain commissures,
this position agreeing best with the position of the
second antennae in Apus.

On comparing this theoretical scheme for the antennal
nerves with Zaddach's drawings, we found, as stated,
that the nerve for the first antenna, which has the
more ventral position, branched off from the brain-
cesophageal commissure dorsally to the nerve of the
second antennae \vhich has the more dorsal position,
so that, if Zaddach's drawings were correct, a slight

1 "Backwards " is morphologically correct ; actually the brain moved
forwards and upwards.

THE APODID&

PART I

crossing of the nerves must take place. Our own
examination of the nervous system quite confirmed
this, as shown in Figs. 19, 20, and 21. These figures

FIG. 19. — Diagram to show the derivation of the central nervous system of Apus from
that of a bent Annelid as in Fig. 18 ; drawn on the assumption that the nerves
of the first antennae of the original Annelid branched from the first ventral
ganglion, i, 2, 3, 4, 5, ganglia of the ist, 2nd, 3rd, 4th, and 5th segments, b,
brain of Annelid ; /3, of Apus ; ce\, cesophageal commissures of Annelid ; o?2, «3,
the two cesophageal commissures of Apus derived from ce\ ; a\> a%, nerves to the
ist and 2nd antennae of the Annelid ; cq, 0.0, ditto of Apus ; e\, e-2, nerves to the
eyes of the Annelid ; 771, rjo, ditto to those of Apus.

should be compared with Figs. I and 2, which show
the positions of the antennae.

This position of the first antennal nerve tells us,
however, nothing definite as to the position of the

SECT. V

THE NERVOUS SYSTEM

73

ganglion. All we can positively affirm is that, if
the ganglion was infra-cesophageal in the Annelid,
the proximal portion of the nerve was carried up
with the cerebral portion of these commissures

FIG. 20. — Diagram showing the same as Fig. 19, drawn, however, on the assump-
tion that the nerves for the first antennae in the original Annelid came from the
posterior end of the brain. Lettering the same as in Fig. 19. x shows the
position of the ganglia of the first antennae according to Pelseneer.

which split off and travelled backwards. This would
explain the apparently anomalous position of the
points of departure of the antennal nerves from
the brain-cesophageal commissure, — they have to cross
each other to reach their destinations. As these

74 THE APODID^E PART I

commissures travelled upwards, describing part of a
circle, carrying up with them the two pairs of antennal
nerves, the pair of nerves which originally had the
more ventral position would naturally come to occupy
the more dorsal position, as shown in the diagram,
Fig. 19.

If now we assume, as shown in the diagram, Fig. 20
that in the original Crustacean-Annelid the ganglia
of the first antennae had already travelled up the
cesophageal commissures to near the brain, then we
have to suppose that these ganglia split away with
the brain-cesophageal commissures, although, by
so moving off with the brain, they were dragged
further from the limb their fibres had to innervate.
This latter assumption, as shown in Fig. 20, agrees
best with the description of the central nervous system
given by Pelseneer. He assumes that a group of
ganglion cells, in the position marked x in Fig. 20,
form the ganglia for the first antennas, and he supports
this claim by the fact that the nerves branch back-
wards, as shown in Figs. 20 and 21. 1 If this reason-
ing is correct, then we may assume either (i) that the
migration of the ganglia had already taken place in

1 Quarterly Journal of Micro. Sc., vol. xxv. Although inclined
to believe Pelseneer's view to be correct, his arguments do not
seem to us quite conclusive. The results of our own research
unfortunately remained neutral. We should much like the point re-
examined ; perhaps the new method of staining the nervous system of
living animals with methyline blue would reveal the actual courses of
the fibres. In our own best hasmatoxylin preparations the fibres
became suddenly quite confused where the antennal nerve joined the
commissure, and we could not say whether they ran on to the brain, or
bent back towards the infra-oesophageal ganglion.

SECT, v THE NERVOUS SYSTEM 75

the original Annelid, and is inherited by Apus, or
(2) that the formation of the compound sensory
nervous centre (the syncerebrum of Lankester) has
taken place in Apus by the wandering of the first
pair of ganglia to join the brain. The former seems
to us the more natural conclusion, considering
the great difference in the distances between the
brain and antennae of an Annelid, and between
the same parts in Apus. We mean that the great
distance between the cerebral position of the ganglia
of the first antennae and the antennae themselves in
Apus, which seems unnatural, is best explained by
assuming that this cerebral position of the ganglia
was derived from the Annelid, where, owing to
the proximity of the antennae to the brain, it is
most natural. On the other hand we think the
second view the less probable, considering (a) the
weak development of the first antennae in Apus, and
(^) the distance of the eyes from the antennae, and the
difficulty of correlating their respective sensations, the
eyes pointing forwards and upwards, the antennae
backwards and downwards.

In the higher Malacostraca, with well-developed
antennae placed close to the eyes and functioning as
auditory, olfactory, and tactile sensory organs, there
would be no difficulty in imagining the migration of
the ganglia to have taken place in the course of their
development. But, as already stated, it is difficult to
imagine this in the case of Apus, and it is easier to
suppose that the fusion of the antennal ganglia with
the brain had already taken place in the original

76 THE APODIDyE PART i

Annelid, We may perhaps find some support for
this view in the fact that the first antennae never
appear in the Crustacea as anything but uniramose
which shows that, in the original Annelid, they had
long lost all traces of the parapodia to which they
primitively belonged, and were nothing but sensory
organs projecting forwards on each side of the pro-
stomium.

Some further light might perhaps be thrown on this
point by a study of the central nervous system of
Limulus, which shows in some respects a more primi-
tive condition than that of Apus, at least as regards
the position of the brain. According to Packard, the
fibres of the first antennal nerve do not come from the
brain, but from the cesophageal commissures near it.
Owing, however, to the great modification of the
cesophageal commissures of Limulus, in consequence
of the lengthening out of the oral aperture, it is doubt-
ful whether this fact supports the view illustrated in
Fig. 20, that, in the original Annelid, the ganglia of
the antennae had moved to near the brain.

There are, however, other points which bear on this
question. On examining the first section of the ven-
tral cord of Apus, we find a long ganglion consisting
of two groups of ganglion cells, and joined by two
transverse commissures. From the long ganglion, the
prostomial-cesophageal commissures run down to
embrace the cesophagus. Before reading Pelseneer's
paper, we were inclined to consider the front group of
ganglion cells, which form part of the long ganglion,
as belonging to the first antennae. It did not occur

SECT. V

THE NERVOUS SYSTEM

77

to us that they belonged to the prostomial com-
missures, or stomato-gastric ring, as suggested by
Pelseneer, because we considered this ring as no
special nerve branching from the cord, but simply as
the remains of the original cesophageal commissures.

FIG. 21. — Central nervous system of Apus. Lettering as in Figs. 17, 18, and 19.
cm, nerve to the eye muscles ; e*, the anterior pair of eyes transformed into the
unpaired " eye " (cf. § on the sensory organs); * the position of the ganglia of
the ist pair of antenna; according to Pelseneer.

As for the two transverse commissures, we consi-
dered them to represent the two commissures of the
two pairs of antennal ganglia. We assumed that each
was due to a fusion of two transverse commissures,
at least if we might conclude from the fact that the

78 THE APODID^E PART i

other ventral ganglia in Apus have two transverse
commissures, that the transverse commissures of the
antennal ganglia were also originally double. On this
supposition, passing from the original Crustacean-
Annelid to the higher Crustacea, we should have two
separate fusions. First, in Apus, there is a fusion of
the double transverse commissures of the two pairs
of antennal ganglia, leaving two transverse com-
missures, one for each pair of antennal ganglia ;
then, in the higher Crustacea, these two single
transverse commissures of the two pair of ganglia
again fuse, so that there is only one transverse com-
missure joining the cesophageal commissure in front of
the infra-cesophageal ganglion. This transverse com-
missure, which is always assumed to be that of the
ganglia of the second antennae, would be, according to
this view, the fused transverse commissures of the
ganglia of both pairs of antennae, though the ganglia
themselves have wandered towards the brain.

We can see no inherent difficulty in thus deducing
the central nervous system of Apus from that of a
bent Annelid ; none of the assumed transformations
are in themselves improbable, if the migration of the
eyes is once admitted. We shall return more than
once to this subject of the migration of the eyes,
especially in connection with the Nauplius and
Limulus, in each case bringing forward fresh evidence
in support of the assumption. In the meantime it
seems to us that the central nervous system of
Apus, taken as a whole, bears incontestable witness

SECT, v THE NERVOUS SYSTEM 79

to the fact that the eyes have thus travelled on
to the dorsal surface. Zaddach's diagram gives the
brain and cesophageal commissures a distinct curve
backwards, so that they come to lie along the
oesophagus, reaching as far back as the mid-gut.
The brain lies between the hepatic-diverticula.

In discussing the appendages, we saw how the
antennae, which were originally metastomial, became
prostomial by the bending of the head. We now
see that the same change of position has taken place
in the case of the points of departure of their nerves
(at least of those of the second pair). By the sweep-
ing round of the cerebral portion of the cesophageal
commissures upwards and backwards, and by their
carrying the antennal nerves along with them, these
nerves have also acquired a prostomial position.

This derivation of the central nervous system of
Apus from that of a bent Annelid throws new light
upon the fact, established by Claus and Dohrn, and
referred to by Balfour with evident surprise, that, in
the Nauplius, the nerves for the second antennae arise
from the infra-cesophageal ganglion. This is of
course what we should have expected, indeed from
our point of view it is necessary to account for the
fact that the nerves of both the pairs of antennae do
not arise from the infra-cesophageal ganglion. We
are obliged to assume that this primitive Annelidan
condition was passed through in embryonic and not
in larval life, i.e., in an early Annelidan, not in a
Crustacean stage.

Of the rest of the nervous system little need be

So THE APODID^: PART I

said ; the Annelidan character of the ventral cord of
Apus has long been acknowledged as an unmistak-
able sign of relationship between it and the Annelida.
Lankester has also called attention to the fact that
the ventral cord resembles more nearly that of a
Chaetopod than that of a Crustacean. He sees its
archaic character in the fact that the longitudinal
strands are separated by a considerable interval.
This reasoning is however doubtful, because in the
rudimentary segments of Apus the ganglia in each
segment are close together. While it is true that
a great interval between the longitudinal halves of the
ventral cord of an Annelid is generally supposed to
denote an archaic condition, this state in Apus has
clearly been secondarily acquired. Further, the pre-
sence of well-developed parapodia, which were
gradually transformed into Crustacean limbs, is con-
clusive evidence that the Annelid from which Apus
was derived was not a primitive form. The drawing
out of the longitudinal commissures of the anterior
ganglia of the ventral cord which, in the bent
Annelid, were massed together (see Fig. 18), is clearly
a secondary modification, due to the travelling
forwards of the brain. It will be referred to again
in the next section in connection with the migration
of the eyes.

Till now, it has never been quite understood why
the ventral cord should suddenly cease with the
limbs, so that no ventral ganglia are developed in
the limbless segments. The explanation of this we
have already seen, viz., that the posterior end of the

SECT, v THE NERVOUS SYSTEM 81

body becomes fixed in a larval stage ; the posterior
limbs with their ganglia remain quite rudimentary,
while the last few segments develop neither limbs
nor ganglia.

Some light is also thrown on the morphology of
the sympathetic nervous system, which is particularly
well developed in the Malacostraca. The second
cesophageal ring formed by the sympathetic nerve is,
in fact, the remains of the original Annelidan ceso-
phageal ring, after the splitting off of the portion
which carried the brain. The present Crustacean
cesophageal commissures, together with the ring
made by the sympatheticus, formed the original
Annelidan cesophageal commissures.

We also get an interesting insight into the mor-
phology of the Crustacean brain. Originally, when
still placed in the prostomium, it consisted of the
ganglia of the two pairs of eyes, and of whatever
other sensory organs may have been on the prosto-
mium, and perhaps also of the ganglia of the first
antennae. These sensory centres (with the exception
of the last, which were probably situated on the
cesophageal commissures) were but collections of
hypodermal ganglia, as is clear from the fact that the
pair of longitudinal muscles which traverse the head
dorso-ventrally (see Figs. 12 and 13) pass between
the brain-cesophageal commissures (#3) ; this shows
that the ganglia must have had a hypodermal posi-
tion, i.e., must have lain between the hypodermis and
the musculature. On the migration of the eyes, the
optic ganglia would separate from the ganglia which

G

82 THE APODID^E PART i

belonged especially to the prostomium, taking the
antennal ganglia along with them on the com-
missures which continue to unite the brain with the
ventral cord.

The transformation of the anterior pair of eyes
into the unpaired " eye " with other sensory functions
would bring about secondary complications.

The gradual wandering of the ganglia of the first
antennae along the cesophageal commissures, or, if
these were already near the brain, their final union
with the same, added further complications.

Lastly, when the antennae, and especially the
anterior pair, adopted a frontal position on the head,
and became important sensory limbs carrying
olfactory, auditory (? directive), as well as tactile
sensory organs, so that their ganglia became large
complex sensory centres at the posterior end of the
brain, its complication was completed, and it
reached the stage found in the higher Crustacea
(e.g., Decapoda).

At first sight, this method of deducing the Crusta-
cean nervous system from that of a bent Annelid
may not appear to the reader altogether satisfactory.
We may therefore perhaps anticipate what we shall
describe in its right place, and mention that when we
drew Fig. 18, to show where the brain was originally
placed in the more Annelidan ancestors of Apus, we
had quite forgotten that this was still the place which
it occupies in Limulus. In Part II. we hope to be
able to show that, if Apus is derivable from a bent
Annelid, Limulus must also have had the same

SECT, v THE NERVOUS SYSTEM 83

origin. This difference in the position of the brain in
Apus and Limulus is one of those cases, referred to in
the preface in which the differences between these two
animals afford more striking proof of their relation-
ship, through a common origin from a bent Annelid,
than any similarity in the position of the brains could
have done.

G 2

SECTION VI

THE SENSORY ORGANS

IN our endeavour to deduce the sensory organs of
Apus from those of a carnivorous Annelid, we must
not forget that the development of an exoskeleton
must necessarily lead to striking modifications.
Such modifications, important in all the organs, are
especially so in those which, like the sensory organs,
lie at the surface in more or less immediate contact
with the outer world. We will take the sensory
organs in- turn, and discuss the changes which took
place in them during the transformation of the
Annelid into the Crustacean.

The antenna, as sensory organs, admit without
difficulty of deduction from the antennae and antennal
parapodia of the first two segments of the Annelid,
as we have already seen in the section on the
appendages.

The round white spot behind the eyes of Apus
has often been taken to be a sensory organ, and we

SECT, vi THE SENSORY ORGANS 85

originally assumed it to be the remains of a frontal
cirrus (as shown in Fig. i) smoothed off to facilitate
swimming. We have, however, discovered that its
functions are entirely excretory (see Section IX., on
the excretory organs, and Appendix III.).

Of the original four anal cirri of the Annelid, two
are retained and two are rudimentary. Those
retained have developed a ringed cuticle for the
greater part of their length, and so far are covered
with setae also arranged more or less in rings, those
on the inner side being longer than those on the
outer. The tips of these cerci are thin-skinned, and
function as tactile papillae ; this is indicated by
shading in the drawing of L. Spitzbergensis given as
frontispiece.

The two rudimentary cirri are reduced to papillae
on the dorsal surface of the anal segment ; they are
thin-skinned, and surrounded anteriorly by a rampart
of thorns ; from the centre of each rises a long branched
tactile hair.

The whole body is covered with hairs. We have
found at least four kinds.

(i) There are very fine hairs in groups of two and
three ; they are apparently longer (ca. 4^) on the inter-
segmental membranes than on the harder parts of
the cuticle (ca. 2/jJ). We have found them chiefly on
the exposed abdominal segments. It is very doubtful
whether they are sensory ; their great numbers and
minute size render it probable that they serve to
prevent the attachment of other organisms which
might hinder free locomotion. On the other hand,

86 THE APODID^: PART I

this very roughness might favour the attachment of
spores.

(2) There are, further, short straight hairs which
seemed to be sensory, but all our attempts to trace
their elements through the cuticle were baffled ; in
one place alone, where the cuticle was split from the
hypodermis, we saw fine processes connecting the
points where the hairs arose with the hypodermis,
and these may have been nerve fibres. These short
hairs are very numerous, especially in the frontal and
dorsal regions of the head.

(3) There are undoubted sensory hairs whose
nerves even with a low power are easy to follow into
the hypodermis, where they probably join the sub-
hypodermal nerve plexus.

(4) The sensory hairs and setae on the limbs may
perhaps be classified as follows.

(a) Tufts of minute hairs on small papillae round
the edges of the endites, and along the outer edges
of the gnathobases.

(ft) Long feathered hairs on the gnathobases, occur-
ring together with sharp tooth-like setae, which latter
help to give the gnathobases the character of jaws
(see Fig. 9, p. 47). The nerves of these highly
developed tactile hairs are easy to follow ; the
ganglia at their roots are compound (see Fig. 31,

P- ISO-

(7) A fringe of similarly feathered hairs round the
flabella, which we. have homologised with the sensory
cirri of the dorsal parapodia. The flabella being
transparent, the nerves can easily be followed.

SECT, vi THE SENSORY ORGANS 87

(8) Hooked or bent hairs on the first antennae
which we homologise with the olfactory hairs of
the higher Crustacea (see Fig. 7, p. 34).

To these may be added : —

(e) Fine sensory hairs thickly covering the inner
surfaces of both upper and lower lips.

The gill, as already stated, has no hairs, since these
would hinder the free flow of the respiratory medium.

Besides these different kinds of hairs, there are
stiff denticulate bristles in rows near the bases, on the
flat surfaces, of the rhomboidal feelers (the endites),
which probably hinder the escape of prey between
the limbs (see Fig. 5, p. 23, and p. 46).

While, perhaps, the finer sensory hairs may be
homologised with similar tactile hairs in the Annelida,
it is not easy to homologise the more highly developed
setae. If any of the original Annelidan setae have
been transformed into the hollow Crustacean hairs,
the transformation would have to be described some-
what as follows. The thickening of the cuticle
supplying a firm base for the seta, it would not be
necessary for it to sink below the integumental surface.
Again, the integument not being liable to be thrown
into folds like the soft skin of the Annelida,
the seta would not require to be movable ; hence
there is no need for it to project from a sac-like
group of secreting cells under the cuticle, provided
with muscular attachments. The Crustacean seta is
a hollow process of the cuticle secreted by a ring of
hypodermis cells, through which the nerve runs into
the lumen of the seta. At the base of the hair, the

THE APODID^: PART i

nerve swells into a ganglion. In the feathered hairs,
a fibre runs into each barb, and the ganglion is a
regular group of cells (see Fig. 31, p. 131). This
nerve may well be the nerve which originally supplied
the setiparous gland of the Annelid, and the ring of
secreting cells all that remains of the sac itself.

On each side of the under lip is a straight longi-
tudinal row of sensory papillae. As these are some-
times found thickly clogged with particles of food,
they probably border the channel leading into the
oesophagus on each side, to hinder juices, &c., from
escaping laterally under the mandibles.

THE EYES.

We are here brought face to face with a problem
of no small difficulty. It is clear that if our theory
is true, we have to attempt to explain the develop-
ment of the eye of Apus, i.e., the development of the
typical Crustacean eye from that of the Annelids. It
is hardly necessary to dwell upon the difficulty of
such a task, since it is but too well known that the
last word has not been said as to the actual structure
of either the Crustacean or the Annelidan eye. Still
we cannot turn from the attempt, especially as we
hope to be able to show that if we do not go
into too many details, and at the same time keep con-
stantly in mind the effect which the development of
a thick chitinous cuticle would naturally have upon
the hypodermal eye-spots, it is possible to sketch a
fairly probable origin of the Arthropodan eye.

SECT, vi THE SENSORY ORGANS 89

We confine our attention at present entirely to the
paired eyes, reserving the unpaired " eye " for special
description later on. We may here say that, whether
this attempt to explain the origin of the Crustacean eye
as a visual organ from the Annelidan eye succeeds or
not, our theory will not be affected, for there are
points in the anatomy of the eye of Apus, such as the
musculature and the space between the eye and the
integument for water, which are easily enough ex-
plained on our theory, and which would, we think, be
very difficult to explain on any other theory. If then
we ourselves fail to trace the rise of the elements of
the Crustacean eye, another, better fitted for the task,
will no doubt be more successful.

As already stated in the introductory chapter, our
original Annelid is supposed to have had (like the
Nereidae) two pairs of eyes on the prostomium, which
we have called simple eye-spots. Von Graber has
shown that these Chaetopodan eye-spots are by no
means simple structures, but are complicated visual
organs. This, however, does not make our task any
the more difficult, because we attribute the trans-
formation of the Annelidan into the Crustacean eye
chiefly to the thickening of the cuticle, which is one of
the Crustacean characteristics of the Nauplius before
the paired eyes are formed. The simple hypodermal
elements of the Annelidan eye have thus had to
develop, in each individual, not under Annelidan but
under Crustacean conditions, i.e., under a thickening
exoskeleton.

When we come to ask what are the most character-

90 THE APODID^E PART i

istic elements in the Crustacean eye, we find them to
be (i) the crystal cones, and (2) the retinulse, z>.,
definite groupings of a certain number of retinal and
pigment cells (see Fig. 23). In the first of these we
have a new structure, whose development must be
accounted for ; the second may safely be assumed to
be merely specialised hypodermal sensory (i.e., visual)
cells ; we have simply to account for the " retinula-
tion " of these cells, as Lankester calls the grouping
of them into retinulae.

We assume, then, that first of all the crystal cones
were but slight irregularities in the thickness and
refractiveness of the developing cuticle. Under
these irregularities, i.e., under those which either con-
centrated the light or otherwise favoured its passage,
the visual cells would naturally tend to group. We
say naturally, because it is clear that, under places
through which the light but feebly penetrated, the
visual cells would be rendered useless. In process of
time, certain definite irregularities of the cuticle would
be selected and further developed as lenses, &c., for
collecting the light. We find in the eye of Limulus
the particular form of cuticular development which
may have given rise to the crystal cones of Apus (cf.
Fig. 22 with Fig. 23). This fact is particularly
interesting because we have already seen that
Limulus has retained the original position of the brain
in the bent Annelid. And here we find the same
animal supplying a form of eye which shows clearly
a possible origin of the Crustacean crystal cones.
We have only to assume that such conical processes

SECT, vi THE SENSORY ORGANS 91

of the cuticle as we find in Limulus projecting inwards
became separated from the cuticle, and thereby,
naturally, surrounded by their secreting hypodermis
cells, and we have at once the Crustacean crystal
cones and cone-cells.1

In the eye of Limulus we further find visual sensory
cells forming groups or retinulae at the tips of the
cones, these latter having been gradually pushed

c

hy '"

r -

FIG. 22. — Section through the eye of Limulus (after Lankester). c, cuticle which
grows into conical papillae directed inwards, and pushing down the hypodermis
cells {hy) ; at the tips of the cones are found the retinulae imbedded in connective

down below the hypodermis into the subhypodermal
connective tissue. We find exactly the same in the
eye of Apus (see Fig. 23), where the crystal cones and
the hypodermis cells form the original hypodermis
layer, the retinulae having been pushed down even

1 Grenadier in his Sehorgane der Arthropoden states that the cones
in Limulus have nothing to do with the Crustacean crystal cones.
Our contention here is, however, that some such conicle projections
of the cuticle, not necessarily exactly similar to those in Limulus, may
easily be supposed to have produced the Crustacean crystal cones, by
being separated from their cuticle.

THE APODID^E

PART I

below the connective tissue layer. In this way we
get the double layer of cells composing the typical
Crustacean eye.

cc

FIG. 23. — Diagram of two single eyes of the paired eyes of Apus. hy, original
undifferentiated hypodermis cells secreting outer cuticular membrane. cc,
crystal cone cells = differentiated hypodermis cells secreting the cones (cr) ; cl,
layer of connective tissue fibres = subhypodermal connective tissue layer ; r,
retinulse, i.e., groups of sensory (retinal) cells (re), with their rhabdoms (rh),
and pigment cells C/1) ; these belong to the hypodermis, but are thrust down
below the subhypodermal connective tissue layer by the cones.

The great advantage of this separation of the cones
is not far to seek. The rounding off of the distal ends
1 See Appendix II. on the pigment in these cells.

SECT, vi THE SENSORY ORGANS 93

of these cones seems one of the simplest methods of
obtaining convex surfaces to act as lenses for the
reception of the light-rays from all directions. The
formation of corneal lenses over these cones is a
secondary and, we think, a much more complicated
specialisation.

Another possible advantage gained in the separa-
tion of the cones from the cuticle is the slight
possibility of movement which the separate omma-
tidia or single eyes would thereby acquire. It
appeared clear to us, during our study of the eye of
Apus, that the separate elements were capable of
slight movement, brought about, no doubt, by the
layer of connective tissue, which is only indicated
by faint lines in the diagram (Fig. 23), but which in
reality is very highly developed. The slight attach-
ment of the crystal cone cells to the cuticle would
not altogether prevent such small movements as we
suppose.

If the Crustacean eye is in this way to be referred
to the formation of the exoskeleton, it seems clear
that no special value can be attached to similarity of
eyes in establishing the relationships between animal
groups. The development of an exoskeleton is
common to the whole class of the Arthropoda, and
there is no reason why very different forms of cuticular
irregularity should not be utilised by the visual cells,
which would group themselves accordingly rn different
ways. We thus see no reason whatever for trying to
deduce the one form of Arthropodan eye from the
other, it being more probable that they are with a few

94 THE APODID^E PARTI

exceptions independent groupings of the sensory cells
under different forms of cuticular irregularities.

Returning to the eye of Apus, it is of special
interest to find that the eyes of all the Crustacea
which we assume to have descended from Apus may
be referred back to different groupings and modifica-
tions of the ommatidia, whose first development in
Apus we have endeavoured to describe.

The formation of the corneal facets above the
crystal cones may be due to a further utilisation
of irregularities in the thickness of the cuticle which
remains after the separation of the crystal cones. In
this way, we think, the gradual development of the
Crustacean eye may have gone hand in hand with
the thickening of the cuticle to form the exoskeleton
characteristic of the class. We have two highly
plastic elements, the hypodermis, with its scattered
sensory and pigment cells, and the thickening cuticle.
We cannot help thinking that it was the latter which,
coming between the sensory cells and the source of
stimulation, took the lead in the formation of the
different kinds of Arthropodan eye.

Before dismissing the subject of the development
of the Crustacean eye we feel that some apology is
necessary for treating it so shortly and so lightly.
We have not attempted to work through the enor-
mous literature on the structure and development of
the Crustacean eye.1 Our object here has been to

1 While these pages have been passing through the press we have
had occasion to read Watase's admirable paper on the " Morphology
of the Compound Eyes of Arthropods." It was especially interesting

SECT, vi THE SENSORY ORGANS 95

describe a possible development of the Annelidan
eye-spots into simple Arthropodan eyes, that being all
that we here need, in our endeavour to show that
Apus is a very primitive Crustacean, and at the same
time but a slightly modified Annelid. What we have
here written is a preliminary suggestion as to the
probable rise of the Crustacean eye. We hope in
another place, and in another connection, to discuss
it more fully, dealing especially with the sensory
elements and their physiological significance.

Some further morphological details relating to the
paired eyes of Apus fortunately admit of more satis-
factory deduction from the Annelida than does the
fine structure of the eyes themselves.

As to their position, we have two remarks to make :
I. We find them on the dorsal frontal surface,
whereas in the original Annelid they were on the
prostomium. It has already been assumed that, on
the fixation of the bent attitude of the five anterior
Annelidan segments, they gradually wandered round
on to the dorsal surface. There is no great difficulty
in this assumption, especially as we have seen, in our
investigation of the central nervous system, that the
position of the brain and the divided cesophageal
commissures indicate that such a wandering of the

to us to find that he had also selected the compound eye of Limulus as
the nearest type of the primitive Crustacean eye. We do not see what
is gained by his assumption of integumental pits. It is not easy to see
how the various stages in the development of these pits could have
functioned as visual organs.

96 THE APODID^: PART i

eyes has taken place. The length of the stalks of
the optic nerve, and the secondary drawing out of
the longitudinal commissures of the anterior ventral
ganglia, point to the same conclusion.

2. We find the eyes close together, i.e., about as
near to one another as they probably were on the
prostomium. This point is important, because it is
often assumed that the eyes of Apus have moved
together from the sides towards the middle line,
whereas, on the contrary, we hold that the eyes of
Apus have kept about the same distance apart as
they were on the prostomium of the Annelid
ancestor, and that it is the eyes of the higher
Crustacea which wander apart and take up positions
at the sides. The gradual reduction of the dorsal
shield, in the majority of the descendants of Apus,
facilitates the wandering to the sides. The case of
Limulus is particularly interesting. Here, as will be
described later, the eyes wandered from the first, not
forwards and upwards, but sideways and upwards, so
that the brain could not follow as in Apus, but, being
drawn in two opposite directions, remained where it
was, the extraordinary length of the optic nerves
showing clearly that the eyes must have wandered
considerably.

Almost more interesting, however, are the water-
sacs which spread out over the eyes of Apus,
between them and the integument. These have never,
we think, been described before, at least in detail,
and here deserve particular attention as lending
support to our theory of the migration of the eyes.

SECT. VI

THE SENSORY ORGANS

97

Fig. 24 is a diagram of these water-sacs. A small
pore, in the shape of a fine transverse slit (see
Figs. 24 and 69, p. 303) in front of the eyes, is, in
large specimens, visible to the naked eye. This leads

oe

FIG. 24. — Diagram to show the water-sacs over the eyes of Apus. s, water-sacs;
c, canal leading into the same ; p, pore ; e, paired eye ; b, brain ; ce, cesophagal
commissure ; og, optic ganglion ; m, eye-muscles. The eye is drawn in the
section ; in reality a median section passing through the pore passes between the
eyes ; e?, unpaired eye receiving a branch from the water-canal.

into a canal, which runs along the dorsal surface of
the unpaired "eye." Here it widens out con-
siderably, its upper and lower chitinous membranes
being in close contact, except in the median line

H

98 THE APODID^: PART I

above the unpaired " eye " (see Fig. 27, p. 105). At the
posterior end of the unpaired " eye " it gives off a
branch which runs into that organ, as will be described
later. The sacs then widen out over the eyes, as
shown in Fig. 24.

This water-layer probably facilitates the move-
ment of the eyes by the ring of muscles attached
round their rims. It is not improbable that the
sacs further serve as lenses, but this cannot be stated
with certainty. As for the mechanism by which the
water is drawn in and out, a contraction of the whole
ring of muscles at once would draw the water in,
while a general pressure of the body fluid under the
eyes, caused by muscular contractions in other parts
of the body, would expel it through the canal.

This whole structure has probably been developed
in the following way. The original head showed con-
strictions between the segments of which it was
composed, i.e., folds of the skin projecting inwards.
The eyes, in travelling backwards, would necessarily
have to pass by these folds. The first fold of all
would be that between the prostomium and the
first segment. It is clear that the eyes must either
be stopped in such a fold, or else carry it back writh
them. This latter is what we suppose took place at the
posterior edge of the prostomium. The eyes came
against the fold of the first segment, which generally
overlaps the prostomium when the body is at all
contracted. Under this the eyes would disappear.1

1 It is clear that while the bending round of the anterior segments
would so stretch the dorsal integument as to obliterate all such constric-

SECT, vi THE SENSORY ORGANS 99

It would be impossible for them to get further
than the posterior end of this fold unless they
dragged it after them, and thus, as we suppose, the
fold has travelled backwards with the eyes, the front
part gradually closing, till only a fine transverse slit
is left.

This derivation of the water-sacs receives some
support from the ring of muscles round the eyes,
which are clearly bands borrowed from the longi-
tudinal musculature. Their sinewy attachment joins
the large muscle which runs down from near the
eyes to be attached near the prostomium. A com-
parison of Fig. 24 with Figs. 12 and 13, p. 56, makes
the origin of the eye muscles very evident. As the
eyes, which are hypodermal structures, travelled
backwards, dragging an intersegmental fold back
with them, they naturally took along with them some
of the longitudinal muscle bands attached to that
fold. This accounts for the way in which a hypo-
dermal structure, such as the original Annelidan eye,
became an independent organ, movable by a special
and apparently highly developed system of muscles.

The development of the stalked eye from the eye
of Apus appears to us by no means such a simple
matter to understand. We are inclined to think that
it may have taken place in two different ways :
(i) By the gradual projection of the eye itself above
the surface of the body (we find such projections of
the eye stalks in many Trilobites). (2) By the
tions between them, this would not be the case with the fold between
the prostomium and the first segment. The labrum of Apus was
probably at first a movable organ.

H 2

TOO THE APODID^E PARTI

gradual diminution of the shield and head, and the
sinking in of the sides of the latter till the eyes,
with their cones of long muscle-bands (cf. Fig. 24, ;;/),
became movable lateral ridges running dorso-ventrally.
Their gradual complete articulation at their bases
would then easily follow. This method of trans-
forming the eyes of Apus into movable stalked eyes
is well exemplified in Branchipus stagnalis, which is
nearly related to Apus. The head region seems in
it to be reduced to a small base for carrying the
enormous second antenna and the stalked eyes.

THE UNPAIRED "EYE."

This organ, which is just visible as a dark spot be-
tween the paired eyes of Apus, is often called the
" rudimentary median eye of Apus." Closer study
of it, however, reveals that it is a highly developed
sensory organ with definite functions of its own. In
describing the water-sacs over the eye, we have
already had occasion to refer to it, and we there found
that through the canal which leads into these water-
sacs its interior is also in open communication with
the external world, that is, if the chitinous fold which
runs into it is really open at the end. It is better to
defer a discussion as to the use of this organ until after
an examination of its general structure and probable
origin.

Beyond the account of the middle eye of a Cope-
pod by Grenacher, this organ has received very little

SECT, vi THE SENSORY ORGANS 101

attention. It is generally referred to as the x-shaped
eye-spot which occurs throughout the Entomostraca,
but disappears during larval life in the Malacostraca.
It is, as a rule, so small, that investigation of its
finer structure is difficult. In Apus, however, the
unpaired eye is so large that its finer details are made
out with comparative ease. This fact is especially
important from our point of view, for if Apus is really
the (or a) primitive Crustacean, then all the unpaired
eyes throughout the whole class are in all probability
only modifications of that of Apus. Hence it is
necessary, for a comparative study of these organs, to
have some accurate knowledge of their original form.
We feel justified in assuming that this organ in Apus
is in its original form, not only because Apus has
retained so many primitive (i.e. Annelidan) character-
istics, but because, as will be described below, this
form gives us a clue as to the origin of the organ out
of an anterior pair of Annelidan eye-spots. In these
pages we must of course confine our attention exclu-
sively to the unpaired eye of Apus, describing in
order (i) its general structure ; (2) its probable origin ;
and (3) its present functions.

Structure. — Two groups of sensory cells, each form-
ing what is in this connection generally called a retina,
yield the two side walls of a cavity which is flat at the
top and rounded below. The top consists of the chiti-
nous fold, already described as forming the water-sac,
while the lower part hangs free in the body cavity.
Anteriorly, the cavity runs to a point along the water-
canal (sec Fig. 25) ; posteriorly, it ends in a blunt

102

THE APODID/E

PART I

point very nearly abutting on the brain. The posterior
wall of the cavity and the posterior half of its floor
are also composed of somewhat similar retinae. The

FIG. 25. — Longitudinal median section through the unpaired "eye," diagrammatic.
vr, ventral; dr, dorsal retina; /, tangle of pigment cells; s, water-space over
the eyes ; c, canal of the same, giving off a branch (Q) into the interior of the
organ ; 6, hrain.

sensory ends of all these retinal cells point inwards ;
the nerves from the outer ends of the cells unite
together to run towards the brain, forming from the
four retinae four nerve strands on which the un-

SECT. VI

THE SENSORY ORGANS

103

paired " eye " seems to stand on the brain, as on four
stalks, between the stalks of the optic ganglia. The
stalk of the ventral retina is distinguished by the pre-
sence of several enormous ganglion cells, apparently
the largest in the whole nervous system of Apus.

.dr

FIG. 26. — Lateral view of the unpaired "eye," diagrammatic, vr, ventral ; dr, dorsal ;
Ir, lateral retina ; b, brain ; S, water-space ; c , canal of the same ; /. tangle cf
pigment cells. Only the outer ends of the lateral retinal cells can be seen.

A cross section of the posterior end of the organ
looks to the naked eye like an X. The retinae, bul-
ging in somewhat towards the interior of the cavity,
give its lumen this form. (See Fig. 27, 3.) The
pigment cells which fill the cavity form a tangle of

104 THE APODID^: PARTI

pseudopodia without any apparent arrangement.
They contain the same minute olive green pigment
granules as the other pigment cells, which spread out
in great numbers over the subhypodermal connective
tissue layer throughout the body. This, however, is not
always the case ; some specimens had brown pigment
in large granules like those of the paired eyes. Some
of the pseudopodia penetrate a long way between the
retinal cells, and, at the surface of the organ where
there are no retinal cells, the pigment cells with their
long processes form together the external surface,
the whole structure having apparently no enclosing
membrane. The pigment cells, in fact, are nothing
more than a plexus of the ordinary pigment
cells which spread out irregularly throughout the
whole body, among the subhypodermal connective
tissue.

In a well-preserved specimen, in which the pigment
did not happen to be very dense, the cells were seen to
send down processes towards the inner sensory ends
of the retinal cells. As these processes were regularly
arranged, and free from pigment, it was difficult at first
to decide whether they belonged to the retinal or to
the pigment cells. We mistook them at first for large
cilia belonging to the retinal cells.

It will be seen from the drawings that the retinal
cells are not all of the same size. The lateral retinae
are composed of two distinct groups, an anterior group
of long narrow cells, and a posterior group of short
thicker cells. The ventral retina is composed solely
of large thick cells, and the dorsal of two groups of

SECT, vi THE SENSORY ORGANS

105

.S

--

• >

I.

106 THE APODIDvE PART i

similar large cells, the smaller group placed dorsally
and slightly anteriorly to the other.

As already described, a fine branch of the water
canal, on which this sensory body is suspended, runs
in towards the angle made by the dorsal and ventral
retinae. This fine canal is shown in Fig. 25 and in
sections 2 and 3, Fig. 27.

This structure of the median eye seems to be com-
mon to all the specimens of the different species of
the Apodidae examined by us. In series of sections
the organ is very likely to be displaced by the tearing
away of the chitinous tube. On this ground it would
require a much more extended study to ascertain
whether arrangements which sometimes appear to be
characteristic of the organ in the different species
are not really due to defects in the preservation of the
animals or in the preparation of the sections.

Enough has here been said as to the general struc-
ture to bear out what we maintain — viz., that the
unpaired eye is no rudimentary organ, but in reality
a highly developed sensory body playing a most
important part in the life of the animal.

Origin. — As to the probable origin of this organ,
everything points to its having been originally com-
posed of an anterior pair of eyes on the Annelidan
prostomium. The two posterior eyes formed the
paired eyes, the two anterior, which were nearer
together than the posterior pair, united to form the
unpaired eye. The evidence in favour of this origin
seems to us to be overwhelming.

When the paired eyes travelled into the end of the

SECT. VI

THE SENSORY ORGANS

107

fold as already described, the anterior pair followed
them, and also disappeared under the fold, but,
naturally, nearer the opening. While the paired eyes,
perhaps with the aid of the water lenses, remained
functional as eyes, the anterior pair of Annelidan eyes
seem to have been dragged out of their hypodermal

FIG. 28. — Diagram to show the origin of the unpaired eye out of two eye-spots, the
retinae being drawn down to form-the lateral retinae of that organ, p, pigment
cells. The pigment in the eyes is not drawn.

position, and to have been arranged so as together to
form the cavity already described. The hypodermis
cells can still be seen secreting the chitinous membrane
of the under side of the water-sac. The retinae now
forming the unpaired eye seem to be connected with
the original hypodermis only through fine connective
tissue and pigment strands, similar pigment cells also

ro8 THE APODIDvE PART I

crowding between the retinae. Fig. 28 is a diagram to
show the way in which we suppose the unpaired eye
to have been formed out of a pair of simple hypo-
dermal eyes. Further investigation must decide
whether the dorsal and ventral retinae and the larger
posterior cell groups of the lateral retinae are later
differentiations of the same two original eyes or new
developments. In the inner ends of the retinal cells,
i.e., in that part of the cells which points inwards,
irregular roundish or oval refractive bodies of different
sizes are found ; these are probably remains of rods or
rhabdomeres originally secreted by these retinal cells.

The whole structure of the organ, its apparently
loose connection with the hypodcrmis, the chitinous
fold which runs down as a branch from the water-canal
into its cavity, the occasional occurrence of brown
eye pigment instead of the olive green connective
tissue pigment, the nerves of its retinae running
separately into the brain, all tend to support the
above view of its origin (cf. also p. 169).

Further corroboration of this theory of the origin of
the unpaired eye from the anterior pair of Annelidan
eyes will be found in the section on Limulus. In that
animal the anterior Annelidan eyes remain as eyes, but
are reduced to ocelli, or eyes with one single large cuti-
cular lens. These ocelli first appear, according to
Packard, on tJie ventral surface^ and wander on to the
dorsal surface in the course of the later development. *
This astonishing fact receives its full explanation if we

1 We shall also find clear traces of a migration of the eyes in the
Crustacean Nauplius, § xi. Figs. 36 and 37, p. 158.

SECT, vi THE SENSORY ORGANS 109

admit the relationship between Limulus and Apus,
and deduce them both from the same bent Annelid, in
which the eyes wander from the prostomium, where
they are useless as eyes, to a position where they can
function as such. In Part II. we hope to show that
the derivation of Apus and that of Limulus from the
same bent Annelid stand or fall together.

Function. — As to the function of the unpaired eye,
we can perhaps with some certainty conclude that in
Apus at least it regulates the position of the body in
the water. Its structure out of four sensory retinae at
once suggests such a function, while further, the loose
tangle of pigment cells would constitute a body free to
be acted on by the earth's attraction. The organ is
perhaps rendered more perfect by the rounded arrange-
ment of the retinal cells, which thus present many
different surfaces to appreciate the movement of the
mass of pigment cells under the action of gravity.

It is difficult to ascertain for certain whether the
chitinous canal opens in the cavity of this sensory
organ or not. If it opens in the cavity to fill it with
water we should rather expect to find a more definite
membrane round the whole organ. If, on the other
hand, it does not open, it may be a structure for
the appreciation of changes of pressure, i.e., of
depth. The end of the tube as shown in Fig. 27
(2 and 3) is irregular, and in section seems as if it
might be a loose empty sac. As the outer pressure
increases, such a fine membranous sac would be the
first part of the body to feel it, and would commence to
swell. But such an appreciation of changes of pressure

no THE APODID/E PARTI

could also no doubt be equally well effected if the
canal were open, the increase of pressure leading
to a rush of water into the cavity. Further, whether
the canal is open at the end or closed, it would serve
well for enabling the animal to have a rapid perception
of changes of temperature in the water, the inlet of
cold water acting at once on the pigment cells.

We may perhaps find some confirmation of our
supposition that this sensory body fulfils various
functions, in the fact that there are at least two kinds
of retinal cells. The diagrams in Fig. 27 show not
only differences in the forms of the cells, but also
different groupings of the cells. The end of the canal
is shown close to the posterior groups of short thick
cells.

We have thus a comparatively simple but extremely
useful organ, probably adapted for the immediate
appreciation of the changes of depth and temperature
in the medium in which the animal lives, and further
for regulating its position in the water. The Apodidae
are, from all accounts, invariably excellent swimmers.
Keeping the ventral surface of the body uppermost,
they dive occasionally with great rapidity, rising
again to skim along just below the surface of the
water. Some organ to regulate such definite move-
ments is clearly necessary.

That this is at least one of the chief functions of the
organ is rendered probable by its early appearance
in the Nauplius larva. The powerful rowing limbs of
this free-swimming larva render some directive body
necessary ; hence the appearance of this organ along

SECT, vi THE SENSORY ORGANS in

with the rowing limbs, and long before the paired eyes
arc developed or needed. It is also worth noting that
the unpaired " eye " is especially characteristic of most
small free-living Crustacea such as the Ostracoda,
Cladocera, and non-parasitic Copepoda.1 In some of
these animals the organ probably combines rudi-
mentary visual with directive sensory functions, the
visual function being secondarily acquired, as there
can be little doubt that it has entirely ceased in
Apus.

This view of the function of the unpaired eye, by
explaining its early appearance in the larva, makes it
unnecessary to suppose that it is, as is generally
assumed, therefore phylogenetically older than the
paired eyes. On the other hand, its appearance in the
larva of all Crustacea rightly leads to the conclusion
that it was present in the original racial form of the class.
According to our theory, Apus being the ancestor of
the majority of the modern Crustacea, the unpaired
eye appeared for the first time as such in Apus.

A further and more exact study of this interesting
organ in Apus, and a comparison of it with the homo-
logous organs in other Crustaceans or Crustacean
larvae, is very desirable. It is impossible here to
follow up the matter further, as it would lead us too
far from the main subject of the book, which is an
endeavour to show how every single organ of Apus
admits of more or less easy derivation from similar
or dissimilar organs of a carnivorous Annelid.

1 Some of the differences between the unpaired eye of Calanella
described by Grenadier, and that of Apus will be referred to in § XV.

SECTION VII
THE ALIMENTARY CANAL

ALTHOUGH it is not, as a rule, possible to draw
any conclusions as to the relationship between animal
groups from the similarity of their alimentary canals,
yet the likeness between the digestive tract of the
Apodidae and that of the Annelida is so striking
that it must be admitted to be of some weight in
establishing the relationship which this book seeks to
prove. Allowing for the bend in the oesophagus,
the alimentary canal runs straight through the body
from end to end, and the mid-gut is lined by the
thread-like ciliated epithelium characteristic of that
of the Annelida.

The bending of the first five segments of our
Annelid, so that the mouth not only lies ventrally
but faces posteriorly, necessarily led to a bend in the
alimentary canal, so that, from the mouth, the
oesophagus would slope upwards and forwards. We
find that it has this position in the Apodidae, and
from the Apodidae it has been handed on to the

SECT. Vii THE ALIMENTARY CANAL 113

whole class of the Crustacea as one of their most
constant characteristics. It is not easy to imagine a
simpler or more likely explanation of this extra-
ordinary bend in the intestinal tract than that here
given ; and if it is true, its importance for the purposes
of classification is at once evident. We shall return
to this subject in the section on the relations of Apus
to Limulus and to the Trilobites.

The oesophagus itself corresponds with that of the
original Annelid, which was probably provided with
a protrusible pharynx. The loss of this proboscis
would naturally follow on the adoption of a browsing
manner of life, and the gradual adaptation of the para-
podia as instruments for pushing food into the mouth.
The oesophagus is very muscular, and is provided
with muscle bands radiating forwards into the fore-
head, and backwards into the sinewy mass already
described. These bands serve to dilate it, while its
powerful circular muscles close it ; when closed it is
thrown into folds. The dilators may perhaps be the
remains of the retractors and extensors of the
pharynx. The oesophagus is lined by a chitinous
intima and provided with setae which project upwards
so as to form a fish-trap apparatus. The paired glands
which open on the under lip close to the mouth (see
Fig. 29) will be described in the section (IX.) on the
excretory and other glands.

The oesophagus is in Apus comparatively simple,
but it is easy to see how a part of such a muscular
apparatus, with its chitinous intima folded, and thrown
into strong movement by every act of swallowing,

I

THE APODID^E

PART I

might become differentiated into a masticatory
stomach, such as we find in the higher Crustacea.
No such differentiation is, however, visible in Apus.
In Limulus we shall find the chitinous ridges used
for masticatory purposes in what is called the pro-
ventriculus, which is homologous with the masticatory
stomach of the higher Crustacea.

The oesophagus projects somewhat into the mid-

FlG. 29. — Diagram of the branched diverticula of the anterior end of the mid-gut (;«),
on the left without the glandular (hepatic) branched cocca, on the right a few of
the latter are drawn, a?, entrance to the oesophagus, on each side of which are
seen two long glands opening together in the middle line, assumed to be the
acicular glands of the vanished parapodia of the first antennal segment.

gut, which is a large sac with lateral diverticula — five
or six on each side ; these unite, in Apus, to enter
the mid-gut together (Fig. 29). These diverticula fill
up the large flat head widened by the ridge run-
ning round the frontal region, as already described.
Diverticula of the mid-gut are common among
Annelids, and serve to increase the digesting surface.
The diverticula of Apus are especially interesting

SECT, vii THE ALIMENTARY CANAL 115

as showing a perfect transition stage between the
simple digesting diverticula of the Annelids, and the
hepato-pancreatic glands of the higher Crustacea.
In Apus, particles of food are found in all the wider
parts of the diverticula which, like the rest of the
mid-gut, are lined with ciliated epithelium. Smaller
branching invaginations from these diverticula (see
Fig. 29) contain large glandular cells, which occur in
great numbers towards their tips. The secretion of
the glands is no doubt forwarded by the ciliated
epithelium, which is everywhere present when not
entirely displaced by the glandular cells. In the
preserved specimens this secretion formed brown
crystals.

In order to turn these digesting diverticula, pro-
vided with glandular cells at their branched distal
ends, into the lobate hepato-pancreas of the higher
Crustacea, we have only to imagine the glandular
cells increasing so as entirely to displace the ciliated
digesting epithelium, and the lumen of the diverticula
themselves narrowed to form glandular ducts.

In Astacus, where the mid-gut has almost en-
tirely disappeared, these diverticula are highly deve-
loped tassel-like hepato-pancreatic glands. Apus
supplies us with a perfect transition stage, showing
the origin of these livers out of the digesting diver-
ticula of the Annelidan mid-gut.

The epithelium of the mid-gut, like that of the
diverticula, is composed of minute thread-like cells
with nuclei near their basal ends. They stand on a
basal membrane, round which run at short intervals

I 2

ii6 THE APODID^: PARTI

fine circular bands of transversely striated muscles.
We find here a striking resemblance with the mid-gut
of many carnivorous Annelids.

The mid-gut passes gradually into the hind-gut ; it
is very difficult to fix upon the exact place where the
chitinous intima of the latter commences. In passing
from one to the other, the muscular layer is more
and more developed ; the epithelium gradually
changes, becoming more and more a glandular
epithelium, with large round glandular cells arranged
in great numbers, and pouring their contents through
pores l in the chitinous intima into the hind-gut. From
the position of these glands we are fairly safe in con-
cluding that they are excretory.

In the anal segment the rectum is attached to the
body wall by radiating muscles, which act as dilators,
while the strong circular muscles keep it closed. As
in the oesophagus, the wall of the rectum is thrown
into folds, which run longitudinally. The anus is
situated at the extreme end of the body under the
caudal plate, where such a plate is present, and
between the caudal cirri or cercopoda.

1 The actual existence of these pores we have not, however, been able
to demonstrate.

SECTION VIII
THE CIRCULATORY SYSTEM

THE actual blood-vessels in Apus are limited to the
long dorsal vessel or heart. Although, among the
Annelida, circulatory systems are found of many
different grades of development, it is not necessary
to suppose that our original Crustacean-Annelid had
a very simple blood vascular system, in order to
account for the above fact. It seems to us probable
that the development of an exoskeleton, which holds
the organs in their places, and protects the inner
parts generally from being squeezed together, renders
special blood-vessels more or less unnecessary, the
blood being able to bathe all the organs of the body
without difficulty. On the other hand, this is not the
case in a soft-skinned strongly contractile body, such
for instance as that of the leech, where it is necessary
-to carry special blood streams between the organs
which are liable to be crowded together. If then our
original Crustacean-Annelid possessed a highly deve-
loped blood vascular system, it would naturally, with

ii8 THE APODID^: PARTI

the development of the exoskelcton, give place to a
simple lacunar system as sufficient for the needs of
the body, the only vessel retained being the contrac-
tile dorsal heart, necessary for propelling the blood
through this lacunar system. It may be objected
that the trunk of Apus under the shield has a soft
skin, and nevertheless the blood system in this part is
entirely lacunar. The trunk is, however, not very con-
tractile, and although the blood flows through lacunse,
there are definite methods of propelling it through
the intestinal sinus, which will be mentioned later on.

It is clear, then, that the absence of nearly all true
blood-vessels from the circulatory system of Apus
makes any comparison of it with that of an Annelid
impossible. Apart, however, from the dorsal vessel
or heart, which is generally recognised as an organ
derived from Annelidan ancestors, there is, in Apus,
a slight trace of" a pair of typical Annelidan vessels ;
these are the short vessels which supply the shell
glands. (See Fig. 30, p. 125.) They branch off
from the dorsal vessel on each side, and descend
towards the dorsal parapodia of the second pair
of maxillae, to dip under the shell gland. They
are necessary for conducting the blood into the
shield in which the coils of the gland lie. They may
be homologous with a pair of lateral arteries from the
dorsal vessel of the fifth segment, such as typically
supply the parapodia and sides of the body in a
carnivorous Annelid.

Attention must be drawn to the tapering away of
the heart to a point towards the posterior end of the

SECT, viii THE CIRCULATORY SYSTEM 119

body. This is in keeping with our general explanation
of the morphology of the body of Apus. As we go
from before backwards, the organs are less and less
developed, the limbs are more and more rudimentary,
and the musculature less and less specialised ; the
nervous system ceases where the rudimentary limbs
cease, and the genital organs gradually diminish in
size and development. The heart is no exception ; it
tapers away in about the tenth or eleventh segment
into a point, not being developed in the more larval
segments which come behind.

We may also perhaps mention that the blood in
many, if not in all Apodidae is coloured red, as is the
case in many Annelids. The direction of the blood
through the body is the same as in the Annelids —
dorsally from behind forwards, ventrally from ^before
backwards. On its way back through the intestinal
sinus, which will be presently described, it streams out
ventrally through fenestrae in the membrane which
forms the sinus. (See Fig. 14, ;;/, p. 59.) It is thus
enabled to stream over the ventral cord, and then
outwards on each side along the ventral side of each
limb. At the end of the limb it turns round to run
back along its dorsal edge and thus passes through
the gills. It passes up thence through the lateral
dermo-muscular sinus of each segment into the peri-
cardial sinus, and thence through the ostia into the
heart.1

We have now to try and trace the origin of the

1 As this account differs from that of Zaddach, a fuller description
with illustrations will be given in Appendix III. p. 296.

120 THE APODID^E PART I

lacunar system of Apus more in detail. Its main
features are very simple, and in this respect it shows
a primitive character. In the main it may be said to
consist of but one membrane, forming a tube which
runs from the anterior end of the first trunk segment
(whore it is attached all round to the body wall) to
the end of the body. This membranous tube sur-
rounds the intestine and genital glands, while between
it and the body wall lie the heart, the ventral cord,
and all the musculature, except the dorso-ventral
bands which run between the intestine and the genital
glands. (See Fig. 14, p. 59 ; cf. also Fig. 66, p. 297.)
The membrane is attached to the body wall, at least
in the first eleven trunk segments, by segmental dis-
sepiments, which correspond with the segmental con-
strictions of the body. These dissepiments extend
dorsally to the points of attachment of the dorso-
ventral muscles, which raise up the membrane in
conical folds. Between these clorso-ventral muscle
rows the membrane hangs free of the dorsal body wall,
thus forming the cardial sinus, in which the heart is
expanded by an arrangement of connective tissue
fibres.

In trying to trace the origin of this membrane and
these dissepiments (see Fig. 67, s, p. 298) from the
internal organs of the original Annelid, we naturally
begin with the latter, as reminding us at once of the
Annelidan septa. Are they the remains of such
septa ? The answer depends on the interpretation we
give to the membrane forming the intestinal sinus.

Glancing at the membrane then as a whole, as a

SECT, viii THE CIRCULATORY SYSTEM 121

tube running through the whole trunk outside of the
dermal musculature, and containing the intestine and
the genital glands, its origin is at once suggested to
us. It appears to be the parietal layer of the coelom
epithelium of the original Annelid, loosened from the
body wall except at certain definite points, viz.,
where it is in contact with the ventral muscle bands,
and laterally along the segmental constrictions, where
it remains attached by means of the septa above men-
tioned, each of which extends dorsally as far as the
point of attachment of the dorso-ventral muscle
bands. (See Figs. 66, p. 297, and 67, p. 298.)

In this way we should at once get just such an
intestinal sinus as we find in Apus, viz., a membra-
nous tube lying just inside of the dermal musculature,
the transverse dorso-ventral muscles being almost the
only muscles found within the tube. Indeed the rela-
tion of the membrane to these muscle bands seems to
support this view, for where these are attached to the
dorsal surface, the membrane itself is raised up into
conical folds in the manner illustrated in the diagram
(Figs. 14, p. 59, and 66,/). This certainly looks as if
the membrane had, as it were, fallen away from the
integument. If this view is correct, the intestinal
sinus corresponds with the body cavity of the original
Annelid, and the dermo-muscular sinus of each seg-
ment is a new formation caused by the loosening of
the epithelium from the body wall.

The dissepiments themselves may be folds of this
membrane grown together. If so, these partial dis-
sepiments must have been secondarily acquired, after

122 THE APODID^: PARTI

the original Annelidan dissepiments had disappeared.
There is no great difficulty nor, indeed, improbabi-
lity in such a supposition. We may indeed find in
these dissepimental folds, attaching the membrane
to the body wall, traces of the former presence of
the old Annelidan septa, which may have originally
run in between the folds just as the transverse dorso-
ventral muscles run up into the conical folds of the
membrane, as already described and marked / in the
figures.

If this latter view is correct, we then explain the
origin of the lacunar system of Apus by the loosening
of the ccelom epithelium from the heart and from the
dorsal surface in the immediate neighbourhood of the
heart so as to form the longitudinal cardial sinus.
On each side of this sinus laterally it remains attached
to the intersegmental folds, being loosened, however,
from the segmental walls, so as to form the lateral
segmental dermo-muscular sinuses, which conduct the
blood from the gills to the cardial sinus.

The separate stages by which this lacunar system
took the place of a blood vascular system, are not
difficult to imagine. The first step in the transforma-
tion would be the gradual degeneration of the separate
blood-vessels and the consequent mingling of the
blood with the body fluid or lymph. The disappear-
ance of the vessels supplying the dermal musculature,
and the general diffusion of haemolymph between
the integument and the ccelom epithelium might
very easily lead to the loosening of the latter from
the integumental musculature, excepting along

SECT. Yin THE CIRCULATORY SYSTEM 123

the intersegmental folds, across which the dermal
blood-vessels of the original Annelid did not run (?).
These two modifications, (i) the degeneration of the
blood-vessels, and (2) the loosening of the peritoneum
from the body walls, are all that is needed to produce
the lacunar system of Apus from the blood vascular
system of an Annelid.

The contrast between the circulatory systems of
the Annelida and of the Apodidae does not therefore
stand in the way of the acceptance of our theory of
their close relationship. Apart from the well-known
fact that, among the Invertebrata at least, blood
vascular systems have little morphological value, we
have here shown how simply the lacunar system of
Apus can be deduced from the blood vascular system
of an Annelid.

We reserve any further discussion of the ccelom
epithelium, a part of which has been here used up in
the formation of the lacunar system, till we come to
the genital glands, when we shall again see what an
important part it has played in transforming the
Annelid into the Crustacean. (See further p. 169 and
Appendix III.)

SECTION IX
EXCRETORY AND OTHER GLANDS

The Shell gland. — THE most conspicuous glands
of the Apodidae, the long coils of which are seen in
the shield, one on each side of the middle line, are
known as the shell glands. These are generally
homologised with the Annelidan nephridia.

From Figs. I and 2 it will be seen that we have
assumed that the acicular gland of the dorsal para-
podium of the fifth segment became excretory, and
grew into the growing shell fold, thus forming the
shell gland. The position of its external opening, at
the tip of the dorsal branch of the second maxilla,
agrees exactly with this supposition. The distal end
of it is a chitin-lined sac running through the shaft
of this limb.1 The proximal end of the gland has
grown into the shield, and is of great length, being
folded six times upon itself. That the inner part of
the gland has grown upwards, and the limb bent
downwards, can be seen by the course of the duct of

1 See Appendix IV. for the finer structure of the gland.

SECT, ix EXCRETORY AND OTHER GLANDS

125

the gland which, on leaving the shell fold, makes a
sudden sharp bend downwards. We thus consider
this shell gland as a striking link between our Annelid
and Apus. The mesoblastic origin of the urinary
canal may mean that this section of the gland is a

FIG. 30.— Diagram of the shell gland, k, the heart
blood is distributed to the gland through

:he aorta cephalica. The
a special vessel on each side, ts,

terminal saccule ; uc, urinary canal ; b, chitin-lined bladder in the shaft of the
dorsal parapodium of the 2nd maxilla (»i), homologous with the original
setiparous gland of the Annelid.

new formation, — the bladder alone representing the
acicular gland.

This homology of the Crustacean shell gland with
the acicular gland of the fifth parapodium of the
Annelid, naturally leads us to ask whether the acicular
gland of any other of the head limbs has been pre-

126 THE APODID^E PART i

served. The mandibles and first maxillae have lost
their dorsal parapodia almost entirely, and with them
all traces of setiparous glands. The antennae, how-
ever, are in this respect especially interesting. We
have homologised them with the sensory cirri of the
vanished or rudimentary dorsal parapodia of the first
two segments. The acicular glands of both, however,
have apparently been preserved in the Crustacea.

The Acicular gland of 'the first segment. — There is
a pair of glands (salivary ?) opening near the entrance
of the oesophagus (Fig. 29, p. 114). We think
that th£se belong almost certainly to the first
antennae ; 1 they open together in a small transverse

1 We were at first inclined to think that these "salivary " glands were
the acicular glands of the parapodia of the second antennae, there being
no antennal glands in Apus. Their opening in the mouth could then
be explained as follows : — When the Annelid first took to browsing, its
ventral parapodia near the mouth would not as yet have developed
teeth. The acicula of the antennal parapodia might then have been
used as piercers and killers of prey. They would thus be turned inwards
towards the opening of the mouth, where their glands might persist as
salivary glands after the development of the ventral parapodia of the
third and fourth segments into jaws and maxillae had rendered the
acicula useless as teeth. It seemed to us more likely that the acicula
of the parapodia of the second segment would be so used, than that the
acicula of the vanished parapodia of the first segment should redevelop
for that purpose. We have to choose then between the following :

(1) These salivary glands are the acicular glands of the parapodia of
the second antennae, the acicula themselves having once functioned as
teeth ; this homologises them with the typical antennal glands of the
Crustacea.

(2) They are the acicular glands of the vanished parapodia of the first
antennae which had redeveloped their acicula as teeth.

(3) They are the acicular glands of the vanished parapodia of the first
antennae which, as glands, need never have disappeared. While we
think the oral position of the opening of the glands is better explained

SECT, ix EXCRETORY AND OTHER GLANDS 127

fissure on the inner side of the under lip, almost
within the mouth. Their whole structure indicates
that they are acicular glands. The ducts are long
chitinous tubes which lead to a chitin-lined sac with
a very fine epithelium, their proximal ends being
attached to the body wall by muscle bands just as
are the setiparous sacs in the Annelids. The figure
shows the course of these long glands. What the
exact function of the two glands, opening together in
the mouth, may be, it is impossible to say. In all
preparations they are found to be strongly contracted,
so that the chitin-lined lumen is to be seen only with
difficulty.

These " salivary " glands, developed out of acicular
glands, are especially interesting as compared with the
salivary glands of Peripatus, which have been shown I
to be transformed nephridia. In both cases the •
openings of the glands have united in the middle line.
In both cases we have to assume that, the acicula or
the secretions of their glands on the one hand, the
excretion from the nephridia on the other, assisted the
jaws in their alimental functions as the first step
towards their differentiation into salivary glands.

The Acicular gland of the antennal parapodia or
Antennal gland. — In Apus we could find no certain
trace of an antennal gland at the base of the second
antennae ; a slight indentation on the basal swelling
seemed, however, to indicate that there had been

by the early use of the acicula as teeth, which would make our choice
fall between I and 2, we think the last view is the most probable,
although we do not reject the second alternative.

128 THE APODID^E PART I

an opening. Its redevelopment in the higher Crus-
tacea is paralleled by the redevelopment of the
dorsal parapodia of the mandibles, although the
latter had disappeared in Apus. It is a well-known
principle that organs which have disappeared may
reappear in the descendants of those who have learnt
to dispense with them.

The absence of the antennal gland in Apus is
perhaps to be explained by the enormous size of the
shell gland. In one specimen of L. Spitzbergensis,
1 1 mm. long to the tip of the caudal plate, the coils of
the shell gland on each side measured over 25 mm.
Such an enormous gland would no doubt be able to
undertake the greater part of the excretion of the
body.

We have thus, in the Crustacea, three setiparous
glands preserved in the head : the salivary (?) gland
of the first antennae (in Apus at least), the antennal
gland, and the shell gland of the second maxillae.

The antennal glands as well as the shell glands are
generally homologised with the Anneliclan nephridia.
From the foregoing account of the origin of these
glands we repeat the following points, which must
render such a homology improbable.

(i) The position of the external opening is on
the dorsal parapodium, — an unlikely place for the
opening of a nephridium, but, on the other hand,
quite a proper place for the opening of an acicular
gland.

(2) The structure of the glands as we find them (in
Apus at least) is exactly that of setiparous glands.

SECT, ix EXCRETORY AND OTHER GLANDS 129

A long chitin-lined duct opens into a similarly lined
vesicle.

(3) In the "salivary" gland the chitinous sac ends
blindly, the end being fastened by muscle bands to
the body wall, exactly as is a setiparous gland of the
Annulata. In the shell gland, however, the sac or
bladder is continued into a long coiled urinary canal.1
The position of this urinary canal in the dorsal fold,
and the finer structure of its walls, seem to indicate
that at least this part of the gland is a new formation.
It in no sense reminds one of an Annelidan nephri-
dium.

(4) These arguments are especially strong if the
rest of our argument holds good, viz., that Apus is
but a slightly transformed Annelid, or, indeed, if
we only claim what is often admitted, that the
Phyllopods stand nearest the racial form of the Crus-
taceans. If even this latter alone is the case, the
shell gland of Apus, if a true nephridium, should
show more likeness to a nephridium than do the
shell glands of the higher Crustacea, which have
departed further from the Annelidan type. \Ye
should expect the shell gland in Apus to be a
transition form between the Annelidan nephridium
and the Crustacean shell gland, just as we found the
" liver " of Apus to be a true transition form between
an ordinary digesting diverticulum such as is common
among the Annelida, and the purely glandular hepato-

1 Grobben says that the whole canal in the antennal gland of Mysis is
lined with a chitinous cuticle. In Apus, however, the intima ceases
with the bladder.

K

130 THE APODID^: PART I

pancreas of the higher Crustacea. But this is
certainly not the case. Neither in position nor in
structure do the glands remind one of Annulatan
nephridia, but, on the other hand, they correspond in
a most remarkable manner with the acicular glands
of the Annelidan parapodia.

(5) We further assume that the habit of browsing
of the bent Annelid was originally acquired by the
adult animals, in which the nephridia in the anterior
segments have generally disappeared in the course
of development ; so that Apus, which represents
such a browsing Annelid, would probably have no
nephridia in the anterior or head segments. In the
trunk segments, on the other hand, we shall find
abundant traces of nephridia.

These considerations, which, taken singly, do not
possess much weight, taken all together make the
nephridial origin of these glands very improbable
compared with that which we here attribute to them.

Sctiparous glands on the trunk segments are hardly
to be expected ; the dorsal parapodia are developed
into complicated limbs covered with setae, and the
ventral parapodia are also thickly beset with setae of
different kinds. We have succeeded, however, in
finding two such glands on the same limb in one
specimen of Apus cancriformis (see Fig. 31), We
could find no similar glands on the gnathobase or
ventral parapodium of the corresponding limb of the
same segment, and only on one other limb. Perhaps
further search would reveal more, but it is certain
that these glands occur quite irregularly. We arc

SECT, ix EXCRETORY AND OTHER GLANDS 131

inclined to consider them as occasional abnormal
returns to the Annelidan method of developing the
setae (see pp. 87, 88). When such an abnormal
setiparous sac does occur, it would in all probability
be utilised for excretory purposes. Chitin itself is
probably an excretory product, utilised for protective
purposes.

FIG. 31. — Part of a section through the gnathobase of Apus mentioned in the text
(p. 130), showing an abnormal reappearance of a setiparous gland containing
a brown secretion. «, the nerves to the hairs ; those to the feathered hairs send
a fibre into each barb and possess small groups of ganglion cells.

This irregular appearance in Apus of glands so
obviously homologous with the setiparous glands of
the Annelida, as an occasional abnormal return to a
former method of developing setae, establishes beyond
all contradiction the usual homology of the leg or
coxal glands of the Crustacea with the setiparous
glands of the Annelida.

K 2

132 THE APODW& PART I

Having so far considered the typical Crustacean
glands, the antennal, shell, and leg glands, and
homologised them with setiparous glands of the
original Annelid, it is necessary, in order to establish
the close relationship which we maintain exists
between Apus and the Annelida, to discuss the
typical excretory organs of the latter (i.e., the
nephridia), and to endeavour to discover their fate
during the transformation of the Annelid into the
Crustacean.

Professor Haeckel, in the last edition of his Natural
History of Creation^ characterises the Crustacea as
segmented animals without nephridia, and the worms
as segmented animals with segmcntal organs or
nephridia, the presence or absence of these latter
being the chief characteristic difference. The stress
here laid by so distinguished a zoologist upon the
nephridia as a class characteristic renders it doubly
necessary either to find the nephridia in Apus — our
primitive Crustacean — or to give some probable
explanation of their absence. Although no expla-
nation of the absence of nephridia was immediately
evident, we were convinced that it would some day
be found. We would not allow that the difficulty
of finding a set of organs in Apus to homologisc
with another set in the Annelida — though no doubt
serious — could destroy the value of the mass of
evidence already obtained as to the relation of
Apus to the Annelids. This reasoning is further
especially applicable in the case of the Annclidan

1 Ed. viii. 1891, p. 570.

SECT, ix EXCRETORY AND OTHER GLANDS 133

nephridia, whose arrangement in the Annelids them-
selves is always very variable.

It at first appeared possible that the absence of
nephridia in Apus could be explained by assuming that
in the original Crustacean-Annelid they were developed
more in the posterior segments (which is in fact often
the case), and that these segments do not attain deve-
lopment in Apus, the enormous shell gland sufficing
for the removal of waste products from the blood.
The weakness of this argument is at once obvious.
It is only when all the segments are fairly well
developed that the permanent nephridia are limited
to the posterior segments. Nephridia or their rudi-
ments are, as a rule, to be found at one time or
another in the course of development in all the seg-
ments. As the posterior segments attain develop-
ment, the nephridia in the anterior segments often
disappear. Nephridia ought therefore certainly to be
found in the developed segments of the trunk of
Apus, and rudiments of nephridia in the larval
segments of which the posterior part of the
trunk of Apus is composed. Fortunately, we
are not driven to take refuge in such a doubtful
explanation.

Knowing, on the one hand, that there are no true
nephridia in Apus, and on the other that in the carni-
vorous Annelids the nephridia are often the ducts for
the transmission of the sexual products, we naturally
tried to overcome the difficulty by the aid of the
genital glands. The study of these glands soon
yielded the desired clue. We have then here some-

134 THE APODID^: PARTI

what to anticipate a description of the genital
glands.

These glands are segmental tubes running dorso-
ventrally on each side of the intestine, inside the
intestinal blood sinus. They are separated from the
intestine only by the incomplete longitudinal dissepi-
ments formed by the dorso-ventral muscle bands
described above. The genital tubes commence in the
first trunk segment, but become gradually shorter and
shorter till they are quite rudimentary in the larval
segments of the abdomen ; they cease to be deve-
loped at all some distance before reaching the last
limb-bearing segment. These segmental tubes are
branched at each end. At the tips of the branches
eggs develop which are found projecting, not into
the tube, but into the body cavity, as will be more
minutely described in the next section ; it is, how-
ever, important for our argument to mention the fact
here.

All the segmental genital tubes on each side arc
connected together by a longitudinal canal which runs
through them all, and acts both as oviduct and shell-
or rather shell-secreting gland (see Fig. 32), so that
eggs coming dorsally and ventrally from the ends of
the branches meet in the middle, and then travel
along the longitudinal canal to near the middle of
its course, where a descending canal leads to the
exterior.

The question is, Can these organs reveal anything
about the vanished nephridia ? It is obvious that
they are not themselves the nephridia ; they are

SECT, ix EXCRETORY AND OTHER GLANDS 135

simply tubes formed out of the germ-bearing epithe-
lium. When we turn to the carnivorous Annelids,
we find that the germ-bearing epithelium is simply
the ccelom epithelium which covers all the internal
organs including the nephridia ; the eggs project from
this epithelium into the body cavity, and, falling off,
ripen in the body fluid (see Fig. II, p. 54) to find
their way out through the nephridia. When we com-
pare this process with what takes place in Apus, we
find in the latter an epithelium from which the eggs pro-
ject into the body cavity. (See Fig. 33, p. 144.) Is not
this epithelium homologous with the Annelidan ccelom
epithelium ? Instead, however, of dropping off into
the body cavity, the eggs are drawn back through the
epithelium and find their way out through the canals
formed by this ccelom epithelium. Are not these
canals, then, in some way, the homologues of the
Annelidan nephridia ? To the first of the above
questions we give an affirmative, to the second a
negative answer ; but we arrive at the conclusion that
the germ epithelium is the original ccelom epithelium
which covered tJie . nephridia, and that the canals
which it now forms once contained the nephridial
canals through which the eggs found their way to the
exterior. In course of time, the nephridial canals
ceased to have any excretory function owing to the
sufficiency of the shell gland, and disappeared, leaving
only their coverings of ccelom epithelium, which, in
proportion as the canals degenerated, itself developed
into the pronounced epithelium of the genital glands.
The development of the longitudinal canal also out of

I36 THE APODID^E PART I

the ccelom epithelium presents no difficulty, as this
epithelium, on the degeneration of the Annelidan septa,
would naturally form such a continuous membrane
through all the segments of the body. The dis-
appearance of all the ncphridial apertures except the
one between the tenth and eleventh segments is a
further very natural specialisation.

In summing up the arguments here used in favour
of this account of the disappearance of the nephridia
in Apus, we have to notice the following points :—

(1) The eggs which develop out of the epithelium
of the genital glands project, as in the Annelida, into
the body cavity and not into the cavity of the genital
glands themselves, as one would naturally have ex-
pected. This epithelium, then, is a part of the original
ccelom epithelium of the Annelid.

(2) The eggs pass again through the epithelium,
and travel down the canal formed by it ; this canal has
therefore probably taken the place of the nephridial
canal which it once covered as ccelom epithelium.

(3) We have to call attention not only to the
segmental arrangement of the organs, but to the fact
that there is a pair in each segment except in the
most rudimentary ; both of these facts agree with
what we know of the typical development of nephri-
dia in each segment of the Annelida.

(4) The position of these genital organs be-
tween the dorso-ventral muscle dissepiments and the
body wall agrees exactly with that of the nephridia
of the carnivorous Annelids, which lie in the lateral
chambers of the body (see Fig. n, p. 54).

SECT, ix EXCRETORY AND OTHER GLANDS 137

(5) The position of the only aperture which re-
mains agrees well with the position of the opening
of the Annelidan nephridium, i.e., it lies laterally on
the ventral surface. The fact that, in Apus, it comes
between two consecutive limbs is due to the bending
round of the parapodial limbs towards the ventral
middle line as already described (see § on Append-
ages).

(6) The genital aperture does not always remain
in the same segment in the Crustacea ; it differs even
in the males and females of the same species.

(7) Lastly, we have to add the fact that the genital
ducts have been generally recognised as homologous
with Annelidan nephridia.

\Yc thus believe that though the nephridia are ab-
sent in Apus, we have found sufficient traces of their
having been once present in the typical manner, one
pair in each segment, functioning, as they do typically
in Annelids, as ducts for the transmission of the sexual
products. The great development of the shell gland
rendering the excretory functions unnecessary, there
remained only the secondary function of conduct-
ing the sexual products out of the body. As this
could be done equally well by simple tubes formed
out of the covering of ccelom epithelium, these latter
alone have been retained, preserving, however, the seg-
mental arrangement and the position of the nephridia
which they had at one time clothed.

These considerations seem sufficient not only to
remove the difficulty caused by the absence of ne-
phridia in Apus, but even to strengthen the evidence

138 THE APODID^ PART

in support of our main argument ; they not only
remove a difficulty, but bear positive testimony to the
truth of our theory. Those who, however, may think
this view of the disappearance of the Annelidan nc-
phridia (with the exception of their peritoneal cover-
ings) far-fetched, should remember that the weight of
all the positive evidence brought forward as to the
relationship of Apus and the Annelida is only really
diminished if we cannot show that a difficulty is sur-
mountable. It is by no means necessary for our
argument cither to remove all difficulties so long as
they are not positive contradictions, or to state exactly
M how such and such a transformation came about, but
i only to show that such transformations are not incon-
ceivable. We believe, however, that in this case we
have not only shown this, but more, viz. that the pro-
cess of the disappearance of the nephridia was what
we have described.

We have now dealt with the principal glands of the
Crustacea and of the Annelida. We have deduced
the Crustacean glands from the Annelidan setiparous
glands, and followed the Annelidan nephridia in their
transformation into Crustacean genital glands.

Of the typical dermal glands of the Annelida we
have found no trace in Apus (except in the dorsal
organ, see below and Appendix IV). The hypodermis
is very thin, and seems to be entirely taken up in
secreting the cuticle in its gradual transformation
into an exoskeleton.

There are very numerous glandular cells in the
hind-gut, which have already been mentioned in

SECT, ix EXCRETORY AND OTHER GLANDS 139

§ vii., where we stated that from their position
they were almost certainly excretory.

The glands at the tips of the diverticula of the mid-
gut were also mentioned in the same section, and
were interesting as forming with the diverticula a
striking transition between the digesting diverticula
of the Annelids and the hepato-pancreas of the higher
Crustacea.

The white oval spot, or dorsal organ, behind
the eyes in Apus we at first thought to be the
remains of an Annelidan frontal cirrus such as that
shown in Fig. i. It appears to be an island of dermal
glandular cells, the last remains of the Annelidan
dermal glands, which the thickening exoskeleton
probably rendered impracticable. This organ will
be described in detail in Appendix IV. It offers
indirect support to our theory of the Annelidan origin
of Apus.

SECTION X

REPRODUCTION

THE carnivorous Annelids arc mostly sexually
separate. The same was originally the case with
the Apodidae, which arc now, however, mostly
hermaphrodite (see Appendix V.). Males, generally
smaller than the females, have been found at intervals
in the best known species, and these seem to suffice
for occasional cross fertilisation.

The sexual elements of the Annelids frequently
develop out of the ccelom epithelium (see Fig. n, p.
54), and then, falling off, ripen in the body fluid.
They arc discharged through the nephridia, which
may or may not be specially modified into sexual
ducts.

In the section on the excretory glands, we have
already briefly described the sexual glands in the
Apodidae. We have here, then, only to describe the
process of formation of the sexual products some-
what more minutely. We repeat, however, for the sake
of clearness what was said above. The genital glands

SECT, x REPRODUCTION 141

are segmental tubes connected by a longitudinal canal
which acts as a common duct for all the glands.
These latter are branched dorsally and ventrally.
The branching is very rich in A. cancriformis and L.
productus, and in L. glacialis more so than in L.
Spitzbergensis. We failed to find that the branches of
the genital glands in one segment anastomosed with
those in others, so as to form the network figured by
Zaddach ; but we see no reason why such anasto-
moses may not sometimes take place.

FIG. 32. — Diagram of a somewhat simplified genital tube of L. Spitzbergensis or
glacialis, the dorso-ventral segmental tubes only slightly branched, a, aperture
between the toth and nth segments ; /, the rudimentary part, i.e. the part
which lies in the posterior rudimentary segments, in which in this species the
epithelium breaks up into sperm cells filling up the whole lumen of the tube.

Although there is no apparent difference between
the epithelium of the longitudinal canal and that
of the sexual glands, it is useful to consider the
two apart. They are both, as we have seen, formed
out of the peritoneal epithelium, and besides having
the same origin, they have the same function, viz., to
secrete the slime which hardens in the brood pouch to
form a covering for the eggs.

It is of considerable importance for us to note first
of all that the sexual glands are segmentally arranged
in Apus. This, as far as we know, is not the case in

142 THE APODIDyE PART i

any other Crustacean, and is in itself evidence of
the primitive or Annelidan character of Apus. The
gradual simplification of the glands from before back-
wards towards the less developed segments is also
significant. The many-segmented ancestors of Apus
developed sexual products in every or nearly every
segment.

In the section on excretion we have already traced
back the epithelium forming the sexual glands to the
ccelom epithelium of the original Annelid, and to that
special part of the epithelium which covered the
nephridia. The nephridial tubes themselves have
entirely disappeared, having probably been rendered
useless by the great size and physiological efficiency
of the shell glands. Their peritoneal coverings, how-
ever, have remained as the sexual glands. The eggs
develop out of this epithelium, not projecting into the
lumen of the gland, but outwards, so that they bulge
out into the body cavity. This agrees with what
takes place in many Polychaetan Annelids ; the eggs
develop out of the peritoneum, and apparently out of
any part of the same, drop off into the body cavity,
and are emptied out through the nephridia. In Apus
slight changes have taken place : the eggs develop
towards the body cavity out of the peritoneal covering
of the vanished nephridia : they do not, however, drop
off, but pass through the epithelium again, to pass out
through the tube formed by this epithelium, just as
they at one time no doubt passed out through the
nephridial tubes. When we consider the great size
of the eggs owing to the accumulation of the yolk, it

SECT, x REPRODUCTION H3

is apparent that the vanishing of these nephridial
tubes could be nothing but pure gain ; the simpler
and the less differentiated the duct which they have
to stretch in passing out, the better.

The eggs appear to develop out of indifferent
epithelial cells at the dorsal and ventral tips of the
genital glands. Their very first stages we have, how-
ever, not been able to trace. They are first recognis-
able as small groups of four cells with large nuclei
(see Fig. 33), embedded among the undifferentiated
epithelial cells. The nuclei show characteristic
differences from the first stage at which we have found
them. One is clear and round, with one or at the
most two germinal spots ; the other three are slightly
larger and quite full of irregular deeply stained
granules. The former is the nucleus of the future
egg, the latter are the nuclei of the nutritive
cells.

The four cells grow together in one compact mass
to a great size, the partitioning membranes being,
however, traceable. They bulge out the membrane of
the genital tube into the body cavity. In successful
preparations, fine nuclei of a tesselated follicular
epithelium can be found between the eggs and this
membrane (Fig. 33, w). When the egg has been dis-
charged down the branch into the genital tube, the
locus of the egg is found as a small shrunken bag full
of minute round bodies which are doubtless these
follicle cells thrown off by the shrinking of the mem-
brane (Fig. 33,6). This epithelium then apparently
plays no very important part, unless, in some way, it

144

THE APODIDyE

PART I

brings about the contraction of the membrane for the
discharge of the eggs.

The nuclei of the nutritive cells grow to an enor-
mous size, and clearly play the chief part in absorbing
material for the formation of the yolk. When the
egg is ripe, these nuclei come to the surface and
gradually disappear.

FIG. 33.— Eggs at different stages. In stage i the definitive egg nucleus is already
differentiated from the nuclei of the three nutritive cells ; the latter are seen to
grow very large and coarse, and then in 5 to move to the side where they
eventually disappear. In 5 the yolk discs fill the whole egg, making the nucleus
difficult to see. At 6 an egg has been discharged ; the follicular membrane has
shrunk, its contents being probably dislodged epithelial cells, t, testes as
occasionally found (e.g. in A. cancrifcnnis); n, nuclei of follicular epithelium.

The egg, in its passage down the genital tube and
along the longitudinal oviduct, gets covered with a
slimy substance yielded by the deep club-shaped
epithelial cells. This substance hardens into a shell
for the protection of the egg, a shell which, as Von
Siebold remarks, looks as if made of hardened foam.
The eggs pass out between the loth and nth pairs of

SECT, x REPRODUCTION 145

limbs into the brood pouch formed by a modification
of these limbs.

The external aperture of the genital ducts corre-
sponds well, as has already been pointed out, with one
of the apertures of the nephridia of the original Crus-
tacean-Annelid. This view, which is now generally
accepted among zoologists, is supported by the fact
that, among the higher Crustacea, the position of the
apertures is not always constant, i.e. they do not always
occur on the same segment or segments. Indeed, the
male and female apertures may occur on different
segments in the same species (e.g. in the Crayfish).
Wliiie there is variation in this point, there is con-
siderable constancy in the position of the openings
relatively to the ventral middle line of the body.
Both these points are important in homologising the
genital apertures of the Crustacea with the nephridial
apertures of the Annelida.

Many if not all of the Apodidae are now, as was
before stated, hermaphrodite. The small oval sperm
cells form out of the epithelium of the genital tubes
in the manner illustrated (Fig. 33, /). In some species1
the whole of the epithelium at the extreme posterior
end of the genital tube breaks up into sperm cells.

The sperm cells of the Polychaeta are always (?)
thread-like, while those of Apus are round or oval :
this is no doubt a secondary modification. But
the round form of the sperm cells in Apus may
perhaps be the starting point for the many peculiar
shapes found among the higher Crustacea. Other

1 L. glacialis and L. Spitzbergensis, see Fig. 32.

L

146 THE APODID^: TART I

groups again (e.g. the Cirripedia) have returned to,
or retained, the thread-shaped spermatozoa of the
Annelida.

In the development of the eggs and of the sperm
here described, we find but little positive evidence of
the relationship which we seek to establish. But we
must again repeat that it is enough for our argument
if nothing actually contradicts it. It rests upon an
accumulation of homologies which are hardly to be de-
nied, some of which, indeed, have long been recognised
though never before carried out in detail. It is enough
if we show how the other parts of the organisation
of Apus can be deduced from organs of an Annelid.

With regard to the origin of the sexual products, we
have shown more than this. We have drawn attention
to at least one point in which Apus agrees with the
Annelid, and that is, in the development of the egg, not
into the genital tube, which did not exist in the
Annelid, but into the body cavity. The point may
seem to be a small one, but every one who has worked
out the anatomy of Apus will, we are sure, have been at
once struck by the fact, that although the genital glands
are large and extensible, yet the eggs bud outwards
and not inwards. It was this striking method of
development of the eggs which first led us to homo-
logise the epithelium of the genital tubes with the
ccelom epithelium of the Annelida.

The egg, as already described, consists of one egg-
cell and three nutritive cells. As the nutritive cells arc
probably modified egg-cells, the eggs of the original
Crustacean-Annelid may have developed out of its

SECT, x REPRODUCTION 147

coelom epithelium in small groups. So also the small
bulgings in the epithelium in which the sperm-cells
develop may be considered to represent small pustules
containing sperm in the peritoneal wall of an Annelid
(see Fig. 33, /), but in this case the organs which
this wall originally covered, i.e. the nephridia, have
disappeared.

Lastly, we repeat the fact, that in no other Crus-
tacean are the sexual glands segmentally arranged ;
one pair in each developed segment, and rudiments
in the more developed of the rudimentary segments.
The value of this fact for our argument can hardly be
over-estimated.

L 2

SECTION XI
DEVELOPMEN T

THE NAUPLIUS

OUT of the egg of Apus is hatched the well-known
Crustacean larva the Nauplius, which, with certain
characteristic differences for each group, occurs in all
essential points the same throughout the whole class.
The general likeness of the adult Apus to the
Nauplius has, as already mentioned, been pointed
out by earlier observers. This likeness, from our
point of view, is very easily explained ; Apus being
the primitive Crustacean, or at least one of the
primitive Crustaceans, the Nauplius is simply the
young Apus, the adult developing gradually out of
the larva without any metamorphosis worth men-
tioning (see Figs. 34, 39, and 41). Thus the
Nauplius larva of other Crustaceans is simply the
Apus-stage in their development. We repeat this,
not as a conclusion only, but in order to use it as
an argument in support of the theory set forth in
this book.

SECT, xi DEVELOPMENT 149

The fact that a Nauplius stage is passed through
by so many Crustaceans — by all, indeed, where the
larva is not hatched out at a higher stage of develop-
ment than the Nauplius, has received great attention.
It led the older naturalists to assume that the primi-
tive Crustacean must have been an animal like a
Nauplius. This view has, however, generally been
given up, on the ground that no such conclusion
can be drawn from a free-swimming larva which
is certainly much modified to suit its own special
mode of life as larva. The whole argument of this
book has, nevertheless, brought us back somewhat to
the old view, i.e. that the primitive Crustacean was
a Nauplius-like animal, viz., an Apus. At the same
time, the modern objections were largely justified, for
the Nauplius is only a larval form of the primitive
Crustacean, in some respects comparable with, but
much more advanced than, the Trochophoran larva
of the Annelids, showing, on the one hand, traces
of its adult organisation, and, on the other, modifi-
cations to suit its own special method of existence
as a free-swimming larva. There were no means
of deciding which features belonged to the adult
and which to the larva as such. The general like-
ness to Apus was never therefore understood to
point to the fact that the Nauplius was nothing more
nor less than an Apus larva, and that consequently
Apus was a primitive Crustacean. And yet there
seems to be no difficulty in this view ; indeed, had
it been put forward alone, it would, we think, have
met with some acceptance as a plausible specula-

150 THE APODID/E PARTI

tion. In this book, however, we have arrived at
such a conclusion from quite another point of
view. We started by endeavouring to show that
Apus, from its many striking Annelidan charac-
teristics, was a transition form between the Crus-
tacea and the Annelida, and hence a primitive
Crustacean. It comes, therefore, as no slight sup-
port to our argument to be able to show that the
higher Crustaceans pass through an Apus-like stage.
That all Crustacea do not pass through this stage
is easily explained by the theory of abbreviated
development, so that this stage is either passed
through in the egg, or else considerably disguised
by the early acquirement of adult characteristics.
As to the case in which the stage is passed
through in the egg (e.g. among the Malacostraca)
it is important to note that this is not the case
in all Malacostraca, a Nauplius stage occurring,
for example, in the development of Penaeus and of
Euphausia.

The theory of the origin of Apus from an Annelid
gives us at once the true relation of the Nauplius to
the Trochophora. It is not necessary to assume that
Apus passes through a Trochophora stage, because
this latter is a stage in the development of the
Annelid specially adapted to a free-swimming larval
life. The equivalent stage in Apus, being no longer
larval but embryonic, does not require to develop
the special characteristics of the Trochophora.

When the young Crustacean is hatched as a
Nauplius, it has already advanced considerably

SECT, xi DEVELOPMENT

beyond the Trochophora stage. A comparison of
the Nauplius of Apus just hatched from the egg
with the figure of a Polychaetan Trochophora shows
at a glance that the former stands at a far higher
stage of development than the latter. That this is
in reality the case is also clear from the fact that
the Nauplius develops three limbs, i.e., the homo-
logues of the parapodia of the first three Annelidan
segments, traces of the dorsal fold which belongs
to the fiftJi segment, and further slight indications
of five trunk segments (see Fig. 34), in all ten
Annelidan segments. A larva so far developed
cannot be compared with the simple ciliated Tro-
chophora, which when hatched probably represents
only two, i.e. the first and last, segments of the adult
Annelid.

With certain characteristic differences for each
group, the Nauplius is essentially the same through-
out the whole class of the Crustacea. Its exact
morphology we shall endeavour to explain with the
aid of the light we have now obtained as to its
origin, as the larva of Apus, or the Apus-stage in
the development of other Crustacea.

When hatched from the egg the Nauplius has
three pairs of Crustacean limbs, the unpaired " eye,"
the dorsal shield, the large upper lip, and what is
not usually mentioned — the bent intestine, or, what
is the same thing, the rudiments for the development
of such a bent intestine (see Fig. 37). We will
take these points in turn.

I. The Nauplius Limbs. — We are not bound to

152

THE

PART I

claim that these in any way resemble the original
limbs of the primitive Crustacean. It is only neces-
sary to assume that they are homologous with the
first three pairs of Crustacean limbs, but modified for
the special needs of a free-swimming larval life. As a
matter of fact we do find that the form of the limbs
can easily be traced to its origin. The first uni-
ramose limb corresponds with the antenna of the

.-€

FIG. 34. — Nauplius of Apus cancriformis just hatched (after Claus). The large
rowing limb homologised with the dorsal parapodium d, carrying the sensory
cirrus c, which forms the most important branch, the smaller branch being the
tip of the parapodium.

Annelids, i.e. with the sensory cirrus of the vanished
parapodium of the first segment. It arises direct
from the body as a uniramose appendage without
any parapodial portion, i.e. unless the slight bulging,
which is seen at its base in Apus (see Fig. 7 A,
p. 34) represents the remains of such a dorsal
parapodium, which we think improbable, as the
sensory cirri of Annelidan parapodia frequently rise
from such papilla-like swellings. Owing to the

SECT. XI

DEVELOPMENT

153

smallness of the Nauplius it is not easy to ascertain
exactly where the limb springs from ; Claus states
that it rises on each side of the prostomium or
upper lip.

ie second nnio is biramose, i.e. besides the sensory
cirrus, the parapodium on which it stands is also re-

FIG. 35. — Nauplius of Lepidurjis productus (after Brauer) ; commencing segmen-
tation of the trunk disguised. A comparison with the and stage, Fig. 40, shows
that the trunk in the Nauplius certainly corresponds to several segments, d,
dorsal parapodium of the 2nd antenna ; c, sensory cirrus.

tained. Repeating the homologics brought forward
in the section on the appendages pp. 32 and 33, the main
stem of this second limb of the Nauplius is composed
of the dorsal parapodium together with its sensory
cirrus ; the true tip of the dorsal parapodium appears
like a small branch. Here clearly the exopodite is the
sensory cirrus, the endopodite the tip of the dorsal

154 THE APODID^E PARTI

parapodium. That this is the true homology we
have little doubt ; the facts that the second antenna
is a sensory limb, that its tip carries long sensory
hairs, that, as a long rowing foot, it requires to be
provided with a fine sense of touch, all tend to sup-
port it. It is difficult to say whether the thorn-like
process at the base of the limb represents the ventral
parapodium ; it is possible that, in order to facilitate
the motion of the rowing foot, the ventral branch has
disappeared, just as in the higher Crustacea, when the
legs become more specialised as such.

Figs. 35 and 36 are two views of the NaupliusofL.
productus. In these the structure of the whole limb
in the manner we have described is particularly clear,
the sensory nature of the larger branch of the second
antenna being marked by the length of its filaments.

The further development of the limb is interesting.
As it ceases to be a rowing limb and to be specialised
as a sensory organ, one of its branches, that represent-
ing the tip of the original dorsal parapodium, dege-
nerates, leaving the other, the sensory cirrus, i.e. the
exopodite, to form the distal portion of the limb. In
Apus a small rudiment of the endopodite remains
(see Fig. 7 B, p. 34, where the lettering explains the
homologies).

If the thorn-like process at the base of the 2nd
antenna is really the homologuc of the ventral para-
podium, we may perhaps sec in it an attempt on the
part of the very first Crustacean to use the ventral
parapodia of all the segments round the mouth for
mastication, an effort which succeeded well in Limulus.

SECT, xi DEVELOPMENT 155

as we shall see in Part II. In the other Crustacea,
however, the greater efficiency of the ventral para-
podia of the 3rd, 4th, and 5th segments, owing to
their easier concentration round the mouth, led to
their specialisation as mandibles and 1st and 2nd
maxillae, so that the masticatory process of the second
antenna was rendered useless and disappeared (see
table p. 250).

The third limb has again essentially the same shape
as the second. We have the dorsal (and ventral ?)
parapodia, with an appendage on the former homo-
logous with the sensory cirrus or the antennal branch
of the second limb. The dorsal parapodium gradually
disappears in Apus, leaving only the ventral as masti-
catory ridge or mandibles. It is however retained
as palp in the higher Crustacea.

We repeat then here what we have learnt from our
study of the limbs of the adult Apus and of those of
the Nauplius larva. The tip of the dorsal Annelidan
parapodium forms the endopodite of the Crustacean
limb, the sensory cirrus the exopodite, and the ventral
parapodium the masticatory process. Applying this
once more to the trunk legs of Apus, we conclude that
the flabellum becomes the exopodite, and the limb
proper (i.e. the dorsal parapodium) is the endopodite ;
the gnathobase or first endite is the ventral para-
podium, which in the typical trunk limb of the
Crustacea disappears, but may be retained as a
primitive feature, as in Apus, Limulus, and the
Trilobites, and, as on the maxillipedes of the higher
Crustacea, as a masticatory process.

156 THE APODID/E PARTI

We repeat further what was stated on p. 50, that
theoretical considerations would also lead one to
expect a retention of the parts mentioned to form the
Crustacean limb, the capacity of sensation being
necessary to all co-ordinated movement. Hence, as
the dorsal parapodium lengthened into a seizing foot
or locomotory organ, it had everything to gain by
retaining its sensory appendage. It is interesting to
note that when the exoskeleton is so developed that
the limbs are protected by an almost stony covering,
and the limb used simply for walking, the exopodite
disappears, while on the other hand it is nearly always
present in soft-skinned limbs, and generally seems to
have retained its sensory functions. As an instance
of this we can compare the thoracic with the abdominal
limbs of the macrurous Dccapoda.

When we come, in Part II., to consider the relation
of Apus to Limulus and the Trilobites, we shall find
considerable confirmation of the homologies here put
forward, the homologising of the limbs of these animals
with those of Apus being by no means the impossible
task it is too often assumed to be.

2. The Unpaired " Eye'' — The presence of powerful
rowing limbs in the larva necessitated some more per-
fect sensory organ than any possessed by the Anne-
lidan larva ; hence the early development of the un-
paired " eye " which in the Nauplius probably still
retains its visual functions, although these have appa-
rently been lost in the median "eye" of Apus. The
structure of this organ in Apus has been described, and
its probable origin out of the two anterior eye-spots on

SECT, xi DEVELOPMENT 157

the original Annelidan prostomium has been discussed.
\\'e also saw how it, together with the paired eyes,
wandered on to the dorsal surface. In support of this
migration of the eyes, we call attention to Figs. 36
and 37, which show the eyes paired and unpaired
far more anteriorly placed in the Nauplius than they
are later in the adult, i.e. midivay between the ventral
position in tJie bent Crustacean- Annelid and tJie dorsal
Crustacean position.

In Limulus, as already mentioned, the ocelli travel
during embryonic life from the ventral to the dorsal
surface. The homology of the unpaired eye of Apus
with the two ocelli of Limulus assumed here and on
p. 1 08 will be further discussed in Part II.

It is important to note that this sensory organ is
present in all Nauplii, and persists throughout life in
all Entomostraca, but degenerates in the Malaco-
straca. In the more highly developed larvae of these
latter, traces of it are also generally, if not universally,
to be found, e.g. in the Phyllosoma larva of Palinurus,
the Erichthus larva of Squilla, and in some, if not all,
Zoaea larvae. Owing to this almost universal presence
of the unpaired eye among the Crustacea it has been
assumed that it was present in the original Crustacean.
This assumption falls in with our theory that it was
first developed in the Crustacean-Annelid.

In addition to what was said on p. 109 as to the
function of this organ, we may say that its form as a
hollow vesicle full of pigment cells seems at first
sight to suggest an auditory organ, but we share
the growing conviction among zoologists that many

158 THE APODID^: PARTI

sensory organs which are now called auditory really
serve for regulating the position of the body in the
water. The position of the organ in Apus seems to
support this view. The feathered hairs fringing the
flabella are far more capable of appreciating and
responding to sound waves than is a plexus of pig-

FlG. 36. — Nauplius of L. prodnctus from the side (after Brauer), showing the position
of the eyes at the frontal end, i.e. in their passage from the ventral to the dorsal
position.

ment cells in a closed vessel suspended inside the
body some distance beneath the outer integument.
There is here, however, abundant room for further
research. It is possible that in the course of the
development of some Entomostraca, its original
function as a directive body may entirely give place
to secondary visual functions, or, as above suggested,

SECT. XI

DEVELOPMENT

159

it may function both as directive and as visual organ.
Indeed, there is no reason why we should not assume
this double function at least during larval life, so long

^

/ — -i

FIG. 37. — Nauplius of A. cancriformis just hatched (Claus) from the side, showing
the unpaired " eye" in its passage from the ventral to the dorsal position. /,
large upper lip ; s, dorsal shield ; J, dorsal parapodium of the second antennae ;
c, sensory cirrus of the same.

as the paired eyes are not developed, and the animal
is transparent. On the other hand, if it is a directive
body its disappearance in the higher Crustacea is
quite intelligible, especially in the Decapoda, which

i6o

THE

PART I

for the most part crawl, and develop " auditory "
organs in the antennules.

We may in this connection mention the frontal
sensory organs which appear in many (or all ?) Nauplii
(see Fig. 39, f) on each side of the unpaired eye.
They disappear throughout nearly the whole class
in the course of development. They may perhaps

FIG. 38. — Nauplius of A. cancrifortnis just hatched, dorsal view (after Claus).
S, posterior edge of the shield ; N, the large

or neck organ.

larval excretory organ, the dorsal

be supposed to represent a pair of feelers rising on
the prostomium of the original Annelids, such as those
found, for example, on the prostomium of the Eunicidae.
In a section of Apus cancriformis we thought we found
traces of them on the frontal surface in a very
short stiff horn-like process of the cuticle, at the base
of which was a group of large ganglion cells.

SECT, xi DEVELOPMENT 161

3. The Dorsal SJiield. — As a dorsal shield is present
in most Nauplii, it has been generally concluded that
the original Crustacean possessed such a protective
covering. When we come to discuss the relation
between Apus and the Trilobites we shall find that
this was by no means the case. Only in so far as
Apus is the original of all living Crustacea (excepting,
perhaps, the Ostracoda1) can it be said with truth
that the racial form possessed a dorsal shield, at least
as a fold of the fifth segment. The shield of the
original Crustacean-Annelid was itself a different
structure. From it not only the shell fold of Apus,
but also the different forms of bivalve shell have been
developed, as will be described in detail on pp. 213-216.
Again, a further false conclusion has often been
drawn from the great size of the shield in the
Nauplius, viz., that it must have been of about the
same size, relatively, in the original Crustacean as in
the Nauplius. Hence it has been concluded that, for
instance, the Estheridae, which have a large dorsal
shield, are more primitive than the Apodidae with
their relatively smaller shield. The great size of the
shield in the Nauplius, however, admits of a much
simpler explanation. The shield is, as we have seen,
a dorsal fold of the fifth segment. Hence, in larvae
in which only the first five or six segments are
developed, it must necessarily appear relatively of
very great size.

1 Our reason for excluding at least some of the Ostracoda from
the other modern Crustacea which we deduce from Apus will 1>e
discussed in a special section of Part II.

M

162 THE APODID^: PART I

The development of the shield is well shown in
Figs. 35 and 36 of the Nauplius of L. productus, which
should be compared with the commencement of the
development of such a shell in the Trilobite Acid-
aspis, Fig. 48, p. 2 1 5, the neck lobe of which, developed
as a thorn-carrier, suggests a very probable origin of
the dorsal shield. We see it again in the Nauplius of
A. cancriformis, Figs. 37 and 38, developing as a
fold.

No great difficulty need be found in the fact that
the fold of the fifth segment should appear in the
Nauplius before any trace of the limbs of the fourth
and fifth segments, i.e., of the two pairs of maxillae. It
is doubtless of considerable advantage to the larva to
develop the shield as early as possible as a protective
covering.

4. The Upper Lip. — The labrum is another very
characteristic feature of most Nauplii. In some,
indeed, it reaches an enormous size (see Fig. 37).
The homologies of this organ have been a great
puzzle to zoologists. Packard suggests the median
frontal tentacle of certain larval Annelids. Its
presence in the Nauplius seems certainly to suggest
that it was a prominent organ in the racial form of
the Crustacea, especially as it is difficult to see what
special advantage it can offer to the larva as such.
Our derivation of Apus and of the whole class of
Crustacea from a bent Annelid, homologises it, as
already described, with the Annelidan prostomium,
which is probably the most important of all the
external organs of the Annelidan body. Its general

SECT, xi DEVELOPMENT 163

presence in the Nauplius larva of so many Crustaceans,
whether it is afterwards retained by the adult or not,
is thus easily explained. Its relatively great size in
the larva admits of the same explanation as we gave
of the great size of the dorsal shield ; the Nauplius
consisting mainly of the Crustacean head of five bent
segments, the prostomium is naturally a more pro-
minent organ in it than it is later in the adult.

5. The Bent Intestine. — As a characteristic of the
Xauplius not often mentioned we have alluded to the
bent intestine or, in other words, the relative position
of the mid-gut and the mouth under the upper lip
(see Fig. 37, which is a side view of the Nauplius of
Apus showing the position of the parts). We lay
stress on this as a characteristic of all Crustaceans,
the origin of which is explained by the bent Annelid
theory.

The development of the liver as diverticula of the
mid-gut is very clear in Figs. 34, 38, 39, 40.

The important fact that the nerves for the second
antennae spring in the Nauplius from the infra-
cesophageal ganglion has already been mentioned in
discussing the nervous system. This fact serves as a
very striking link between Apus and the Nauplius,
Apus showing in this respect a very primitive condition,
for though the nerves of the second antennae branch
off from the cesophageal commissures, there can be
no doubt that the ganglia are infra-cesophageal (cf.
Section V.).

We may perhaps here briefly summarise what has
been said about the morphology of the Nauplius

M 2

1 64 THE APODID^: PART i

larva. So far from its being comparable with the
Trochophoran larva of its original Annelidan an-
cestors; it is essentially a Crustacean larva, con-
taining from six to ten of the original Annelidan
segments, five of which are bent round to form the
head. This method of development by the appear-
ance at first of the head and the gradual addition of
the new segments has been clearly inherited from the
Annelids, and is, in this connection, very important.
This free-swimming larval Crustacean head develops
chiefly those organs which are necessary to it, those
not especially needful remaining rudimentary. These
useful organs are the anterior pair of sensory antennae
and the second pair of rowing antennae, which are also,
as we have seen, sensory organs. The use of the
third pair of limbs is not clear, unless they serve for
holding on to stationary objects ; it seems necessary
to attribute some function to them, since, if they had
no such larval function, they would probably appear
more in their definitive form. The two pairs of
maxillae have no function to perform in the larva and
are only developed later. It is generally said that
the second pair of limbs degenerates ; this is not
strictly true, only relatively so. They are precociously
developed in the larva, and, according to Brauer's
measurements for L. productus, continue to grow,
not, however, in proportion to the growth of the larva.
The slight change they undergo is due to a change of
function. The shield which belongs to the fifth seg-
ment, being a useful organ, is visible from the first
The unpaired "eye" is developed before the paired eye,

SECT, xi DEVELOPMENT 165

not because it is phylogenetically older, but because
its functions are more useful to the free-swimming but
not at first independently feeding larva. In the larva
of L. productus rudiments of the paired eyes are dis-
tinctly visible. The excretory functions of this larva
are entirely carried on by the large round or oval patch
of glandular hypodermis called the neck- or dorsal-
organ (see Fig. 38, and Appendix V.). Lastly, return-
ing to the gradual development of the Nauplius into
Apus without metamorphosis, we cannot help repeat-
ing that in itself it is a strong argument in our favour
that the Nauplius is but the young Apus, and Apus
but an adult Nauplius (cf. Figs. 34, 39, 41, and
Frontispiece).

One of the chief features, however, in the gradual
development of the Nauplius into an Apus is the
regular formation of new segments in front of the
anal segment as in the Annelid, and the cessation of
growth in Apus before the full number of inherited
and rudimentary segments are fully developed. Apus
is thus, even when adult, little more than a large
Nauplius with its posterior segments in front of the
anal segment fixed throughout life in their larval con-
dition. The significance of this fact is very great,
it shows so conclusively that Apus is a primitive
form, that we cannot refrain from repeating our
explanation of its morphology. The very fact
which has been supposed to be an index of the high
specialisation of the Apodidae, />., the great number
and peculiar arrangement of the limbs, is in reality
one of the strongest proofs of the undifferentiated

166

THE

PART

primitive character of the genus. The number of
limbs is far in excess of the rings in the body, and if
we once recognise that the rings do not correspond
with segments except in the fully developed anterior
trunk region, but that each pair of limbs having its own
pair of ventral ganglia corresponds with a true segment

FIG. 39. — Second larval stage of Apus cancriformis (Claus), showing the gradual
development of Apus out of the Nauplius, the liver as diverticulaof the mid-gut.
/, frontal sensory organs.

either developed or rudimentary, we have an animal,
say Apus cancriformis, with from 60-65 segments.
There are other species with from 40-50 segments.
In all other Crustacea the number of segments is
for each group either absolutely or very nearly con-
stant. The type is fixed. In the Apodidae, as we

SECT. XI

DEVELOPMENT

167

have seen, this is not the case ; the number of
segments varies not only in the different species of
the genus, but, as it appears, in different individuals
of the same species. These two characteristics of the
Apodidae, the great number and the varying number
of the segments, ought almost of itself to constitute
them the natural transition form between the Annelids
and the Crustacea. In the Annelids we have a large
and variable number of segments, in the higher

FIG. 40. — Second larval stage of Lepiditms productns (after Brauer).

Crustacea a comparatively small, and for each group
a fixed, number of segments. Between these two the
Apodidre form the true link, having a diminishing
number of segments, diminishing, that is, by a con-
siderable number remaining undeveloped, and so
rudimentary as to be useless to the animal, and there-
fore liable to vanish.

In this section on the Nauplius we have appealed
to the developmental history of Apus in support of

i68

THE APODID^E

PART

the arguments founded upon anatomical and morpho-
logical comparisons brought forward in the previous
sections. We may, we tfiink, safely maintain that the

FIG. 41. — Fourth larval stage of Apus (Glaus), the diverticula of the mid-gut com-
mencing to form the glandular invaginations (/).

bulk of the evidence to be deduced from the Nauplius
is decidedly in favour of our theory. To us it seems
so strong, that on it alone the theory might almost be

SECT, xi DEVELOPMENT 169

based. All that our argument requires is, that, while
positive evidence is strong, the difficulties should
'not be insuperable.

\Yc have had to limit our remarks to the larval
history of Apus, as observations on its embryological
development are unfortunately wanting. We ma}-,
however, here mention one or two facts in the embry-
ology of the Crustacea which bear upon our theory. It
is stated,1 for instance, that the median eye develops
from paired rudiments — an observation which lends
some support to our account of the origin of this
organ out of an anterior pair of eyes. Still more
important for our theory is the fact that, while in the
development of a few Crustacea there is a tendency
in the mesoblast to form paired, segmented mesoderm-
streaks, in the majority of cases the mesoderm cells
form irregular lacunar spaces.2 The significance of
these two facts taken together cannot be over-estimated,
that is, if we are right in assuming that the latter
method of development of the mesoblast is gradually
displacing the former, and is therefore coenogenetic. It
will be remembered how, from purely morphological
reasoning, we came to the conclusion that the greater
part of the ccelom epithelium (the parietal layer) of the
original Crustacean-Annelid went to form the mem-
brane of the lacunar blood system of the Crustacea.
\Ye find then this acquired rearrangement of the meso-
derm shifted back to the earliest embryonic stages.

1 Grobben, " Die Entwickelungsgeschichte derMoina rectirostris.''

2 Korschelt und Ileider, " Lehrbuch der vergleichenden Entwicke-
lungsgeschichte der Wirbellosen Thiere."

170 THE APODID^ PART i

GENERAL CONCLUSION BASED UPON THE ARGU-
MENTS CONTAINED IN THE FOREGOING SECTIONS.

In pre-Cambrian times, of which there arc now
no fossil remains, a browsing carnivorous Annelid ac-
quired the habit of keeping its " head," i.e., its first five
segments, bent round so that the mouth faced ventrally
and posteriorly, and used its parapodia for pushing
food into its mouth. The antennae, antennal para-
podia, and parapodia developed gradually into Crus-
tacean antennae, mandibles, maxillae, and limbs. For
the protection of the exposed anterior dorsal surface,
a shield, to be more accurately described later on,
was developed out of a fold of the tergum of the fifth
segment, the posterior edge of which grew perhaps as
a carrier of defensive thorns. At the posterior end
of the body, the inherited number of Annelidan
segments gradually ceased to be developed, and
remained in a rudimentary or larval condition.
The gradual development of a thickened cuticle led
to transformations of outer and inner organisation
sufficient to change the Annelidan into the Crustacean
type. The modern representative of this Crustacean-
Annelid is Apus.

We have now to see if it is possible to deduce
the principal groups of both living and extinct
Crustacea either from this racial form or from a
similar Crustacean-Annelid. This is clearly the best
test of the truth of the morphological and anatomical
reasoning contained in this first part.

PART II

PART II

SECTION XII

RELATION OF APUS TO THE OTHER CRUSTACEA

Ix Part I. we have endeavoured, on morphological
and anatomical grounds, to deduce Apus from a car-
nivorous Annelid. We have shown that the trans-
formation of the latter into the former was in
adaptation to a new and very simple change in the
manner of life of the Annelid. If the reasoning of
Part I. is correct, we feel justified in concluding on
the ground of probability that the transformation of
Annelids into Crustaceans only took place once, and
that therefore our bent carnivorous Annelid must form
the root of the whole Crustacean system. Further, it
is clear that the Apodidae must stand very close to
this root. This reasoning leads us at once to find an
infallible test for our whole theory. We have two
lines along which to work, both of which are capable
of leading to a positive answer, negative or affirma-
tive. We shall first take the archaic forms and see
whether they, like Apus, are capable of being deduced

174 THE APODID^: PART 11

from our bent Annelid. And, secondly, we shall see
whether Apus forms a probable starting-point for the
modern Crustacea. In both cases we shall find that
our theory stands the test. We shall find that the
transformation of the carnivorous Annelids into Crus-
tacea did not result in only one form of primitive
Crustacean, but in several. It was, however, the same
Annelid, with the same number (five) of anterior
trunk segments bent round towards the ventral sur-
face, which gave rise to the whole class.

The most important and apparently the most suc-
cessful modification in early times was the Trilobitcs,
that is, if we may judge from the extraordinary num-
bers and varied development of these early Crus-
tacea in palaeozoic times. They, however, all died
out, leaving, perhaps as their sole modern represen-
tatives, some families of the Ostracoda.

Other modifications of the original Crustacean-
Annelid were the Eurypteridse and Xiphosuridaj, to
which latter the still living king-crab belongs.

All these groups, however well adapted to their
palaeozoic surroundings, have, with the exception of
the last-named (and the Ostracoda, which we think
may have come direct from the Trilobites), entirely
disappeared, and it was the Apodidae which became
(with the above exceptions) the sole ancestors of
the now living Crustacea, surviving mainly, we think,
on account of the advantages afforded by the develop-
ment of a dorsal shield.

We have, then, to try to show first, that these Crus-
tacean forms are deducible, like Apus, from the bent

SECT, xii RELATION OF APUS TO CRUSTACEA 175

carnivorous Annelid, and, secondly, that the living
Crustacea, excepting Limulus and (?) the Ostracoda,
may be easily deduced from the Apodida).

If these points can be established, they necessarily
involve a rearrangement of the present system of
classification. The discovery that an animal, which
has hitherto been considered as a very specialised
form of a special group, is in reality one of the
simplest and most original forms of all the groups,
supplies at once the starting-point for the classifi-
cation of the Crustacea which has hitherto been
wanting. It is at present impossible to find points of
connection, sufficient for a natural system of classifi-
cation, between many of the different groups. We
shall now find that the acceptance of our Annelid
ancestor of Apus as the original form enables us, for
the first time, to sketch, at least in outline, a natural
order, not only embracing the Entomostraca and
Malacostraca, but also Limulus, the Eurypteridae,
and the Trilobites. This new classification we shall
attempt, that is, if we are justified in calling that
" new " which is in reality only a further development
of views expressed many years back by the older
zoologists, and notably by Burmeister.

Although we have set ourselves this double task, it
is clearly impossible, in a small work like this, to gc
into many details, especially in our comparison of the
Apodidae with the many living Crustacean forms. It
will, we think, be granted, that a successful grouping
of the Apodidae with the Xiphosuridae, the Trilobites,
and other early forms as common derivatives from a

176 THE APODID^E PART n

bent Annelid, will establish our main argument be-
yond contradiction. We shall therefore devote our
chief attention to endeavouring to explain the mor-
phology of these ancient forms from this point of view,
making, as we believe, many points clear which have
never been properly understood.

As to the second part of our task, the deduction of
the modern Crustacea from the Apodidx, and the
formation of a new system of classification, we shall
have to leave the working out of the details to others,
and content ourselves with a short collection of
notes, to suggest the possible ways in which the
modern Crustacea may be deduced from our bent
Annelid, either through Apus or through the Trilo-
bites.

APUS AND LIMULUS.

We begin with Limulus because, being still extant,
its anatomy is well known. It is to the works of
Anton Dohrn, Kingsley, Lankester, Milne-Edwards,
Packard, and others, that we are indebted for the
details of its organisation here brought forward.

The likeness between Limulus and Apus is so
great, not only in external form but in inner organisa-
tion, that almost all the older zoologists classed them
together in one genus. The temptation to draw com-
parisons between them is traceable in the writings of
all who have dealt with either of them. But, in spite
of this unmistakable likeness, all idea that the two
animals could possibly be related has in later times
been steadily repudiated. Indeed no general agree-

SECT, xii RELATION OF APUS TO CRUSTACEA 177

•

mcnt has been arrived at as to the true zoological
position of Limulus. Many eminent zoologists, such
as Van Beneden, maintain that Limulus is not a
Crustacean at all;1 and the able attempt of Lan-
kcster and others to demonstrate that Limulus is
an Arachnid is familiar to all zoologists.

The difficulties in the way of connecting Limulus
and Apus seem to be the following, (i) The limbs
in the two animals are differently arranged on the
body, besides differing in number and form. This
point is rightly considered of great importance,
because it was chiefly the close study of the limbs,
and of their homologies in the different Crustacea,
which enabled zoologists to arrange the class into
the natural groups of our present classification. (2)
Whereas the Xiphosura bear markedly the charactcr
of an archaic group, whose nearest allies are to be
sought for in the earliest geological strata, and which
in development pass through a so-called " Trilo-
bite" stage, Apus has, comparatively speaking, no
geological record, and is, so far as we can learn from
palaeontology, rather a highly specialised tertiary
form. (3) The young of Limulus do not pass
through any stages which appear to correspond with
the stages of Apus. Packard's attempt to discover
the Xauplius stage in the embryological development
of Limulus has met with no favour.

We here have, as far as we can find, the chief

1 " Les Limules ne sont pas des Crustaces — ils n'ont ricn de commun
avec les Phyllopodes." Journal de Zoologie, par P. Gervais. vol. i. p. 42.
Paris. 1872.

X

178 THE APODIDyK PART n

reasons why zoologists have not allowed themselves
to be influenced by the extraordinary morphological
likeness between the two animals, and why they
have maintained that this likeness is merely a
remarkable case of analogy.

On the other hand, the morphology of Apus has
been such a perpetual puzzle, that its likeness to
Limulus, even combined with its acknowledged re-
tention of Annelidan characteristics, gave no key to its
systematic position, just as its likeness to the Nauplius
failed to suggest that it is itself the proto-Nauplius.
When once, however, we recognise the essentially
Annelidan and therefore primitive character of Apus,
and thus regard it as an archaic form, i.e., as a sur-
vival from early geological periods, its likeness to
Limulus takes on at once a new meaning. The diffi-
culties above mentioned deserve to be re-examined ;
fortunately they arc not insuperable. Before going
into a detailed comparison of Limulus, let us briefly
indicate the way these three difficulties may be met.

I. In the first place, the difficulty as to the difference
between the limbs of Limulus and Apus depends
entirely upon an exaggeration (a very natural ex-
aggeration) of the importance of limbs for the
purpose of classification ; we say, a very natural
exaggeration because, as above stated, it has been
by a close study of the homologies of the Crusta-
cean limbs that so much has been done to arrange
the Crustacea into natural groups. On the other
hand we ought not to lose sight of the fact that
of all organs of the body the appendages are the

SECT, xii RELATION OF APUS TO CRUSTACEA 179

most plastic ; the slightest alteration in habit of
life, and every change in size and form of the
body, bring about some corresponding change in the
limbs. So that while, on the whole, stability of
type is wonderfully exemplified in the Crustacean
limbs, too much weight must not be laid upon it,
since the same class supplies us with equally won-
derful examples of extreme plasticity. Specialisation
for some particular habit of life leads often enough
to modification which altogether obliterates the type.
It is not safe, then, to conclude, because the limbs
of a Crustacean do not now show the typical form,
that there is no way of connecting them with
typical limbs. We thus maintain that the assumed
failure of Limulus to show the typical Crustacean
or Phyllopodan limbs ought not for a moment to
weigh against the positive likeness between it and
Apus.

Further, while Apus has, as we have seen, retained
the more primitive form of limb, not far removed from
the Annelidan parapodium, the manner of life of
Limulus has led to a specialisation of its limbs, but
not, it is important to note, to such extreme
specialisation that no points of resemblance with
the limbs of Apus are retained. On the contrary,
the likeness, in some respects, is so great that one
might almost be tempted to leave the limbs out of
account in the question of relationship ; they speak
equally strongly both ways. When we come to
compare the animals in detail, Apus having supplied
us with the clue, it will not be difficult to deduce

N 2

i8o THE APODID^E PART 1 1

the limbs of Limulus from the parapodia of our
Annelid, and to explain the transformations which
have taken place.

The first difficulty as to the form and order of
the limbs is thus, we think, fairly satisfactorily met
for the present by the following four considera-
tions :—

(1) That the possibility of homologising the limbs
with typical Crustacean limbs must not be too much
insisted upon, in the face of the well-known plas-
ticity of these organs.

(2) That the limbs of Limulus are in many points
as strikingly like the limbs of Apus as, in other
respects, they are unlike.

(3) That the modification of the Xiphosuran limb
out of the Phyllopodan or Annelidan is fairly easily
traceable to the manner of life of the animals.

(4) As to the number of the limbs — our whole
theory makes the number of limbs or segments
developed of no real importance. The method of
the development of new segments is such that few
or many may be developed according to the needs
of the genus.

II. Turning to the geological difficulties, we think
these of even less weight than those founded upon
the dissimilarity in the form, number, and order of
the appendages. The habit of life of Apus from
earliest times must have been such that it could be
very seldom preserved in a fossil state. It was
probably first shut off from the ocean in brackish
lagoons, and was gradually driven by the struggle for

SECT, xii RELATION OF APUS TO CRUSTACEA 181

existence into small fresh-water pools, where alone
it was able to hold its own, shut off from compe-
tition with almost all the rest of the animal king-
dom. It is in this way, as already stated, that we
account for the preservation of its primitive charac-
teristics. Now, in such a record as this, what are
the probabilities of its leaving any fossil remains ?
The marine carnivorous Annelids of palaeozoic
times have left only their hard, chitinous teeth,
so that the Apodidae of those times, with a skele-
ton not much harder than that of the Annelids,
would hardly be likely to be preserved. Their
comparative softness is thus one element to be
taken into account in discussing the probability
of their being preserved as fossils. But, further,
when once they had adopted their fresh-water life
in shallow pools, the chances of their preservation
would be smaller still. They would at this time
belong to the land fauna. There would thus be
very little chance of their remains being preserved.
In the first place the dead bodies would have
decayed before there was any chance of their being
covered by a deposit ; there is, as a rule, very little
suspended matter to fall in the isolated fresh-water
pools which we suppose the Apodidae to inhabit.
And in the second place, land surfaces are, as is well
known, seldom if ever preserved. There is therefore
very little chance of any Apus being preserved
excepting under very exceptional circumstances.

Further, although there may be no true Apodidae
recorded from Palaeozoic strata, yet there are abun-

1 82 THE APODHLE PART n

dant remains of Phyllopods, many of which show
such a striking resemblance to the Apodidae that we
are justified in claiming them as nearly related forms.
This fact lends distinct support to our argument that
Apus is a very ancient form, in spite of the deficiency
in its own geological record.

III. The embryological difficulty is even of less
account than the two others. Packard, whose studies of
the embryology of Limulus entitle him to speak with
authority, states that it is evident that the metamor-
phoses are all undergone within the egg, in order
that the young may enter at once on the manner
of life of the adult. And we may repeat here
what has been affirmed in other connections, that
it is enough if the embryology of Limulus does not
directly and plainly contradict our theory ; we say
plainly, because we do not lay much weight on the
passing hints which an animal in its development
may give as to its 'ancestors, unless these hints are
supported by other evidence.

If these are not completely satisfactory answers to
the difficulties which stand in the way of any close
relationship between Limulus and Apus, they at any
rate weaken those difficulties to such an extent that
they are of little value in comparison with the positive
evidence based upon the anatomical and morphological
likeness between the two animals, taken together with
all the evidence brought forward in the first Part to
show that Apus has retained most of the characteristics
of a primitive Crustacean, and, in fact, is as truly an
archaic form as Limulus itself.

SECT, xii RELATION OF APUS TO CRUSTACEA 183

Having prepared the way, let us commence the
detailed comparison of the organs of Limulus and
Apus.

The first point on which we fix our attention in
order to test the relationship between the two animals
may not appear very important, but the longer it is
considered the more convincing, it seems to us, is the
argument founded upon it. It is as follows :

Our main argument is that Apus is a bent Annelid.

FIG. 42. — Section of Limulus rot ntidicauda to show the bend in the intestine with
the sinewy sternal plate in the angle to be compared with that of Apus Fig. 13,
p. 56. b, brain ; //, heart ; /, openings of the hepatic ducts in the mid-gut ; sj>,
the sternal plate. From Bronn's Klasscn und Ordnttngen des ThierreicJifs.

In this way we explained the bend in the intestinal
canal which is so characteristic of the Crustacea.
This bent intestine is very marked in Limulus (see
Fig. 42), and suggests the same origin. This, how-
ever, was not enough. We argued that if we find in
the bend of the intestine of Limulus a sinewy mass
such as we find in Apus, referable there to the
clumping together of the abdominal musculature,
the coincidence can hardly be a mere case of
analogy. Such a sinewy mass is found in Limulus, in

184 THE APODID^E PART n

essentially the same place as in Apus, and is known
as the sternal plate. If we have given the right
explanation of these two points, the bent intestine
and the sinewy mass in the bend in Apus, there can
hardly be any other explanation to be given of their
occurrence in Limulus. Given, then, the derivation of
Apus out of a bent Annelid, in the manner described
in the first part, it is hardly within the range of pro-
bability that Limulus, in which these two essential
marks of such a derivation occur, should have had a
different origin. These two points of resemblance,
occurring together, are, in our opinion, conclusive as
to the essential relationship of Limulus to Apus,
through their common origin from a bent Annelid.
This alone without further positive evidence was
sufficient to convince us that Apus and Limulus
were at least branches of the same stem. But, as
will be seen in the following pages, the whole
organisation of Limulus admits of direct com-
parison with that of Apus, the very differences
between them affording striking confirmation of
our theory of their common origin from a bent
Annelid.

With this decisive evidence in favour of our theory
we naturally proceeded with considerable confidence
in attempting to homologisc the limbs which have
hitherto presented the chief difficulty in connecting
Apus and Limulus. Before commencing a detailed
discussion of the limbs of Limulus, there arc many
points of resemblance in external organisation which
should be mentioned. It will also be useful to

SECT, xii RELATION OF APUS TO CRUSTACEA 185

ascertain here the general principles on which Limulus
has been modified.

On comparing the external form of Limulus with
that of Apus, we find that the head region, while
retaining essentially the same form, is yet far
larger in proportion to the size of the body in the
former than in the latter. The eyes are further
apart, and two ocelli take the place of the unpaired
" eye " or directive sensory body. The dorsal shield
does not stand out from the body as a fold like that
of Apus. Its frontal and lateral edges are produced
downwards and outwards, so that the anterior part of
the animal proper is raised from the ground, and,
under the shield, has room for the movement of its
limbs. 1 The dorsal shield, in fact, forms a sort of roof
under which the animal lives. It is as if the head
and anterior trunk segments of an Apus had been
pushed upwards and backwards under the shield,
being somewhat squeezed up in the process. The
whole life of Limulus is passed under cover, every
possible entrance being shut, or carefully guarded,
against enemies.

The fusing of the head with the dorsal terga of a
certain number of trunk segments naturally deprives
these segments of any power of movement one upon
another. The abdomen is also quite rigid, although
in its embryonic state it shows distinct external traces

1 This is not shown in the section of Limulus rotundicauda, Fig. 42,
which, being a median longitudinal section, passes through the forehead.
The vaulted shape of the shell is best shown in transverse sections, cj.
the sections of a Trilobite Fig. 54, p. 230.

1 86 THE APODID^E PART n

of segmentation. The length and rigidity of the
caudal spine, taken together with the rigidity of the
anterior portion of the body, would render the flexi-
bility of the middle part of little use. It is true that
there are Trilobites with rigid shields and pygidia
and yet with flexible segments in the middle region
of the body ; but the pygidium could, in these animals,
be used as a covering in the rolling up of the body.
In animals possessing a caudal spine there could
be no question of its being put to any such use ;
hence, probably, the rigidity of the middle or ab-
dominal region of the body.

There are, comparatively speaking, but a small
number of segments in the Xiphosuridse, at least as
compared with Apus ; but this is a matter of com-
paratively small importance if we take into con-
sideration the method of development of the early
Annelidan-Crustaceans. The hind part of the body
might become fixed at almost any stage of develop-
ment, more or fewer segments coming into existence
according to the degree of specialisation of each
group. Apus developed, comparatively speaking, a
large number (50-60), Limulus a small number
(ca. 1 6).

The metamerism of Limulus is probably to be
reckoned as follows :

Ceplialotliorax. — -This is composed of five segments
of the bent Annelid forming the head, each segment
retaining its appendages, and of two trunk segments
bearing two pairs of limbs, the posterior pair forming
the operculum ; in all seven segments.

SECT, xii RELATION OF APUS TO CRUSTACEA 187

Abdomen and Caudal Spine. — These probably re-
present nine 1 segments, of which the first five carry
leaf-like gills, four (the last of which develops into the
caudal spine) remaining limbless ; these latter are to
be compared with the five or six limbless segments
of Apus. The caudal spine is a development of
the anal segment homologous with the tail plate of
Lepidurus. i.e., of those Apodidae in which the anal
segment is produced posteriorly into a flat plate.

According to its external organisation, therefore,
Limulus is an Apus-like animal, especially adapted
for living on mud under a shell. The shell is vaulted
and the body correspondingly compressed against its
roof, so as to allow the limbs, &c., to function. In
this way we think that the chief differences between
Apus and Limulus can be explained. It is important
to bear in mind this general principle on which
Limulus has been modified as a key to its special
organisation.

It is worth pointing out that Packard, quoting
from Dr. Gissler, describes the method of moulting
in Limulus and Apus as being essentially alike.
This, however, need be no more than a case of
analogy.

In the following detailed comparison we shall find
that in some points Apus is the more specialised, in
others Limulus, but there can be little doubt that, of
the two, Apus stands nearer to the original Crusta-
cean-Annelid.

1 Packard gives this number for the abdomen of Limulus.

ISS

THE APODID/E

PART ll

THE LIMBS.

As already stated (p. 179) we find that the differ-
ences between the limbs of Apus and of Limulus are

FIG. 43. — Ventral surface of Lininlus moluccanus 9 (after Van der Hceven), showing
the ventral parapodia of five pairs of limbs, viz., the second, third, fourth, fifth,
and sixth (first trunk limb), working between the prostomium and the under
lip ; also the differentiation of the sixth, or fir.st trunk limb, for locomotory
purposes.

as significant as are the likenesses. First as to number,
we need only repeat what was said about the difference
in number of the segments ; instead of a large number
of segments with a large number of limbs at different
stages of development, from the parapodium-like limb

SECT, xii RELATION OF APUS TO CRUSTACEA 189

of the Phyllopocla to the Crustacean leg, we have in
Limulus a small number of segments with a small
number of limbs, showing essentially the same
differentiation as we find in the legs of Apus, but more
specialised in adaptation to its manner of life. We
have anteriorly the more typical Crustacean limb,
posteriorly the more parapodial limb, the transition
between the two, however, being not gradual but
sudden.

In trying to homologise the limbs of the two animals,
we shall have to utilise the conclusions arrived at in
Part I.

The first pair of limbs of Limulus occupies a place
corresponding to that of the first pair in Apus, viz.,
at the sides of or close to the labrum, and is homo-
logous with the first pair of antennae. In general
form these limbs do not differ much in the two animals,
as may be seen by comparing Fig. 43 and Frontis-
piece. The chief difference is that in Limulus the
bends have developed hinges, owing to the greater
development of the exoskeleton, and there are chelae
at the tips. We need hardly say that neither of
these points is of very great morphological worth.
For instance, within the Arachnoidea we find one
and the same limb, the pedipalp, in one group (the
Araneidae), as a simple feeler ; in another (the Scorpio-
nidae), it develops powerful chelae ; and within the
group of the Spiders themselves the tips of the pedi-
palps in many males undergo even more wonderful
modifications for the purposes of copulation.

With regard to the second antenna, we were at first

190 THE APODID^E PART n

disposed to think that it had entirely disappeared.
In Apus it has not yet quite disappeared, but it is so
rudimentary that it appears to be in the act of disap-
pearing. It seemed to us that the condition of this
limb in Apus helped us in pointing out a missing limb
in Limulus. But maturer thought led to the homolo-
gising of the second pair of limbs with the second
antennae of Apus. In the first place the position
agrees very well (cf. Fig. 43 with Frontispiece).
This was not, however, the real reason for our change
of opinion, which was due to a comparison of the sixth
limb in Apus with the sixth limb in Limulus, and, to
anticipate somewhat, with the sixth limb in the Eury-
pteridae and in the Trilobites. The sudden specialisa-
tion of this limb in all these animals must be admitted
to have some common significance. That given on
p. 44 seems the most probable, viz., that, taking five
segments to form the bent head, the sixth wa.s the
first free segment, and its parapodium was thus free
to develop into a limb for locomotion or for some
other function. Thus, taking the sixth limb of
Limulus to represent the first trunk limb, the full
number (five) of head limbs were left to be homologisecl
with the typical head limbs of Apus and the other
Crustacea. The second antenna is therefore present,
exactly corresponding in position with the homologous
limb of Apus. It is, however, a well-developed and
highly functional limb. In the female it is chelate,
but in the male it develops a seizing hook. It is in-
teresting to note that the same pair of limbs in the
male of Branchipus, which is closely related to Apus,

SECT, xii RELATION OF APUS TO CRUSTACEA 191

also develop powerful seizing hooks. The homology
of the second pair of limbs of Apus with the second
antennae of the Apodidae is doubly interesting because
we here find the ventral parapodium retained as
masticatory ridge. The great number of masticatory
ridges in Limulus will be referred to again, and com-
pared with the number of homologous ridges in Apus
and the fossil Crustacea.

The third pair of limbs of Limulus offers a most
interesting comparison with the mandibles of Apus.
In the latter, the ventral parapodium alone is retained,
the dorsal having entirely disappeared. In Limulus,
both have been retained, the ventral parapodium as a
very pronounced masticatory ridge, the dorsal as a
long jointed chelate leg.

The fourth and fifth head limbs have nearly the same
form as the third, and are homologous with the two
pairs of maxillae of Apus. One interesting feature,
however, deserves special attention in connection
with our deduction of Apus and also of Limulus
from an Annelid. In Limulus, the ventral para-
podium, which stands out much more pronounced
as a ventral parapodium than it does in Apus, has
retained distinct traces of its sensory cirrus (see Fig.
44). In this respect Limulus is more primitive than
Apus. On the other hand in Limulus, the sensory
cirrus (or exopodite) disappears from the dorsal
parapodium of the head limbs ; this is the exact
opposite of what we find in the typical Crustacean
limb, where the dorsal parapodium as endopodite and
its sensory cirrus as exopodite are alone preserved, thq

192 THE APODID^E PART 11

ventral parapodium being only occasionally retained
as a mere rudiment. This very striking difference
between Apus and Limulus is thus at the same time
a link, since it connects them both with our Crustacean
Annelid.

The sixth pair of limbs has already been homologiscd
with the first trunk limbs of Apus and the other Crus-

FIG. 44. — Basal part of one of the 2nd pairs of limbs of Limulns molnccanus (from
Bronn's Kiassen und Ordnimgen dcs Thierreiches) to show the well-developed
ventral parapodium, retaining the remains of its sensory cirrus (c)\ d^ dorsal
parapodium ; v, ventral parapodium.

tacea. It differs characteristically from the head
appendages. Its ventral parapodium is well developed
as a masticatory ridge, and functions as a jaw, in front
of the under lip, with the masticators of the last four
head segments. Its dorsal parapodium, however, is
developed in a peculiar way. It is a powerful limb for
pushing the animal forward in its burrowing operations ;
its tip is provided with a group of sensory feelers

SECT, xii RELATION OF APUS TO CRUSTACEA 193

comparable with the sensory endites on the ventral
edge of the trunk limbs of Apus ; in the middle of
this group of sensory processes is a small pair of
chelae. This whole limb seems to combine loco-
motory with protective functions. Its tip lies close
to the lateral gap between the cephalo-thoracic and
abdominal shields, so that no enemy could enter with-
out being immediately perceived and thrust out. This
first trunk limb seems to have preserved its dorsal
sensory cirrus, also no doubt as a guard against the
entrance of enemies which could not easily be ejected
if once lodged on the dorsal edges of the head limbs^
i.e. between the limbs and the shield.

The last pair of limbs of the cephalo-thorax is highly
modified as a flat cover or operculum for the abdominal
feet. Its form is essentially Phyllopodan. In Limulus,
the passage from the flat Phyllopodan limb to the
Crustacean leg is abrupt ; there are no transition forms
as in Apus. The first trunk limb is leg-like, the second
Phyllopodan. As the form of the latter is essentially
the same as that of the abdominal limbs, we reserve
our description of it till later.

Before leaving these cephalo-thoracic limbs we wish
once more to call attention to their arrangement,
which is well explained by the manner of life of
the animal. The animal, i.e. its anterior end, seems
as if fixed up in the vault of a roof, the mouth
being in the centre and the limbs hanging down all
round. The most anterior and most posterior limbs
do not function as jaws, but the five limbs between
these two, i.e. the last four head limbs and the first

O

194 THE APODID/E PART n

trunk limb, have powerful jaw pieces, which are
arranged in two rows, one on each side of the oral
aperture, the dorsal parapodia being developed into
an outer ring of chelate feet for seizing prey and
bringing it between the jaws.

It is clear that the efficiency of these long limbs,
already limited as to space for free movement, would
be materially lessened if on their dorsal edges they
had to carry gills, &c. ; hence these appendages have
entirely disappeared, respiration becoming localised on
the abdominal limbs, which have been especially modi-
fied for the purpose. In the Trilobites the movement
of the limbs is also limited by a large shield running
the whole length of the body, but in their case both
the gills and the cirri are retained, the reason being
very simple, viz., that the ambulatory legs of the
Trilobites do not require to carry out the complicated
movements of the limbs arranged round the mouth of
Limulus, but are simply ambulatory legs.

Two processes from the ventral surface of the
body bring these limbs to a close. The mouth parts,
i.e. the masticatory ridges, work between the labrum
at the one end and these two processes at the
other. Although authors have recognised that these
processes function as an under lip, they hesitated to
assert that morphologically they were the true under
lip. We can, however, hardly doubt that they are to
be homologised with the under lip of Apus. An
examination of the special modifications of Limulus
explains the position of the organ ; the mouth has been
lengthened out longitudinally so that the labrum has

SECT, xil RELATION OF APUS TO CRUSTACEA 195

been forced outwards and forwards, and the under lip
backwards, so as to admit of the working of the five
pairs of jaws between them. If it at first sight seems
unlikely that the paragnatha should move so far back
as to come behind the first pair of trunk feet, it must be
remembered that, when the mouth of the Crustacean-
Annelid first stretched out so as to admit of the working
of the five ventral parapodia as jaws, the parts were
more flexible. It is also some confirmation of this
homology to find that the sternal plate, the sinewy
mass of the musculature originally in the angle of the
bent intestine and thus close above the under lip, is
also drawn back as far as this under lip, showing that
the whole region has been drawn out of its original
shape. The origin of the division of the under lip
has been already explained (p. 40).

In Limulus, as already said, we do not find, as in
Apus, a gradual change in the limbs from the more
Crustacean form anteriorly to the more Annelidan,
i.e. parapodial, posteriorly. The transition is sudden.
The limb corresponding with the second trunk limb
of Apus forms the flat operculum to cover the follow-
ing five pairs of Phyllopodan (i.e. parapodia-like)
limbs.

The typical parts of these abdominal limbs can still
be more or less clearly recognised. The ventral para-
podia of each pair of limbs have fused in the middle
line, forming the basal plate ; the dorsal parapodium
is represented by a row of four joints approaching
the middle line (see Fig. 45). On the outside of
these come the large gill and somewhat smaller flabel-

O 2

196

THE APODID^E

PART II

lum fused with one another and with the basal plate,
but still distinct. On the well-known principle of the
increase of the respiratory surface by the formation
of integumcntal folds, the dorsal (i.e. morphologically
posterior) surface of the gills has developed a row of
leaf-like folds.

It is of importance to note that it is only that part

FIG. 45. — One of the abdominal limbs ofLimtdmfor comparison with a Phyllopodan
limb. ?/, ventral parapodia fused together; ci, dorsal parapodia (endopodites) ;
Jl, flabellum or sensory cirrus (exopodite) ; g, gill, the posterior surface of which
carries leaf-like integumental folds.

of the limb of Limulus which morphologically cor-
responds with the gill of the parapodium (or of the
Phyllopodan leg) which is thus modified. One would
have been inclined to think that the increase of the
respiratory surface could have been better obtained
by using the whole surface of these leaf-like limbs.
We have here a most interesting case of the strict
localisation of function. The increase of respiratory

SECT, xii RELATION OF APUS TO CRUSTACEA 197

surface required, in consequence of the suppression of
the gills on the anterior limbs, is obtained by a com-
plicated increase of the surface of the gills on the
other limbs, and of the gills only.

We have, then, compared the appendages of Limu-
lus with those of Apus, and shown how they throw
light upon one another, how. they are both deduciblc,
some along one line of special differentiation, some
along another, from the parapodia of our original
Crustacean-Annelid. The type is clearly the same
•in both, as is also the type of the whole organisation
of the two animals.

The Musculature. — The musculature of Limulus is
very specialised, in accordance with the specialisation
of the outer body and the high development of
the exoskeleton. We will not here venture on the
task of endeavouring to trace the separate muscles
from their Annelidan origin. In all such attempts,
the limitation of the movements of the body must
be borne in mind. The fact that the body of Limulus
is no longer capable of any bend, except in the
sagittal plane, would alone bring about very profound
changes in the musculature, which originally moved
the Annelid freely in any plane.

It must here suffice to refer to what was said above
(p. 184) about the sinewy mass found within the bend
of the intestine for the attachment of the muscles.
For the origin of this mass from the Annelidan ven-
tral muscle bands we refer to the derivation given in
Part I. of the similar mass found in the same place in
Apus. And further we can point to the entapophyses

198 THE APODID^: PART n

in Limulus as homologous with the points of attach-
ment of the dorso-ventral muscles in Apus, marked
/ in figures 66 and 67.

T/ie Nervous System. — The nervous system of
Limulus is especially important and interesting. In
many respects it is more primitive than that of
Apus, for example, in the position of the brain. On
the other hand, again, owing to the greater special-
isation of the whole body, it is in some respects
more specialised.

In describing our bent Annelid we naturally found
it necessary to assume that the brain was originally
in the prostomium or labrum. In Apus, owing to
the wandering of the eyes forwards and upwards, the
brain followed the eyes, splitting the cesophageal com-
missures into a sympathetic ring and a cerebro-ceso-
phageal ring. In Limulus the brain has retained its
original Annelidan position. It need hardly be said
that this is a very striking confirmation of our deriva-
tion of Apus from a bent Annelid. We had quite lost
sight of this fact when we stated that in the original
Crustacean-Annelid (shojyn in Fig. 18, p. 69) the
brain was in the prostomium, and that in Apus it had
wandered from its place through secondary adapta-
tions. Thus the very difference in the position of the
brains of Apus and of Limulus affords a conclusive
proof of their real relationship as derived from the
same bent Annelid.

It is almost equally important- for our argument to
note that, as R'ay Lankcster. pointed out, the brains of
Apus and Limulus are alike in constitution, both

SECT, xii RELATION OF APUS TO CRUSTACEA 199

being very nearly pure archicerebra. They consist
almost wholly of the ganglia for the eyes and ocelli in
Limulus, and of the eyes and unpaired sensory body
in Apus. In Limulus, according to Packard, the brain
is not complicated by the presence of the ganglia for
the antennae ; in Apus, however, according to Pelse-
neer, the ganglia for the first antennae have joined the
brain. In the great simplicity of the brain, these two
animals are, Lankester states, almost unique.

The eyes in Limulus, in wandering forwards and
outwards, were unable to take the brain with them,
but are simply connected with the brain by long,
and not very important, nerve fibres. These nerve
fibres have the same relative position on the brain as
the stalks of the optic ganglia of Apus. Between the
two optic nerves, a pair of nerves is found running
to the pair of ocelli which lie anteriorly near the
middle line. \Yc have already referred to these
median ocelli of Limulus, as some support for our
argument that the median sensory body in Apus
arose out of an anterior pair of eye-spots on the
prostomium of the original Crustacean-Annelid.
The position of the points of departure of the
nerves to these ocelli agrees exactly with that of
the nerves to the unpaired " eye " of Apus. In
Limulus there are other nerves leaving the brain from
between the optic nerves besides those to the ocelli.
In Apus we found that the sensory body is com-
posed of four retina:, with four nerves running to the
brain. If we homologise the lateral retina; with the
ocelli, the nerves from the postero-dorsal and ventral

200 THE APODID^: PART n

retinae might correspond with a pair of the other
nerves just mentioned which leave the brain near
those of the ocelli. The different shape and grouping
of the sensory cells of the postero-dorsal and ventral
retinae from those of the lateral retina: seem to
indicate that they must have been derived from some
other sensory organs.

Owing to the backward prolongation of the mouth
and the oesophagus, and the arrangement of the
limbs round the former, the anterior nervous system
is very concentrated ; the nerves for the anterior
antennae and the five pairs of limbs branch out
radially from the thickened cesophageal commissures.
It is as if the oesophagus had forced its way back-
wards between the two longitudinal commissures of the
nerve cord, forcing apart the separate pairs of ganglia
of the first five pairs of limbs, the four transverse com-
missures of which arch over the slanting oesophagus.

Between the nerves to the fifth pair of limbs and
those to the operculum is a pair of nerves to the
chilaria or under lip. If the homology of the
chilaria with the under lip of Apus is correct, these
nerves have been carried back with the under lip,
in the drawing back of the mouth.

The ventral cord of Apus is more primitive than
that of Limulus, which, at its posterior end, is much
modified. This specialisation of the ventral cord of
Limulus is in correspondence with the great concen-
tration of its body as compared with that of Apus.
In Apus the posterior end was found in a rudi-
mentary and larval condition.

SECT, xu RELATION OF APUS TO CRUSTACEA 201

The arterial envelope surrounding the nervous
system will be referred to later in the paragraph on
the circulatory system.

The Sensory Organs. — We have already (Fig. 22,
p. 91) described and figured the eyes of Limulus in
order to explain the origin of the typical Arthropo-
dan eye of Apus from the Annelidan eye-spots. It
is of no small interest to remark that we had selected
the eye of Limulus as a guide towards explaining
the origin of the Crustacean eye at the very outset
of our investigation, when we were entirely occupied
in attempting to deduce Apus from an Annelid, and
long before it occurred to us that Limulus was
probably related to Apus. The establishment of
the relationship between the two thus lends con-
siderable support to the theory put forward in Part I.
as to the possible development of the Arthropodan
eye out of an Annelidan hypodermal eye-spot by
the thickening of the cuticle. If this deduction is
correct, then the eye of Limulus is more primitive
than that of Apus. This indeed we might expect
from the manner of life of the two, the free-
swimming form naturally having the more perfect
visual organs, while Limulus, which burrows in mud
or sand and lives practically under a roof, has eyes
comparatively weakly developed

The wandering of the eyes from the ventral surface
on to the dorsal, which we found indicated by the
bend of the cerebro-cesophageal commissures in Apus,
is here shown in an equally interesting way by the
upward, forward, and outward bend of the long optic

202 THE APODID^: PART n

nerves. Their very length, when compared with the
usual distance between eyes and brain throughout the
animal kingdom, is a clear indication of displacement.

The anterior pair of Annelidan eyes, which in Apus
went to form the unpaired " eye," are represented in
Limulus by a pair of ocelli. The wandering of these
ocelli on to the dorsal surface can still be traced in
the course of the animal's development. According
to Packard, the ocelli at their first appearance in the
embryo are on the ventral side, and travel on to the
dorsal side before the young animal is hatched. The
true significance of this fact has already been dwelt
upon, and has been compared with a similar, though
not so pronounced, wandering of the eyes in the
Nauplius as shown in Figs. 36 and 37. The presence
of the ocelli on the ventral surface of any ancestor
of Limulus would be difficult to explain by any
other theory than that of our bent Annelid. The
nerves to these ocelli branch from the brain from the
same place as do those to the unpaired " eye " in
Apus, i.e. from between the optic nerves.

Judging from the lateral retinae of the sensory body
of Apus, and also from the fact that the posterior
eyes are compound, we should have expected com-
pound eyes and not ocelli as the anterior pair in
Limulus. In certain Trilobites (e.g. Harpes), accord-
ing to Barrande, these ocelli are not single but com-
posed of groups.1 It seemed to us that these might
perhaps form an interesting connecting link between

1 See Packard's paper on the structure of the eye of Trilobites.
American Naturalist^ July 1880.

SECT, xii RELATION OF APUS TO CRUSTACEA 203

retinulated compound eyes and the single ocelli of
Limulus. We find, however, that the ocelli of
Limulus, according to Lankester's and Bourne's
figures, are only ocelli in the sense that they have
but one large cuticular lens ; the retinal cells under
them being grouped in retinulae. The presence of
retinula^ essentially of the same shape as those
under the conical cuticular projections in the paired
eyes (see Fig. 22) suggests that this large cuticular
lens has arisen by the coalescence of a number of
such crystal cones ; otherwise, according to our
view, it would be difficult to account for the reti-
nulae, which we think first arise by the grouping of
the sensory visual cells round the tips of the conical
refractive processes. If this is the case, the ocellus
of Limulus is not due to an independent utilisation
of a special form of cuticular thickening, as we
think is the case in such an eye as that of the
Dytiscus larva, but, as stated, to a coalescence of
the separate crystal cones to form one large lens.
The original compound eyes with their separate
cones probably form2d weak spots in the anterior
shield, and therefore gradually developed large single
lenses by the concrescence of the cones. One con-
sequence of the change is, according to our theory,
clear, and that is that the retinulae, being no
longer grouped round crystal cones, are, as reti-
nitlce, comparatively useless. We turned, therefore,
with great interest to Lankester and Bourne's
account of these retinulse, and found what we
expected, that they are by no means so definite as

204 THE APODID^ PART n

those of the lateral eyes, their irregularity suggesting
their slow disorganisation.

The very differences then which we find between
these sensory organs in Limulus and Apus arc in
reality more confirmatory of our theory than any
exact similarity could possibly be. Similarity could
only help to establish the relationship between the
two animals. As it is, we have a sufficiently strik-
ing likeness with just those differences which arc
only to be explained by deducing both animals from
a common Annelidan ancestor, in the way described
in this book.

The alimentary canal has, as already described, the
important bend which we refer both in Limulus and
in Apus to the bending round of the whole Annelidan
body. The chitin-lined cesophageal portion is more
highly differentiated than in Apus ; its oral portion is
lengthened out posteriorly (or morphologically ante-
riorly), showing the same longitudinal folds of its
intima as we found in the oesophagus of Apus. Its
anterior portion is widened out to form the so-called
pro-vcntriculus, the chitinous folds of which are so
pronounced and differentiated that they probably help
in the trituration of food. We here have the homo-
logue of the masticatory stomach of the higher Crus-
tacea. The posterior end of this projects like a
conical crater into the mid-gut, as it does to a much
slighter extent in Apus. The mid-gut runs almost to
the end of the body, receiving in its course, on each
side, two hepatic ducts from the much branched
" livers," which fill up a large portion of the cephalo-

SECT, xii RELATION OF APUS TO CRUSTACEA 205

thorax. In Apus we have a more primitive stage,
in that the livers are still clearly little more than
digesting clivcrticula of the mid-gut, at whose branched
ends only are found the hepatic glands. In Limulus,
the glandular portion is far more pronounced, and the
diverticula themselves are diminished to bile ducts, as
is the case in the higher Crustacea.

The very difference between what we find here and
in Apus is instructive ; perhaps, from the fact of there
being two ducts on each side, we can conclude that
there were originally two or more intestinal diverti-
cula in Apus. The general form of the liver of Apus
certainly looks as if it consisted of two or more diver-
ticula run together at the places where they open into
the mid-gut. Embryologically (according to Packard^
the livers of Limulus begin as simple biliary tubes,
the branchings following later. The development of
the liver as outgrowths of the mid-gut is well shown
in Glaus' figures of the Nauplius (Figs. 39, 41).

The rest of the alimentary canal offers nothing
special for remark ; like that of Apus it has a short
rectum, the chitinous intima of which is thrown into
longitudinal folds by the musculature. It is worth
noting that, whereas the anterior half of the mid-gut
is very thin-walled, it gradually gets thicker and
more muscular as it approaches the rectum ; there is
no sharp division between the two. This is exactly
what .we found in Apus.

The circulatory system of Limulus is very highly
specialised. We do not, as already stated in Part I.,
lay much value upon it from a morphological point

2o6 THE APODID^: PART n

of view. This specialisation of the blood vascular
system in Limulus is a very good illustration of the
principles stated on p. 117. The compression of the
body of Limulus against the vault of its own shell
would lead to the development of special vessels to
supply those parts which, because of compression,
would not otherwise receive their proper share of
blood. Thus we may consider the circulatory vascular
system of Limulus either as a modification of that of
the original Crustacean-Annelid, or as secondarily
acquired. The latter view is more probably the
correct one. In the first place, the type of the system
is hardly that of an Annelid, and in the second place,
the arterial envelopes surrounding the nerves are
clearly secondary specialisations in adaptation to the
peculiar physiological needs of the animal.

The Anneliclan character of the long dorsal vessel
with eight pairs of ostia needs no special notice ; it
speaks for itself in showing that at least in this respect
Limulus is not so far removed from the Annelids as
its highly specialised form would have led us at first
sight to imagine.

T/ie genital organs in Limulus are considerably
more specialised than in Apus. The comparative
shortness and flatness of the body hinders the primi-
tive metameric arrangement which we find in the latter
animal. The eggs appear to develop towards the
lumen of the gland instead of outwards towards the
body cavity. This advance on Apus is what we should
expect from the compression of the whole body, and
the consequent diminution of the body cavity.

SECT. XIT RELATION OF APUS TO CRUSTACEA 207

The spermatozoa are filiform as in the carnivorous
Annelids, but this fact is of no great morphological
importance. The genital aperture is situated on the
posterior face of the operculum, i.e. on the second
trunk limb ; in Apus it is between the tenth and
eleventh trunk feet. There were originally nephridial
openings between the limbs of all the more developed
trunk segments ; hence this difference between Limulus
and Apus is of no importance.

Development. — We have already pointed out that
the absence of the Nauplius stage in Limulus is no real
difficulty. We should only expect a Nauplius stage
in Limulus inasmuch as the Nauplius is the larva of
the original Crustacean-Annelid. The great speciali-
sation of Limulus, apparently so unlike its Annelidan
ancestor, readily explains its direct development
without passing through any such stage. Its meta-
morphoses are all passed through within the egg ;
we thus learn nothing of its early ancestors. Its
so-called " Trilobite stage " receives, however, a new
interest from our theory, which includes the Trilobite
also among the descendants of the same bent Annelid.

We conclude, then, from the comparison between
Apus and Limulus that both animals have developed
from the same bent Crustacean-Annelid ; hence the
similarity in their organisation. Although their further
development has travelled along slightly different
lines, their striking differences are in most cases easily
explained by the one having retained more primitive
Annelidan characteristics than the other.

THE APODID^: PART n

Returning to the subject of the shield, while in
Apus the dorsal integument of the fifth segment
developed a large shell fold, we see no need for
believing that in Limulus there was ever a dorsal
shield projecting backwards as a fold. When we
come to consider the Trilobites we shall find reason
to believe that the frontal ridge was in all these
animals older than the dorsal shield, and had a dif-
ferent origin, the dorsal shield itself being a later
development. In Limulus, as in many Trilobites, the
ridge round the front of the head is produced back-
wards on each side to form two horn-like processes.
But we reserve the further discussion of this most
interesting subject for the next section, where it will
be more in its place, as in the Trilobites almost every
possible variation of the same essential type of cepha-
lothoracic shield is found, for the defence of the
anterior bent, and therefore exposed, segments. We
shall then see some reasons for concluding that only
those primitive Crustacea which developed shields, i.e.
either dorsal folds like the Apodidae, or bivalve shells
like the Ostracoda, survived, Limulus being probably
the only exception to this rule. In many modern
Crustacea, however, these shields have again second-
arily disappeared.

SECTION XIII
THE TRILOBITES

Ix this appeal to the ancient Crustacean forms to
ascertain whether they lend any support to our theory
of the origin of the class, we began with Limulus, not
because it is more nearly related either to Apus or to
our bent Annelid than are the Trilobites, but because
its anatomy is so well known that it admitted of closer
comparison, and further because its relation to the
Trilobites is fairly well established. It thus formed
a sort of link for the purposes of our comparison, to
connect the Apodidae with the Trilobites and the
Eurypteridae. That Limulus and the Trilobites are
closely related is now generally acknowledged.

Having shown that Limulus is, like Apus, derivable
from a bent Annelid, if we can only show that the
organisation of the Trilobites is also best explained
by attributing to it a similar origin, we shall be able
to group the Xiphosuridae, the Trilobites, and the
Apodidae for the first time in a natural system.

It is important to bear in mind that the Trilobites

P

210 THE APODIDAE PART n

are the earliest known Crustacean forms. A special
interest therefore attaches to our endeavour to prove
that they were nearly related to the Apodidae. Start-
ing from a purely morphological and anatomical
Standpoint, we endeavoured to show that Apus was a
modified Annelid, and, therefore, a primitive Crus-
tacean. Our finding that the Nauplius, or the earliest
known larval stage in Crustacea, is but a young Apus,
went far to show that our reasoning was correct. If
now we can further show that the earliest known
Crustaceans are easily connected with the Apodidae
as related forms, it seems to us that our case is
established. Such concurrent testimony from deve-
lopmental history and from palaeontology is almost
without parallel.

The relationship of the Apodidae and the Trilobites
has already been assumed by the earlier zoologists.
Burmeister,1 indeed, tried to reconstruct the Trilo-
bites on this assumption, and attributed to them the
typical Phyllopodan limbs, and described them as
swimming about in the palaeozoic seas. Although
Burmeister's reconstruction was not correct, yet his
assumption of a relationship between the two was
justified. The fact that the Apodidae have rowing
limbs does not in any way oblige us to assume that
if the Trilobites were related to the Apodidae they
must have had similar limbs. As a matter of fact we

1 Cf. " Die Organization der Trilobiten aus ihren lebenden Verwand-
ten entwickelt," and further the historical review given by Walcott in
his paper, "The Trilobite. Old and new evidence relating to its
organisation."

SECT, xin THE TRILOBITES 211

now know that the Trilobites had ambulatory limbs
(see Fig. 51). The two sorts of limbs are, as we
shall see, but different modifications of the Anne-
lidan parapodium.

The Annclidan character of the outer form of the
Trilobites is not so much disguised as at first sight it
seems to be. But for its large head-shield it might
well have passed for a flattened Annelid. Anteriorly
we have the crescent-shaped head, followed by a
variable number of movable segments, and poste-
riorly a number of more or less rudimentary seg-
ments, often fused together to form a tail-plate.

Taking the three parts separately, and deducing
them from our primitive Crustacean-Annelid, we shall
find that much light is thrown upon many hitherto
obscure points in their organisation.

(I.) The Head. — The Trilobite head is composed of
the five anterior segments of our Annelid, bent round
so that the mouth opens ventrally and faces poste-
riorly, as described for Apus. The large labrum was
originally the prostomium of the Annelid. Fig. 46
is a longitudinal section through a Trilobite, which
we had not seen till the first part of this book was in
MS., and which afforded a most unexpected confir-
mation of our argument. Anteriorly and dorsally
the bending of the soft cylindrical body gives rise to
the glabella, as the characteristic swelling in the
median line of the Trilobite head is called. This is
the convex surface of the bent Annelidan body, and
is retained only in the Trilobites. In Apus it is
completely disguised by the growing together of the

212 THE APODID^E PART n

frontal ridge and the dorsal shield, but it is always
more or less visible in the Trilobites, which formed no
such dorsal fold. Round the glabella is developed the
remarkable crescent-shaped ridge which runs round
the front of the head, such as we found in Apus as a
prolongation of the lateral edges of the shield. In the
Trilobites, this ridge is often very pronounced, form-
ing a wide margin round the head, with horns some-
times stretching back far beyond the posterior end of
the body (see Figs. 47 and 57, p. 257). The origin
of this ridge is probably to be sought in the folds

FIG. 46.— Longitudinal section through Ceraurus pleurexanthemus (after Walcott),
showing the intestinal canal and ventral membrane, and the bend in the head.
Cf. Figs, i and 2.

which would naturally arise ventrally and laterally in
the bend of the soft body ; the bend is so sudden that
we may well imagine the folds forming projecting
angles at each side like the angles formed by the
bending of an india-rubber tube. This comparison
would be almost exact if we imagine the convex
curve of the tube so stretched as not seriously to
diminish the size of its lumen, as must have been the
case in the bent Annelid to prevent compression of
the viscera. That the sides of the angle of the bend
did thus project we conclude from the position of the
second antennae both in Apus and in Limulus, where

SECT. XIII

THE TRILOBITES

213

they lie outside the longitudinal line which joins the
other limbs. The development of hard cuticular
points, and thence of thorns on such lateral projec-
tions, would be but a matter of time. From these
points also the gradual development of the ridge
round the front of the head can easily be imagined.
In some Trilobites it remains quite inconspicuous, but

FIG. 47. — Dionide formosa (Barr), showing the giabella and the gradual rudimentary
character of the posterior segments.

in others, as already stated, it projects as a great
shovel-shaped margin. We here find, then, the origin
of all forms of the Crustacean shell, which we have
deferred discussing till now ; we may summarise our
conclusions as follows :

Round these lateral projections, due to the bending
of the cylindrical bod}-, all the shapes of the Trilobite

214 THE APODID^: PART n

head-shield play. We are inclined to think that the
formation of the ridge round the front was the primi-
tive variation, because of its great use as a belt-like
shield round the unprotected head of the browsing
animal, especially if it went hand in hand with the
thickening of the cuticle of the frontal surface.
The lateral processes and the frontal ridge thus
formed the primitive head-shield of this whole group
of Annelidan-Crustacea, and every form of shell-
covering may have been developed out of this
primitive shield. As a matter of fact we find almost
every possible variation of this ground form. The
cephalothoracic shield of Limulus is one form,
due to its fusing with the two anterior trunk seg-
ments. But by far the most important of all these
variations was the development of this head-shield
backwards over the trunk to form a cover such as
that of Apus. We have already described (p. 15)
the probable origin of this shell as a fold of the
tergum of the fifth segment developed to carry thorns
for the protection of the exposed dorsal surface, the
head being bent round ventrally. A Trilobite, Acid-
aspis Dufrenoyi (Fig. 48), shows us the neck-lobe
developed into the kind of thorn-carrying fold we
had imagined. Such a fold as that possessed by
Acidaspis, if a little wider and carrying more thorns,
could very easily develop backwards over the trunk
into a shell fold, such as that possessed by the Apo-
didae, the thorn-carrying function eventually giving
way to that of forming a cover for the dorsal surface.
But this is not the only form of shell which can be

SECT, xin THE TRILOBITES 215

derived from the primitive head-shield above de-
scribed. The bivalve shells of the Ostracoda can
also be deduced from the same by the clapping
together of the two wings of the crescent-shaped
ridge against the sides of the body as illustrated in
ig- 57> P- 257- When this crescent is large, owing

FIG. 48. — Acidaspis Dufrenoyi (Barr), Upper Silurian (after Barrande, from Zittel's
Handbnch). Showing the fold of the skin carrying two prongs projecting
backwards just behind the glabella, to demonstrate the probable origin of the
dorsal shield of Apus.

to the great development of the shovel-shaped ridge
round the front of the head, the lateral folding
of these wings round the rolled-up body would yield
a bivalve shell. Another obvious method of pro-
ducing the bivalve shell is by the folding down of the
edges of a dorsal shell such as that of Apus. These

216 THE APODID/R PART n

two origins arc, however, clearly quite different ; we
shall find later that they help us greatly in under-
standing the striking difference between the Ostracoda
and the other Crustacea possessing bivalve shells.

In the meanwhile the development of shells directly
from the primitive head-shield seems to have taken
place in two directions.

(1) By the development of the posterior edge of a
neck-lobe, or dorsal fold of the fifth segment, at first
carrying the thorns, as shown in Fig. 48, and later
forming a covering for the back.

(2) By the growth and folding down of the wings
or horns of the crescent-shaped head-shield against
the sides of the body.

To these two shell formations wre shall however
return in discussing the probable origins of the
modern Crustacea. We shall also have again to refer
to the importance of the formation of such shields
protecting the whole body, and to the advantages
which they offered over all the other variations of the
head-shield.

We conclude then that the shield of Apus was not
the primitive formation ; the ground type was, we
think, the head-shield, every variety of which we find
in the Trilobites. Thus although, in Apus, we spoke
of the ridge round the head being the prolongation
of the lateral edges of the shield, strictly speaking the
ridge and the shield were two independent develop-
ments of the primitive Trilobitan head-shield, the
former starting forwards from the lateral projections
necessitated by the bending of the cylindrical body,

SECT. XTII THE TRILOBITES 217

the latter the posterior development of the neck-lobe
as shown in Fig. 48.

(II.) T/ie Trunk Segments. — The greatest difficulty
in homologising these segments with Annelidan seg-
ments is that we find the crescent-shaped head
followed by segments repeating, in their pleura, the
form of the head, whereas at first sight we should
expect the head to be followed by a row of Annelidan
segments as in Apus, i.e. a continuation of the gla-
bella alone, as was no doubt originally the case. The
gradual acquisition on the part of the trunk segments
of their highly developed pleura repeating the charac-
teristics of the wings of the head is probably to be
explained as follows: — As soon as the typical Trilobite
head-shield became an important factor in the struggle
of each species for existence, it would tend to appear
earlier and earlier in the larva ; the Trilobite Nauplius
would then be little more than a generalised Trilobite
head with an anal segment. Between these two parts
the segments were gradually differentiated, so that
the characteristics of the head might very well make
themselves felt in the development of the segments,
and in this way spread gradually backwards to the
posterior end of the body. This, indeed, we find to
be the rule in many Trilobites ; the most specialised
segments are immediately behind the head, while
posteriorly they are more and more simple. In this
way then the segments of the Annelidan trunk were
gradually provided with the pleura characteristic of
the Trilobites ; their pleura being segmental repe-
titions of the lateral projections of the head-shield.

218 THE APODID^: PART n

The variation in the number of trunk segments is
also a point of no small interest While some authors
have tried to classify the Trilobitcs according to the
number of the trunk segments, Barrande has shown
that even within the same genus the number is quite
inconstant, the different species varying greatly in this
respect, in Olenus 9-15, Cyphaspis 10-17, &c. This
is exactly what we find in the Apodidae, where the
number of segments varies greatly : from 60-65 m
A. cancriformis, to 40 in L. glacialis. We have already
discussed the importance of this inconstant number of
the segments in our argument that the Apodidae stand
half way between the Crustacea, with their small con-
stant number of segments, and the Annelida with
their large inconstant number. But the argument has
not the same weight here as it had in our endeavour
to show that the Apodidae were very primitive
Crustacea, because in the case of the Trilobites the
fact is already apparent from their geological
position ; still it is an important characteristic which
they have in common with the Apodidae, and as
such is so much positive evidence in favour of our
argument that both are derived from the bent
Annelid.

(III.) The Pygidium is a more or less constant
characteristic of the Trilobites. It is the posterior
region of the body, composed of a varying number of
segments fused together, so that the whole region
forms a stiff plate, a sort of tail-shield answering to
the anterior head-shield. The morphology of this
pygidium has been as little understood as that of the

SECT, xin THE TRILOBITES 219

posterior end of the body of Apus ; our explanation
of the one also explains the other.

\Ye find, in fact, almost the same as we find in Apus,
that the posterior segments remain in an undeveloped
or larval condition ; although the gradual tapering
away and diminution in length of the segments is not
visible in all species, yet where it is no longer visible
it must be assumed to have secondarily disappeared.
In some cases these rudimentary segments develop
sufficiently to hinge upon one another and to bend
in the sagittal plane, or perhaps the bending may
have been effected as in Apus by the development
of rings which do not correspond with true seg-
ments. In very many cases, however, the segments
are so rudimentary that they are unable to bend upon
one another, and hence together form the stiff plate
under discussion — the pygidium (see Fig. 50). We
thus deduce the pygidium not strictly from fused
segments but from segments too rudimentary to bend
upon one another.

It has been noticed as a somewhat remarkable fact
that the trunk segments appear after the pygidium,
the young larva consisting of the head and the pygi-
dium, and between these two the thoracic segments
are gradually interposed. This is a most interesting
case of the shifting back on to the larva of important
characteristics. The pygidium, being probably useful
in the rolling up of the larva, is thus very early deve-
loped, and is then analogous to the anal segment in
the Trochophora larva, although morphologically it
is composed of a number of rudimentary segments.

220

THE APODID^E

PART II

We have, as has already been pointed out by many
authors, the parallel case of the Zoaea,- in which the
abdomen which is useful to the larva for swimming is
developed before the posterior thoracic segments.

FIG. 49. — Asaphos megistos Hall (after Walcott), showing the well-developed
ventral parapodia, and the gradual simplification of the limbs from before back-
wards, as in Apus. Cf. Frontispiece ; /, pygidium.

The Limulus larva, in which the same thing occurs, is
on this account called by Packard a Zoaea.

Having mentioned this habit of rolling up, we may
as well here point out that it also forms a link of
connection between the Trilobites and the Crustacean-
Annelid, it being easily explained as the perfection

SECT, xin THE TRILOBiTES 221

of a very natural action which we may safely assume
went hand in hand with the development of the
primitive head-shield already described. At the
approach of an enemy the forehead would be pressed
against the ground, the thorns, if there were any on
the posterior dorsal fold of the fifth segment, would be
somewhat erected by the bending under of the head
or humping of the back. In such simple movements
we have the first step towards rolling up.

This method of defence by rolling up is one of
considerable biological interest (see Fig.54). In one way
it is a very perfect method of defence, but in another
it is very fatal. Its perfection is clear from the periods
of geological time through which the Trilobites lived ;
its fatality in the fact that it admits of no further
development. Hence the Trilobites, at least all which
failed to develop shells, have died out, as unable to
protect themselves from new and more powerful
enemies, or from old enemies when these latter had
once learned to overcome this method of defence-
The development of shell folds, which, except in the
case of bivalve shells, are clearly inconsistent with the
habit of rolling up, render it unnecessary. They
make it possible to develop new and more plastic
methods of defence, to which we owe the preservation
and the rich and varied development of the whole
class of modern Crustacea.

T/ie Trilobite Limbs. — In spite of the great progress
which has been made in our knowledge of the limbs of

222 THE APODID^: PART n

the Trilobites, chiefly through the patient researches of
Walcott, they are still shrouded in a certain amount of
mystery. We believe that it will be found that our
derivation of the Trilobites from a bent Annelid will
throw considerable light upon the beautiful series of
sections made by Walcott, by giving a new clue to
the interpretation to be put upon them.

One difficulty, for instance, which has been found
in classifying the Trilobites with the Crustacea is the
absence of any trace of limbs (i.e. of antennae) in front
of the mouth. This, however, from our point of view
is no real difficulty. In reality the antennae of Apus
are hardly in front of the mouth but in a line with
it, and both are more or less rudimentary, from being
caught in the angle of the bend. This same bend was
equally sharp in the Trilobites (see Fig. 46). Why
may not the antennae have been in this bend, and as
rudimentary as they are in Apus ? We shall try to
answer this question in the following pages.

We have, in Walcott's restoration (see Fig. 50),
posteriorly to the labrum, three small limbs with mas-
ticatory processes, followed by a large pair of loco-
motory limbs with especially large ventral parapodia
for mastication. For reasons given above (pp. 44, 190)
we homologisc these large locomotory limbs with the
sixth pair of typical Crustacean limbs, i.e. with the
first pair of trunk limbs. The three pairs of limbs
anterior to these are therefore homologous with the
mandibles and the two pair of maxillae of the typical
Crustacean head. In front of these and behind the
labrum, we have, in Walcott's restoration (Fig. 50), a

SECT, xin THE TRILOBITES 223

space in which we think the antennae should have
been drawn. That they were present we have little
doubt, probably somewhat reduced, as in Apus, and
pointing backwards. Our reasons for thinking that
there must have been two pairs of antennae as here de-

FIG. 50. — Ventral surface of Calymene Senaria restored by Walcott (from Zittel).
Assuming that the large pair of locomotory limbs are the sixth or first trunk
limbs. The two pairs of antennae are missing — they should probably be drawn
in on each side of the prostomium projecting backwards, as in Apus.

scribed, are two, apart, that is, from the general reasons
founded upon our theory of their relationship to Apus
through common descent from a bent Annelid.

(i.) Figs. 51 and 52 are sections passing through
the prostomium (labrum or hypostoma) of two Trilo-

224 THE APODID^E PART n

bites. In the second of these they are cut through
along the line shown in Fig. 53. These transverse
sections through the head and labrum certainly seem
to indicate the presence of such antennae as we have
described, at least they seem to show that there were
appendages of some sort starting out sideways from
each side of the labrum, just as in Apus. It is
perhaps possible to interpret all these fragments of
limbs shown in the sections, both those seen springing
from the sides of the labrum, and those scattered about
the section, as parts of the limbs of the hind-body,

FIG. 51. — Sections through Ceraurus pleurexanthemus (after Walcott) passing through
the prostomium, showing traces of limbs springing out from each side of the
same, which we assume to be homologous with the antennae of Apus ; the
fragments of limbs at the sides may be those of trunk limbs brought near the
mouth by the rolling up of the animal.

which when the animal is rolled up are naturally
brought up to the mouth. This, however, does not
seem to be so probable as our supposition, founded
upon a comparison with Apus, that those actually
starting from the sides of the prostomium are traces
of true antennae, because :

(2.) We think that, if the place assigned by
Walcott to the three posterior head limbs is correct,
some form of antenna must have been present, if not as
antennae then as mouth parts of some kind. Accord-
ing to our theory, one of the chief advantages of
the bending round of the anterior segments was the

SECT, xni THE TRILOBITES 225

possibility of using the parapodia as instruments for
pushing food into the mouth ; and indeed, whether
our theory is correct or not, we doubt if any case
will be found of a Crustacean mouth without limbs
as mouth parts closely bordering it. \Yalcott's restora-
tion, given in Fig. 50, is therefore so far incomplete.
The mouth, which is covered by the large labrum, must
have had some kind of appendages bordering it on
each side. When therefore we find clear traces of such

Fir,. 52.— Sections of Calymene Senaria (after Walcott) passing through the prosto-
mium, showing traces of limbs springing out from each side of the same, which
we assume to be homologous with the antennae of Apus ; the fragments of limbs
at the sides may be those of trunk limbs brought near the mouth by the rolling
up of the animal. The section passes along the line shown in the next
figure.

limbs in the sections (Figs. 51, 52), we think we are
justified in claiming them as such.

It will no doubt be objected that these two reasons
are only sufficient to show that there were limbs as
mouth parts on each side of the mouth, near the
labrum, but not that they were the homologues of the
Crustacean antennae. This homology depends, it is
true, upon the truth of our main argument that the
Trilobites, like Apus, were originally bent Annelids,

226 THE APODID^: PART n

and further, upon our homology of the large loco-
motory limbs with the first trunk limbs ; to this
latter point we shall return. In the meantime
we assume that these two reasons, taken together
with our whole argument, are sufficient to establish
the fact that the Trilobites possessed two pairs of
antennae like the Apodidae and the typical Crus-
tacea.

FIG. 53.— Rolled-up specimen of Calymene Senaria (after Walcott) ; the line through
the head is the line of the sections in Fig. 52.

When now we come to ask how these antennae
were developed, we can only conjecture that in some
way or other they must have supplied the opening of
the oesophagus with jaws, or perhaps with simpler
instruments for pushing in food. It is improbable
that the first antennae should develop their ventral
parapodia as jaws, firstly because it is almost certain
that the original Annelid-Crustacean had already lost
all traces of the parapodia of the first segment, the

SECT, xin THE TRILOBITES 227

first antennae being simply sensory cirri ; and secondly
because in no other group of early Crustaceans do
the anterior antennae show any traces of ventral para-
podia as masticatory ridges. In Eurypterus, where
the method of life we attribute to the animal would
certainly have developed them into jaws had it been
possible, they almost entirely disappear. On the other
hand, we have examples of the second antennae
developing their ventral parapodia as masticatory
ridges, not only in Limulus but also in Eurypterus.

We are thus disposed to complete Walcott's resto-
ration by adding a small pair of anterior antennae
on each side of the labrum, and a pair of posterior
antennae, developing, probably as their most important
part, a pair of jaws strong enough, if not to crush and
destroy, at least to push food into the opening of the
oesophagus ; whether the sensory part was developed
or not is not so easy to decide.

It may be noticed that it was not so necessary for the
Trilobites to have large crushing jaws under the
labrum, as the masticatory ridges of the first trunk
limbs were, as in the Eurypteridae, highly developed
to function as chief mandibles. The crushed food
would have to be forwarded towards the opening
of the oesophagus, and then pushed in by special
appendages at the sides of the opening. This
point is almost as interesting from a biological as
from a morphological point of view. We have
already had two entirely different combinations of
head appendages as jaws. In Apus, the third and
fourth head limbs form the mandibles (or chief

Q 2

228 THE APODID^E TART n

jaws) and maxilla?. In Limulus we have five pairs
of nearly equally important jaws, on the four last
head, and first trunk, limbs. In the Trilobites we
find the mandibles, or chief jaws, between the
first trunk limbs, and masticatory ridges for pushing
the food into the mouth, as in Limulus, on the four
posterior head limbs. In the Eurypteridae \ve shall
find further combinations. We may perhaps find
in these different attempts to develop the best
arrangements of mouth parts almost as important a
factor in the development of the class of the Crustacea
as we think we have found in the development of the
shield. There can be no doubt that while it offered
some advantage to use the ventral parapodia of the
most powerful limbs as jaws, this must have been
attended by certain disadvantages. To this important
subject we shall return.

. We repeat here what we said on p. 43, that we might
with some safety establish a rule that the closer the
forehead was pressed against the ground the less likely
would the antennae be to function as antennae ; they
might cither degenerate as they have done in Apus,
and, according to Wralcott's restoration, in the Trilo-
bites, or they might function as seizing organs or mouth
parts, as in Limulus. We shall have occasion later to
see the converse of this rule, and shall find that the
raising of the head leads not only to the further
development and pointing forwards of the antennae
as sensory organs, but also to the travelling of the
antennae themselves towards the anterior end of the
body, an advantage for the animals which has enabled

SECT, xin THE TRILOBITES 229

them to hold their own to-day, whereas Limulus and
Apus are probably the only surviving Crustacea which
retain the original position of the Annelidan antennae.

As above pointed out, our interpretation of the head
limbs of the Trilobites rests largely upon our homo-
logising the large locomotory limbs with the sixth
pair of Annelidan parapodia, or with the first trunk
limbs of the Crustacea. Our adoption of this large
locomotory limb throughout all the primitive Crus-
tacea as the first trunk limb, for reasons given p. 44,
receives some support from Walcott's restoration,
where it lies behind the line which runs from side to
side, through the widest part of the head, which is
morphologically the line round which the body bent.
\Yc have already seen, further, that Limulus — and we
shall see that the Eurypteridae, with some exceptions —
not only possessed the two pairs of antennae, but also
the large locomotory limbs as the sixth pair, i.e.
the first pair of trunk limbs.

It should be mentioned that so far as these con-
clusions are based on the few sections published
in Mr. Walcott's paper, his conclusions are undoubt-
edly of much greater value than ours, inasmuch as
they were based upon a much more extended study
of sections, and of the whole Trilobite problem. We
have, however, to set, as against this, our claim to have
found in Apus a key to the true understanding of
the morphology of all these primitive Crustacea.

The form of the trunk limbs in the Trilobites does
not at first sight admit of any close comparison with

230

THE APODID/E

PART 11

those of either Apus or Limulus, but by closer study,
and by referring them back to the original Annelidan
parapodia, their common origin becomes evident.
We find the limbs much specialised, the habits of
life of the animal leading to certain modifications.
In the first place, the creeping motion along the
ground required the development of legs. In the
second place, the habit of rolling up requires that
the limbs should take up as little room as possible,

FIG. 54. — Restored transversed section through Calymene Senaria (after Walcott),
showing the spiral gills, the exopodite (= the sensory cirrus of the dorsal para-
podium), the endopodite, or ambulatory foot (= the dorsal parapodium), and the
thigh piece or coxal joint, the ventral projection of which corresponds with the
ventral parapodium, cf. Fig. 49.

first, to render the rolled-up attitude mechanically
possible, and second, in order that as much of the
respiratory medium as possible may be enclosed.
The special form of the limbs can thus be under-
stood. We find (Fig. 54) a large basal joint, the
inner ventral part of which is almost certainly to be
homologised with the ventral parapodium of the
Annelid. This again originally functioned as a
gnathobase or accessory jaw for the holding and
forwarding of food to the mouth, i.e. on a certain

SECT, xni THE TRILOBITES 231

number of limbs not too far from the mouth. It
was, no doubt, as in Apus, much reduced in other
parts of the body, in order not to occupy much
space.

The ambulatory limb, if our homology is correct,
was the prolonged tip of the dorsal parapodium, and
thus homologous with the endopodite of other Crus-
tacea. The exopodite was the sensory cirrus ; and
here, no doubt in correspondence with the needs of
the animal, it retained its position close to the gills,
and its filiform shape ; it did not travel along the
prolonged dorsal branch of the parapodium, or
develop into a rowing flabellum as in Apus. The
modification of the gills into spirals, &c., is also very
easily explained on the grounds given above. The
animal required respiratory organs which afforded as
large a respiratory surface as possible while occupy-
ing the smallest possible space, such respiratory
organs being essential to the habit of rolling up.

Before dismissing the subject of the form of the
Trilobite limbs, we wish to return for a moment to
Burmeister's assumption, that if the Trilobites were
related to Apus they must have possessed Phyllopo-
dan limbs. This, however, is by no means necessary.
What is generally known as the typical Phyllopodan
limb is but one of the ways in which the Annclidan
parapodium developed, the Trilobite ambulatory leg
being another and quite independent modification
the parapodial type being visible in both. The
modifications are due to adaptations to the different
manners of life adopted by the different groups.

232 THE APOD ID/1-; PART n

The development of the ventral parapodium into
mandibles and gnathobases is a common specialisa-
tion in all the groups, this being the most primitive
modification according to our deduction of the Crus-
tacea from a carnivorous Annelid, which caught prey
between its ventral parapodia and forwarded it on into
the mouth, bent round to receive it. The dorsal para-
podia, being chiefly used for locomotion, have how-
ever been differently developed according to the
different methods of locomotion adopted. In Apus
they are specialised as rowing plates (except a few
anteriorly for raking prey together), in the Trilobites
as ambulatory legs. The former modification requires
no description ; it results simply in a further develop-
ment of the flat leaf-shaped parapodia, the sensory
cirrus alone, perhaps, requiring to change its form
from a cirrus into the flat flabellum. The ambulatory
leg of the Trilobitc may be supposed to have arisen
as follows. A strip running from the tip of the
parapodium, where it rested on the ground, to the
body, would tend to be strengthened, and would
eventually bear the weight of its share of the
body. On each side of this strip the leaf-like para-
podium would be useless, and would gradually dis-
appear, this disappearance being accelerated in the
Trilobites by other and special causes which we have
already described, such as the necessity of having limbs
which, in the rollcd-up body, would occupy as little
space as possible. Thus we may safely assume that
the parapodia, if used for walking or crawling, would,
by a simple biological law, turn into ambulatory legs.

SECT, xin THE TRILOBITES 233

The question as to whether the leaf-like feet per-
sisted at the hinder end of the body is an interesting
one. We have no certain data on the subject, but,
from our point of view, we do not think it at all
probable. We have seen that in Apus even the most
rudimentary limb repeats the Phyllopodan type. We
are also inclined to believe that the more rudimentary
Trilobite limbs would naturally repeat the Trilobite
ambulatory type. The presence of flat leaf-shaped
limbs in the Eurypterida^ and Limulus, accompanied
by highly specialised anterior limbs, may perhaps be
used as an argument in favour of their presence in
the Trilobites also. On the other hand the highly
developed gills on the trunk limbs of the Trilobites
rendered it unnecessary to concentrate respiration on
a few broad gills at the posterior end of the body as
in Eurypterus and Limulus, which in this respect
compare with some modern Isopoda.

The first trunk limb, according to Walcott's
restoration, has both its locomotory dorsal branch
and its masticatory ventral branch specially strongly
developed (see Fig. 50). It is, in some respects, very
natural that the masticatory ridge of a powerful
locomotory limb, if it possessed any function at all,
should gradually come to be the chief jaw, as we
shall see to have been the case also in the Eury-
pteridae ; the disadvantages of this arrangement will,
however, be pointed out later.

We have already shown why the first trunk limb,
being the parapodium of the first free segment, not
taken up in the formation of the head, should be

234 THE APODID^: PART n

highly developed. The use of such a specialised
limb in the Trilobites, however, is difficult at first
sight to see. In Apus we find it developed as a
sensory organ on the principle of the division of
labour. In the Trilobites it is clearly locomotory, and
as such seems rather out of place among the smaller
and less powerful crawling legs of the other trunk
segments. In discussing the manner of life of Eury-
pterus and Pterygotus, we shall find that they throw
some light on the probable use of this limb in the
Trilobites.

It is especially interesting to find the gradual
simplification of the limbs from front to back,
which is evident towards the posterior end of the
body (Fig. 49, cf. with the Frontispiece). There can
hardly be any doubt that the gradual dwindling of
the limbs in the Trilobites admits of the same
explanation as a similar dwindling of the limbs in
Apus. Such a singular morphological occurrence,
in two animals so like in other respects also, can
hardly be a case of analogy.

TJie Eyes. — Packard has shown that the hard part
of the eyes of Trilobites, which alone have been pre-
served in the fossils, are identical with those of Limu-
lus. As we have already seen in discussing the eyes
of Apus, we consider the eye of Limulus as a more
primitive stage in the development of the Crustacean
eye out of the Annelidan eye-spots. In this respect
Apus is more highly developed than both Limulus
and the Trilobites, as indeed we should expect from
its free-swimming life.

SECT, xin THE TRILOBITES 235

The Alimentary Canal has already been referred to.
It has the very pronounced bend on which we lay so
much importance (see Fig. 46). Although we think
our proof is not much weakened by our not finding
any traces of the sternal plate, still it would be
interesting if it were to be found, as it must without
doubt have been there, i.e. if there is any truth of our
deduction of these animals from bent Annelids. The
habit of rolling up would lead to a strong development
of the ventral muscle bands, and consequently of this
sinewy mass for their attachment (cf. p. 261).

\Ye think, then, that we have here made it highly
probable that if our deduction of Apus from a bent
carnivorous Annelid holds, the Trilobites must have
had the same origin. This fact, that the most primi-
tive Crustacean known to the palaeontologist should
show so many points in its organisation directly
deducible from the Annelids, i.e. deducible after the
Apodidae have supplied us with the key to their cor-
rect interpretation, is one of those confirmations of a
theory which we think amounts almost to a demon-
stration.

The Trilobites, then, are nothing but specialised
carnivorous Annelids, browsing under cover of the
dorsal integument, which, starting from the head-
shield, gradually spread out like a flattened jointed
roof, covering all the segments. Every imaginable
variation in the sculpture of the surface of this roof,
and in the thorns for its protection, are to be met
with in the Trilobites.

236 THE APODID^ PART ii

This development of great multitudes of armoured
browsing carnivorous Annelids in the palaeozoic seas,
supplies us with abundant matter for biological specu-
lation. It was perhaps in defence against these
powerful marauders that so many Ccelenterata per-
fected their nematocysts or stinging cells, that 'the
Corals built their stony ramparts, and that many of
the Mollusca developed their shells. It may indeed
have been the perfection of these defences which
led to the dying out (with the exception of Limulus)
of these early Crustacea, especially of the giant forms.
Whatever the cause, all except Limulus, the Ostra-
coda, and the Apodidse (looked upon as the
racial form of all other existing Crustacea) gradually
died out.

The first and the last of these still fortunately retain
the clearest traces of their origin, and, more or less
modified, the browsing habit of life.

SECTION XIV

THE EURYPTERID^E

THIS last group of the Gigantostraca need not
detain us long. By the general consent of all the
zoologists who have recently studied these animals,
they are classed with the Xiphosuridae and the
Trilobites. The exact relationship, however, has not
hitherto been very clear ; we now find it in their
common origin from our Crustacean-Annelid.

We have imagined our Crustacean-Annelid develop-
ing first of all a kind of crescent-shaped protection for its
bent head, arising primarily from the lateral projections
due to the bending of the cylindrical body. This
shield develops in almost every possible way. In the
Apodidae it forms a dorsal fold to cover the rest of
the cylindrical and unprotected Annelidan body ; in
the Ostracoda it forms the bivalve shell in a way to
be described later, or it gives rise, as described on p.
217, to the flat jointed dorsal roof extending over
the whole body in the Trilobites and the Xiphosuridae,

238 THE APODIDyE PART n

In the Eurypteridae, however, even this primitive
head-shield seems wholly or almost wholly to have
disappeared, and the flattened Annelidan segments
relied almost entirely upon the stronger development
of the exoskeleton for protection. Like the Xiphos-
uridae, they developed comparatively few segments,
ending in a caudal spine or plate. In this limited
number of segments they show considerable special-
isation. The whole structure of the animal is clearly
adapted for a free-swimming life, the first trunk limbs
forming powerful oars.

The limbs develop as Crustacean limbs only on the
head and first trunk segment ; on the other trunk seg-
ments they remain leaf-shaped, i.e. more like the
original Annelidan parapodia. The gill portions of
these limbs may have had their surfaces increased by
means of numerous integumental folds like the leaves
of a book, as in the Xiphosuridae.

We feel some confidence in the following homology
of the head limbs, because we have learned, from all
the groups hitherto discussed, that the large rowing
limb is probably the first trunk limb ; we need not
here repeat the reasons already given for this conclu-
sion. All that lies in front of these large rowing
limbs therefore represents the head.

Before, however, attempting to examine the parts
in detail, we are at once struck by the difference
between the heads of these animals and those of the
Apodidse, Limulus and the Trilobites. The mouth
parts are in fact so specialised that it is not easy to
compare them with those of the above-named groups.

SECT. XIV

THE EURYPTERID^

239

The head limbs are different in the two groups,
Pterygotus and Eurypterus (see Figs. 55 and 56).
They arc, however, only different modifications in

FIG 55. — Pterygotus Osiliensis, upper Silurian, after F. Schmidt (from Zittell), show-
ing five pairs of cephalic limbs, the enormously developed first pair of antennae, and
first trunk limbs in which the dorsal and ventral parapodiaare greatly developed
as locomotory and masticatory limbs respectively. The second pair of cephalic
limbs sometimes disappear, as in Pterygotus Anglicus Agassiz.

adaptation to slight differences in the manner of
life. Judging from the forms of these remarkable
animals, we think the following method of explain-

240 THE APODIDyE PART u

ing the modifications they show will not be far
wrong.

The earliest Crustacean-Annelids possessed large
labra or prostomia projecting backwards, still retained
in the Apodidae and Trilobitcs. This labrum almost
necessitated a very deliberate manner of browsing.
The animal would creep along, and would have to run
some way over its food before it could get it into its
mouth, the whole process, it seems to us, necessitating a
number of small movements backwards and forwards.
Small living prey would very often escape, owing
to the fact that the animal's mouth and jaws were
not ready in position for them when first perceived.
The labrum necessitates the animal passing forwards
over its prey, then darting backwards to follow it with
its jaws. We here see how useful the gnathobases of
Apus must be in catching and holding prey which
has been thus passed over. Indeed the whole arrange-
ment of the limbs of Apus with the sensory endites,
forms an excellent trap to catch prey over which the
labrum has passed. The legs and pleura of the
Trilobites, and the large vaulted shield of the Xiphos-
uridae may serve the same purpose, although in the
latter case the labrum is much modified. In this re-
spect, however, the Trilobites were not so well equipped
as are the Apodidae ; hence perhaps the development of
the large locomotory limb, which enabled the animal
to dart backwards after prey thus run over, with great
rapidity. We here see the use of the two kinds of
limbs figured in Walcott's restoration, ambulatory
crawling limbs for slow and deliberate forward move-

SECT, xiv THE EURYPTERID^E 241

ments, and one pair of springing limbs for short
sudden dartings backwards.

It is clear, then, that the possession of these large
labra was attended with certain disadvantages in
feeding. It is therefore not improbable that some
of these primitive Crustaceans should show various
modifications. Smaller upper lips being an advantage,
the labra might almost disappear, so that the opening
of the mouth would be ready for its prey as soon as
it came in a line with it.1 A natural concomitant
change in the under lips would also take place ;
they would develop into the large metastomata found
in the Euryptcridae, which clearly helped to prevent
prey slipping past the mouth as the animal darted
forwards. The more rapid the forward dart after
prey, the larger should the metastoma be ; otherwise
prey once shot over would be almost sure to escape
before the animal could turn round ; the animals
have no trap-like arrangement of trunk limbs in
which prey could be caught. We do not, it is true,
find from comparing Figs. 55 and 56 that the larger
rowing limb is accompanied by the larger lower lip,
still we think the above reasoning to be correct, and
that other factors, such as the higher development of
the sensory organs, compensate in this case for the

1 On p. 40 we discussed the origin of the division in the under lip of
Apus so that it should not form a barrier to the pushing of food forwards
into the mouth. We now see that the divided upper lip of some Trilobites
there referred to (and well illustrated Fig. 49, p. 220), was also probably
intended to shorten the way into the mouth, only in this case round the
labrum from in front. The thi'ee small pairs of posterior cephalic limbs
may have assisted in this latter process.

R

242 THE APODID^£ PART u

comparative smaliness of the metastoma. Thus then
the entrance to the mouth may have come to have
almost an anterior-ventral instead of a posterior-
ventral aspect. This explains the enormous man-
dibles developed by the ventral parapodia of the
first trunk limb. In Pterygotus, Fig. 55, we have, in
fact, an arrangement almost exactly the opposite of
that found in the other primitive Crustaceans ; the
under lip forms the analogue of the upper lip,
the masticatory ridges of the first trunk limbs arc
analogous to the mandibles, while those of the four
posterior head limbs probably function as maxillae,
their dorsal parapodia doubtless helping in the catch-
ing and holding of prey. And lastly, the first antennae
developed into large chelate feet. It is almost as if we
had the typical mouth formula of a modern Crustacean
turned quite round.

These changes clearly went hand in hand with the
acquisition of more rapid motion in feeding. A spring-
ing or darting movement forward is most suitable
for an arrangement of mouth and jaws facing ante-
riorly, for the sudden seizure of the prey which
comes in the way. Further, it seemed to us that
the more rapid the movement the more delicate
should be the sensory organs for the rapid percep-
tion of what was food and what was not. A com-
parison of the rowing limbs of Eurypterus and
Pterygotus quite confirmed this supposition, and lent
unexpected support to this method of explaining
the morphology of these animals. Eurypterus (Fig.
56), which has all its anterior head limbs developed as

SECT, xiv THE EURYPTERID^: 243

highly sensitive antennae, has larger rowing limbs in
proportion to the size of the body than Pterygotus
(Fig. 55), which does not seem to be so well provided
with such organs. The latter animal moved more slowly
and caught its prey with its powerful pincers. The
former darted forward with great rapidity and caught
its prey at once between its numerous jaws.

We consider then the Xiphosuridae as early Trilobites
specialised for slow deliberate browsing ; the Eury-
pteridae on the contrary for a rapid darting method of
capturing prey. That the Trilobites did employ the
springing movement which we have here assumed
purely on morphological and biological grounds, has
been lately confirmed by the discovery of a Trilobite
track, which, according to Ringueberg the discoverer,
could only have been produced by a series of
jumps.1

This description of the manner of life of these
animals (the Eurypteridae) renders it not so necessary
to describe the limbs of the two animals ; still, as there
are points of great interest in their morphology, a
short account of them will not be out of place.

Taking Pterygotus first, we have the first antennae
developed into long chelate seizing feet, like the first
antennae of Limulus, but much more highly developed.
The analogy of the Scorpionidae will at once suggest
itself, where for the same purpose the palps have
developed in the same way. These chelae of Ptery-
gotus were probably richly provided with sensory
hairs, since the limb on which they were developed

1 Proc. American Association, 1886.

R 2

244 THE APODID.^E PART n

was, as a sensory limb, richly innervated. The eyes
also seem to have been highly developed.

The following four limbs, which correspond with
the second antennae, mandibles, and first and second
maxillae of Apus and of the other Crustacea, resemble
the ordinary Trilobite limbs. Their dorsal branches
probably functioned as palps or tasters, as perhaps
was the case in the Trilobites, or perhaps as limbs
for holding prey brought by the chelae in the right
position for the mandibles to crush, just as the fore
legs of a caterpillar hold the leaf in the best position
for the jaws to work upon it, only in this latter case,
of course, the jaws lie in front of the legs instead of
behind them.

The masticatory ridges of these four limbs probably-
functioned as maxillae, but, as already mentioned, lying
anteriorly to the mandibles, not posteriorly as in all
modern Crustacea.

The first trunk limbs have already been mentioned
as large rowing limbs. It was in one sense natural
that the powerful limb should also develop a powerful
ventral parapodium functional as a jaw, but the union
of the two functions is not easily comprehensible, and
we are more than ever inclined to think that the two
may have been separately articulated with the body.

The limbs of Eurypterus differ markedly from those
of Pterygotus. In front of the large rowing limb, i.e.
the first trunk limb, we have only four limbs visible in
the figure, all of these appearing to be sensory, and
thus affording a striking contrast to the head limbs of
Pterygotus, none of which appear, at first sight, to be

SECT. XIV

THE EURYPTERID^:

245

sensory. From our point of view, according to which
the large rowing limbs belong to the first trunk seg-
ment, we should have had to conclude that one pair of
limbs had disappeared. Such a supposition is however
not necessary, as F. Schmidt has found and described

FIG. 56. — Eurypterus Fischeri Eichw. : Upper Silurian, natural size, after F. Schmidt
(from Zittel's Handbnch der Paheontologie). Between the first pair of feet,
Schmidt found a fine pair of feelers, corresponding with the Antennules of the
other Crustacea.

a pair of rudimentary antennae between the first pair,
so that Eurypterus possesses the typical number of
head appendages. It is a fact generally accepted that
the pair of large rowing limbs corresponds with that
of the sixth segment. There is, however, no general
agreement as to whether these first six segments form

246 THE APODID^: PART II

a head or a cephalothorax. Our homology of the large
limb with the first trunk limb, throughout all these
primitive Crustacea, shows that the six segments of
the Eurypteridae form a cephalothorax, and not only
a head.

This degeneration of the anterior antennae in
Eurypterus is hardly what we should have expected
theoretically. The rapid forward movement for feed-
ing would seem to require highly developed antennae
pointing forwards. We attribute it to the fact that
the manner of life of the animal, as above described,
required that the sense of touch in a limb should be
immediately followed by an act of seizing, by means
of its masticatory ridges. The anterior antennae had,
however, entirely lost the power of developing their
parapodia even in the original Crustacean-Annelid,
and thus became of very secondary importance in the
life of Eurypterus.

As to the other limbs of the head, Eurypterus
resembles Limulus in having the masticatory ridges
on the last four head limbs and the first trunk limb well
developed, and working as jaws round the mouth, which
was apparently not the case in Pterygotus, where the
importance of the masticatory ridges of the first trunk
limb over those of the head limbs was very evident.
The exact morphology of the limbs themselves it is im-
possible to describe with certainty; it is not improbable
that those of the head, in Eurypterus, are the sensory
cirri alone of the original parapodia. We sec no reason
why this should not be the case. Nature seems to
delight in every possible variation, and indeed in the

SECT, xiv THE EURYPTERID^E 247

limbs of the modern Crustacea we have almost every
possible combination of the parts of an Annelid para-
poclium. We have, for example, the sensory cirrus
alone in the antennae, the gills alone in many Crustacea
(e.g. Caprella), the dorsal parapodia alone in the
ambulatory limbs of the Decapoda, the ventral para-
podia alone in the mandibles, and all these parts
together in the typical Phyllopodan limb. Other
combinations, such as the dorsal parapodium with the
sensory cirrus, the dorsal parapodium with the gill,
will no doubt suggest themselves to the reader.

What was said above as to the first trunk limb of
Pterygotus applies equally well to the first trunk limb
of Eurypterus. We may further add that their
form as rowing limbs is just what is required to give
the animals the forward darting movements which we
have assumed to have led to the modifications of
their mouth parts. Whether they kept up a continual
rowing motion like the common free-living Copepoda,
or lurked at the sea bottom to dart out in pursuit of
prey which happened to come within reach, it is
difficult to say ; we incline to the latter as the more
probable habit of life.

Again, as already described, the use of the large
limb in the Eurypteridae throws some light on that
made by the Trilobites of their large first trunk
limb. It functioned as a kind of springing foot to
supplement the more deliberate method of crawling.
The animal kingdom supplies us with many ex-
amples of special arrangements for such a sudden
and more energetic method of locomotion, developed

248 THE APODID.K PART H

in animals whose ordinary progression is slow and
deliberate.

In our general account of the probable manner of
life of these animals we have described the change
which we think took place in the upper and lower
lips, the former almost disappearing, while the latter
develops into a large fold projecting anteriorly, and
bearing exactly the same relation to the masticatory
ridges of the first trunk limbs as the labrum of Apus
does to the mandibles, only pointing exactly in the
opposite direction. The position of this metastoma
corresponds exactly with that of the under lips of
Limulus. This fact seems to suggest that this wa^
also the position of the under lips in the Trilobites.

The leaf-shaped abdominal limbs we have already
mentioned as undoubted links between these animals,
Limulus, and our bent Annelid.

We must now leave these highly interesting animals,
wrhich in point of size reached the highest develop-
ment of all the Crustacean descendants of our car-
nivorous Annelids. The exact relationship of the
group to the Trilobites and the Xiphosuridae, and to
one another, we cannot pretend to settle. It must be
left to those who have made the special morphology
of these fossil forms a life-long study. We must con-
fine ourselves here to the suggestion made above, that
the Xiphosuridse and Eurypteridae are early Trilobites
modified for two different and opposite methods of
feeding. We shall be more than satisfied if we have
been able to contribute something to our knowledge
of the groups, by tracing their origin to the Annelids.

SECT, xiv THE EURYPTERID^E 249

In bringing to a close these comparisons of the
fossil Crustacea with Apus and with our Crustacean-
Annelid, it may be interesting to see, set out in a
table, the various ways in which the parapodia in the
first six Annelidan segments have been developed — a
representation of the attempts of Nature to find the
best combination of head and mouth parts.

The limbs used as jaws are in larger type, so that
the different masticatory arrangements may be seen
at a glance.

A study of this table shows us that all the animals
which retained the early primitive arrangement of
crushing the food between the ventral parapodia of
the first trunk limbs, which were the strongest in the
body, have, with the exception of Limulus, died out.
It is not difficult to see that it is a great advantage to
have the mandibles as close to the opening of the
oesophagus as possible, otherwise the greater part of
the juices of the crushed animal would be lost before
it could reach its destination within the oesophagus of
its clevourer. The enormous metastoma or under lip
of the Eurypteridae may have been partly an attempt
to avoid this loss. It does not seem improbable,
therefore, that the ultimate selection of the third pair
of ventral parapodia as mandibles may have assisted
in leading to the survival of the modern Crustacea.
On the other hand, the enormous growth of some of
these ancient forms (Pterygotus anglicus sometimes
being more than a metre in length) shows that they
did not apparently suffer from lack of nourishment on
account of the arrangement of their jaws. When,

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therefore, we ask why these animals died out, in spite
of their having acquired the habit of free swimming,
we can only suggest that the very perfection of their
specialisation may have been fatal to them ; the line
of their development ended in a cul-de-sac. They
were not plastic enough to adapt themselves to some
great change or other which took place in their sur-
roundings, such as the perfection of the protective
arrangements of their prey, and consequently died
out. The theory which deduces the Arachnida from
them through the scorpions seems to us to be
very improbable in the face of this extraordinary
specialisation. But to this important and interesting
discussion we shall return in a final section dealing
with the other division of the Arthropoda, viz. the
Trachcata.

SECTION XV

ON THE NEW CLASSIFICATION OF THE CRUSTACEA
NECESSITATED BY THE THEORY

OUR work is so far finished. We endeavoured first
of all to show that Apus was easily derivable from a
bent carnivorous Annelid. If this was really the case,
we at first concluded that Apus must be the primi-
tive Crustacean. In order to test this, we appealed
to such an archaic form as Limulus, which is still
extant, and to the palaeozoic Trilobites and Eury-
pteridae. These have offered unexpected confirmation
of our theory, amounting, as we have said above, to
a demonstration. But at the same time we have had
to modify our conclusion that Apus was the primitive
Crustacean, these forms not being derivable from
Apus, but rather from the same bent Annelids.
This accounts, at the same time, for their remarkable
resemblances and for their many differences.

Besides the palaeozoic Crustacea which we have
so far mentioned, viz. the Trilobites, Xiphosuridae,

SECT, xv CLASSIFICATION OF CRUSTACEA 253

Eurypteridae, and Phyllopoda, there occur numerous
remains of Ostracoda and Cirripedia. If we can
in any way connect these latter with the above
named, we shall have solved the difficulty expressed
by Barrande and felt by many, that the Crustacea first
appear in the geological record in several widely dif-
ferent groups, almost simultaneously, and without any
transition forms either leading up to them or linking
them together. Our derivation of the former groups
from bent Annelids with no hard chitinous skeleton
which could have been preserved, explains the
sudden appearance of these groups. We have still
then to show that both the Ostracoda and the
Cirripedia are deducible from these forms. As, how-
ever, these two groups have modern representatives,
we shall treat them in order among the other living
forms.

We have then to ask the question, From which of
these primitive Crustacean forms did the modern
Crustacea arise ? For some groups, fortunately,
the answer is clear ; as to others, however, we can
only guess.

The attempt which we here make to sketch out a
new classification of the Crustacea must be understood
to be quite provisional. In establishing the bent
Annelid as the origin of the Crustacea, we have done
nothing more than lay the foundation stone for the
construction of a complete and final classification of
the Crustacea, including, for the first time, the hitherto
enigmatical palaeozoic forms. It is, however, as
completely out of the sphere of this book as it is

254 THE APODIDiE PART n

beyond the abilities of the writer to attempt to carry
this out in detail.

In arranging the Crustacean groups, we propose
to ignore the usual division into Entomostraca and
Malacostraca. The Entomostraca we take to mean
all those groups which do not clearly belong to the
natural group of the Malacostraca. We therefore
prefer to divide the class into Phyllopoda, Malacos-
traca, Copepoda, Cirripedia, and Ostracoda. From
what follows it will be seen that we divide these five
into three groups, the first consisting of the Phyllo-
poda and Malacostraca ; the second, of the Copepoda
and Cirripedia ; and the third of the Ostracoda. We
believe that the first group is derived from the
Apodidae, the second from a larval Apus, and the
third, at least partly, from a Trilobite. This group-
ing, however, requires considerable investigation
before it can be definitely accepted. We leave it to
others who have made the different groups of the
Crustacea their special field of research to carry it
out in detail. We confine ourselves here to giving a
diagram representing the way we propose to construct
a natural order of the Crustacea based upon our
theory. We can, unfortunately, offer but little in
the shape of proof of this new classification, and
must content ourselves with appending a few discon-
nected notes on the different groups, which tend to
support our views.

Taking the groups in the order in which they
occur in the diagram from left to right, we may at
once dispose of the Eurypteridae and the Xiphosuridae,

SECT, xv CLASSIFICATION OF CRUSTACEA

255

From the Trilobites, however, we are inclined to
deduce at least a part of one important group of
modern Crustacea, the Ostracoda. We think it

Li,m, Ins Ostracoda Cirri.

Apus Phyllopoda Malacostraca

Nebalidcc

Ceratiocaris

The Crttstacean Annelid

Proposed genealogy of the Crustacea. It will be seen from the text that though we
have here given only one root for the Ostracoda it is probable

had at least a twofold origin.

le that they have

probable that the Crustaceans in question may be
deduced both from Trilobites and Phyllopods. The
strong likeness between these early forms, especially in
their larval stages, now perpetuated in the Ostracoda,

256 THE APODID^E PART n

accounts for the general resemblance of the latter to
one another.

THE OSTRACODA.

These animals occur in company with the Trilobites
in the very oldest fossilifcrous strata. Balfour sug-
gested that they may have had an origin independent
of that of the other Crustacea. As, however, we find
them possessing the bent intestine, clear traces of the
entosternite, the paired and the unpaired eye, we
must, according to our theory, deduce them from our
bent Annelid.

There are two ways in which the origin of the
bivalve shell may be explained : either (i) that
shown in Fig. 57, where it arises simply by the
folding together of the horns of the crescent-shaped
ridge round the front of the head, or (2) when it arises
through the folding down of the two halves of a
dorsal shell such as that in Apus. These two
methods are quite distinct ; the former bends the
dorsal integument of the head-shield alone along the
middle line, the latter bends only the dorsal shield as
far as its junction with the body. There is, however, a
method of combining these two modifications if, after a
dorsal shield has been developed, both the head-shield
and the dorsal shield are bent.

We were at first inclined to attribute only the first
method of origin to the bivalve shells of the Ostracoda,
and to deduce them from some such form as Harpes
ungula. It would be an obvious advantage to an animal
given to the habit of rolling up for defence, to be able to

SECT, xv CLASSIFICATION OF CRUSTACEA

257

continue to feed and breathe, and yet remain rolled up
and sufficiently protected against its enemies. It is clear
that this end would hardly be attained in those cases
in which a large solid pygidium closed against the
head-shield. But on the other hand, it would be
quite possible by the longitudinal folding of the lateral
wings of the head-shield, as shown in Fig. 57 B.
\Ve may well suppose that some Trilobites adopted
this method of protecting themselves, since, besides

FIG. 57. — Harpes ungula 'Sternb. A, dorsal view; B, rolled up in profile (from
Bronn's Klassen ttml Ordnungen des Thierreichs) ; B, to show the probable
origin of the Ostracoda, the head-shield with the enormously developed frontal
fold, shown here in profile, only requires to bend in the dorsal middle line to form
a bivalve shell.

the great advantages already mentioned of allowing
the animal still to use its limbs and to move about
and feed while remaining almost perfectly enclosed,
it is also clear that the closing of such bivalve
shells, which would never be very wide open, would
be a much quicker and simpler process than the
rolling up of the whole body in the sagittal plane.
In assuming this origin for the bivalve shell of the
Ostracoda, and not that from the folding down of a

258 THE APODID^E PART ii

dorsal shell of an Apus-like animal, we have been
mainly influenced by the following considerations,
which must be admitted to be of some morphological
importance.

(i) The position of the head in the shell seems to
point decidedly to such an origin. If the shell had
been formed by the bending down of the sides of a
dorsal fold, the head would either project anteriorly
as in the Cladocera, or, if it came between the shells
at all, could only do so by itself bending round ven-

FIG. 58.— Cypris fasciata (from Bronn's Klassen und Ordnungen des Thierrdchs)
to show the position of the head in the shell for comparison with the following
figures and with Fig. 57.

trally,as shown in different stages in Figs. 60, 61, and
62, or by the growing forward of the halves of the shell
so as to cover the head ; this latter method is, for many
reasons, not a very probable one. In the Ostracoda
we find the " face " deep back in the shell, pointing-
forwards in a way difficult to explain on any other
hypothesis than that which we put forward. These
projecting parts of the shell are the lateral halves of the
shovel-shaped ridge which projected so far forwards
in the original Trilobite ancestor of the group. If we

SECT, xv CLASSIFICATION OF CRUSTACEA 259

take the Trilobite figured in Fig. 57^, and fold the
ridge round the head along the dorsal middle line, the
face (which lies under the glabella) would come to
have almost exactly the position which it has in the
Ostracoda.

(2) The ridge of the head-shield is, like the ridge round
the head of Apus, simply a fold of the integument, and
contains a part of the general body cavity. Probably
as in Apus and Limulus, it contained the hepatic

FIG. 59. — Diagrammatic transverse section of an Ostracod, showing the body cavity
continued into the valves of the shell, into which also the hepatic diverticula
penetrate. The closing muscles are seen to radiate from a central sinewy mass,
the sternal plate. /, intestine ; /, hepatic diverticula.

diverticula of the mid-gut. In a cross section through
an Ostracod the observer is at once struck by the fact
that the space between the laminae of the shell is con-
siderable, and that it is a continuation of the body
cavity. Not only do the hepatic diverticula penetrate
into it, but in some genera the genital glands also
(Fig. 59). While this is exactly what we should
expect from the bending of a head-shield with a
pronounced frontal ridge, we should hardly expect to
find it from a bending down of a dorsal fold.

26o THE APODID^: PART n

(3) We have further the fact, already mentioned,
that the Ostracoda are found among the Trilobites in
the Silurian strata, and may thus well have been a
modified Trilobite form.

It is, however, clear that these arguments do equally
well to establish a deduction of the Ostracoda from a
primitive Phyllopod with a developing dorsal shield.
We have only to assume that both the head- and
dorsal shields were bent along the dorsal middle line.
The extraordinary likeness between the shells of some
of the early Ostracoda (e.g. Leperditia) to the shells
of such Phyllopods as Ceratiocaris Salteriana make a
Phyllopodan origin for at least some of the Ostracoda
very probable. We have further satisfied ourselves
by dissections, that at least in some Ostracoda the
ligament uniting the two halves of the shell runs back-
wards posteriorly beyond the point of junction of the
abdomen with the shell. We do not, however, give
up our first impression that some of the Ostracoda are
deducible from Trilobites. In addition to such a
significant form as that given in Fig. 57$, we would
call attention to the fact that the shells of many early
Ostracoda are marked by lobes and grooves which
Barrande compared to the glabella and intervening
furrows, &c., of Trilobites. The presence also of the
" ocular " tubercle on each shell in some Ostracoda
may well signify what its name implies ; the ocular
tubercle of the original Trilobite showing just as well
on the folded, as on the flat, head-shield. The part
played by the habit of rolling up will again be referred
to.

SECT, xv CLASSIFICATION OF CRUSTACEA 261

As to the general truth of our theory that the
Ostracoda are little more than folded Crustacean
heads with large head-shields, and with or without
a rudimentary dorsal shield, there can, we think, be
little doubt. We found strong confirmation of the
theory in the form of the closing muscles. It seemed
to us that if our view were correct, the closing muscles
must be modified from those which radiated from the
sternal plate in the transverse plane, and that they
ought, therefore, to show this origin. This surmise
was fully supported by the facts. The sinewy part
of the muscle is found in the centre — the remains
of the sternal plate, from which the muscle fibres
radiate to the outer walls of the shells. If then
it is established that any of the Ostracoda are
descended from Trilobites, we have in this double-
headed closing muscle very clear proof that the
Trilobites possessed the sternal plate which we have
elsewhere assumed for them.

As to the causes of the modification of some of
these primitive Trilobitic or Phyllopodan Crustacea
into Ostracoda, we may perhaps make the following
conjecture, borne out by the rudimentary condition
of the abdomen, and the small number of trunk
limbs. We have only to assume that in some of the
larvae of these primitive Crustaceans with head-shields,
the gradual thickening and stiffening of the chitinous
head-shield did not keep pace with the developing
muscles, whether the powerful mandibular muscles of
an early Apus, or the muscles of the masticatory
and springing first trunk limb of a Trilobitc. This

262 THE APODID^: PART n

uneven development is not much to ask, and if it
occurred as described, it could hardly fail to lead to a
bending of the head-shield along the dorsal middle
line, every time, for instance, a larva sought to put
in practice its inherited tendency of contracting its
muscles for the purpose of rolling up. The failure
to develop a head-shield stiff enough to counteract the
pulls of muscles lying in the transverse plane, may
have thus led to the conversion of the head-shield into
the bivalve shells, which have, in the long run, proved
a better defence than rolling up.

We thus explain the rudimentary state of the
abdomen and trunk. It was only in comparatively
young animals in which but few trunk segments
had been developed, that the bending was likely to
take place, and, when once acquired, it would be
clearly an advantage to keep the abdomen in a larval
stage, in order that it might be quite enclosed within
the halves of the head-shield.

We therefore suggest that the Ostracoda have had
more than one root, and may in fact be derived from
the larvae of any of the primitive Crustacea with
large head-shields, whether Trilobites or Phyllopods.
There seems to be some evidence for both these
origins.

COPEPODA. -

The origin of this very rich group of Crustacea
is very obscure. The general opinion is that they
must be ranked as perhaps the lowest of all the class.
We have now to try to suggest a possible origin for

SECT, xv CLASSIFICATION OF CRUSTACEA 263

the group in the light of what is known as to the
origin of the whole class from a bent Annelid. We
find, then, no group of early Crustacea from which we
can actually deduce them. They are distinctly lower
in the scale of development than any of the early
groups which we have already described, and proved
to be the most primitive. We are thus driven to the
conclusion that they must have originated from some
larval form. There is no difficulty in this sup-
position. Among the enormous number of free-
swimming and independently feeding Nauplii, it
would almost certainly be an advantage to some to
remain but little advanced beyond the Nauplius, the
more pronounced character of the adult bringing them
at once into danger. If we assume that they are
modified larvae of early Apodidae, the conditions, as
far as we know them, would be fairly well satisfied.
The Apodidae were driven from the open sea by some
foe or foes, and would have been exterminated had
they not, in the manner described in the early part of
this book, taken refuge in shallows and lagoons, and
finally in freshwater puddles. We may well suppose,
therefore, that while one division of the Apodidae
thus retreated inland and were able there to develop
into adults, another probably found safety in remain-
ing at the larval stage, their smallness, their trans-
parency, and the rapidity of their motion rendering
them comparatively safe. Whether the organisation
of the Copepoda can be explained on this hypothesis
we are not able to decide. The view that they are
really equivalent to larvae finds some support in the

264 THE APODID^: PART n

fact that they fail, excepting in a few rare cases (e.g.
Argulus), to develop the paired eyes. The unpaired
eye is always present, at least in the free-swimming
forms ; the paired eyes appear as rudiments, only to
disappear again later. The characteristic caudal fork
of the Copepoda might well be a further development
of the fork which appears at the early larval stages of
Apus (see Fig. 41, p. 168). The characteristic ovisacs
may be a modification of the habit of Apus of carry-
ing its eggs about in a brood pouch, necessitated by
the fact that the more larval Copepoda do not develop
enough segments to reach the inherited place of exit
of the genital products, i.e. between the tenth and
eleventh segments.

This theory also is quite in accord with the fact
that so many Copepoda are parasitic. The same
danger which, loosely speaking, drove the adult Apo-
didae into the land, and the larval to remain at the
larval stage, would tend to differentiate the larvae
themselves, as the wind has differentiated the beetles
in the island of Madeira.1 These insects arc cither
strong fliers or else have given up the habit of flying
altogether, the strong winds having swept away all
intermediate grades. The Copepoda are similarly
very markedly divided into two groups, the free and
powerful swimmers, and the parasites who have almost
or entirely given up the habit of free locomotion, except
in the earliest larval stages when seeking new hosts.

The chief difficulty in the way of this derivation of the
Copepoda from an Apus larva is, perhaps, the form of

1 Darwin, Origin of Species, p. 109.

SECT, xv CLASSIFICATION OF CRUSTACEA 265

the limbs. In consequence, however, of our method of
deducing the limbs from the Annelidan parapodia we
do not ourselves experience this difficulty. The typical
Phyllopodan limb is, according to our view, composed
of the dorsal parapodium carrying on the dorsal side
the gill and the sensory cirrus, and on the ventral side
a row of sensory endites, with the remains of the
ventral parapodium as masticatory ridge. The parts
of these limbs which would be useless to the Copepod
would naturally degenerate, i.e. I, the gills, because the
animal breathes through its integument ; 2, the sensory
endites, because the animal would no longer require
to use its limbs in the way Apus uses them to rake
together prey into the ventral middle-line ; and 3, con-
sequently also the masticatory ridge which in Apus
forwards food thus raked together towards the mouth.
On the degeneration of these parts we have left only
the dorsal parapodium with the sensory cirrus, i.e. the
endo- and exopodite of the typical Copepod Ihnb.

Grenacher's account of the unpaired " eye " of
Calanella differs somewhat from that of Apus ;
although there can be little doubt that the two are
homologous. In Calanella only three "retinae" arc
developed, each consisting of comparatively few
retinal cells. There is no trace of crystal cones
or rhabdomeres, and the pigment is in the centre
of the group. The nerves from the retinal cells
come from their inner ends, their sensory ends
pointing outwards. A comparative study of these
unpaired " eyes " has long been a desideratum.

We thus suggest the deduction of the Copepoda

266 THE APODID>£ TART n

from a larval stage of Apus, dating back to the time
when the Apodidae could no longer develop fully in
the open sea, and only those larvae which were acci-
dentally shut off and isolated in lagoons were able to
grow into adult animals.

CIRRIPEDIA.

These animals are now generally supposed to be
related to the Copepoda. What we have said of
the latter applies in great part to them also. We
can deduce them from no original adult Crustacean
form derivable from our bent Annelid. We are there-
fore driven to deduce them, as we have done the
Copepoda, from some larval form. We think it pos-
sible that the Cirripedia may have been one of the
extraordinary lines of development adopted by the
original Copepod, i.e. larval Apus, which sought
safety in a stationary life. As larvae of Apus, it was
always possible for them to develop the shell-fold or
mantle if necessary, the later calcification of which,
perhaps as protection against the browsing Trilobites,
led to the beautiful shell arrangements characteristic
of the group.

We now come to groups the origin of which can be
established with less appeal to the imagination than
was necessary in the former groups. The manner in
which the other Phyllopoda have been derived from
the Apodidae will afford some capable zoologist a
field for research which cannot fail to be rich in

SECT, xv CLASSIFICATION OF CRUSTACEA 267

biological observations of extreme interest1 We limit
our own contribution to the subject to a few points
of some interest and importance.

THE CLADOCERA.

One specimen of Lepidurus glacialis in our collec-
tion was in the act of casting its skin. Shining
through the shell was a white mass, which turned
out to be a group of eggs, thrust in as far as
possible under the neck. It was clear that this
was not accidental ; the eggs were there in order
to develop under the shelter of the cast-off cuticle.
The origin of this arrangement may well have been
accidental. The Apodidae swim on their backs,
so that eggs from the brood pouch might very
easily fall into the large dorsal shell, and this
would be the more likely, the larger the shell in
proportion to the length of the body ; every diving
movement of the animal would tend to lodge the
eggs further up between the shell and the back. The
young hatched out of such eggs may easily be sup-
posed to have derived some advantage from their
position. We have two cases to consider, first, that
in which the eggs hatch out before the cuticle is cast,
and develop under the shell of the parent, and sfecond,
that in which the eggs do not develop before the
shell is cast, the Nauplius swimming about for a time
under cover of the exuvia of the parent.

1 A suggestion as to one of the changes which explain the origin of
Branchipus out of Apus will be found on p. 100.

268 THE APODID^: PART 1 1

In the first case, if this arrangement proved of any
real advantage to the young, it would certainly bring
about such modification in the parent animal as would
lead to the formation of new species, differing from
those which did not so shelter their young. This
may seem a small point around which to mould a
new species, but not if we give to the reproductive
function its true value in the economy of life. Every
other function is in fact subordinate to it, and it is
therefore capable of modifying every part of the body
in order to ensure its own efficiency. Hence, given a
certain number of Apodidae which have inherited a
tendency to drop their eggs under the dorsal shell,
because in this way a greater number are able to
develop and survive in the struggle for existence, these
animals would, in course of time, be modified so as
to perfect this arrangement. The shield would grow
further down at the sides so as to press more closely
against the body, and the hinder part of the body
would come into closer contact with the hinder edge
of the shield, both alterations serving to prevent
the eggs or embryos from slipping out from under
their cover. It is also probable that processes of
the terga might grow up so as to close the
posterior opening (sec Fig. 60).

On the other hand, again, these very alterations,
which make the falling out of the eggs more difficult,
at the same time make the falling in of the eggs
more difficult ; hence the gradual movement of the
genital aperture up the sides under the shell so as to
ensure the egg finding its way into the cavity under

SECT, xv CLASSIFICATION OF CRUSTACEA

269

the shell in which it is to develop. All the altera-
tions which we have here described are exactly what
we find in the related Cladoccra, for instance in the
well-known Daphnia pulex or water flea (see Fig.
60).

- - -- -be

FIG. 60. — Daphnia (after Glaus), showing the brood cavity (be) between the back of
the animal and the dorsal fold ; also the position of the head. projecting freely
from the folded valves of the shell fold. Cf. Figs. 61 and 62.

Again, as to the great difference in size between
the Cladocera and the Apodidae, it is perhaps worth
suggesting (i) that it would originally be only very
young Apodidae, whose shells were specially large in

270 THE APODIM: PARTII

proportion to the length of the body, into whose shells
the eggs would be likely to fall as they swam on their
backs ; the older the Apodidae are, the longer the
body grows in proportion to the shell, and an egg
dropping out of the adult brood pouch would be
hardly likely to lodge under the shell, but would fall
straight to the bottom of the water, — (2) that the
arrangement is not calculated for the development of
many eggs at a time, such as one finds in the brood
pouches and ovaries of adults ; it could only be
advantageously used by the young animals at the
first commencement of their reproductive activity,
when comparatively few eggs issue from the genital
apertures. In this way perhaps we may explain the
small size of the Cladocera, and also the relatively
enormous size of the shield.

The second case in which the skin is shed with the
unhatched eggs in it does not appear to require any
special modification. It may be a custom among
the Apodidae to collect eggs under the loosening
cuticle ; this certainly seems to be the case from
the specimen of L. Spitzbergensis above mentioned.
It did not bear any appearance of being accident.
About six large eggs were packed in so tightly that
they had to be picked out singly with a needle.

It is, however, to be expected that the habit of
hatching vggs under the shield would naturally lead to
some special arrangement for times of ecdysis. Hence
the ephippium of the Cladocera, in which a differen-
tiated part of the cuticle containing two eggs is occa-
sionally cast off as a modified form of ecdysis.

SECT, xv CLASSIFICATION OF CRUSTACEA 271

THE ESTHERID^E.

This is the only other group of the Phyllopoda
about which we have a few words to say. The forma-
tion of the bivalve shell of these animals has already
been noticed. The question is, How can a perfect
bivalve shell, enclosing the whole body, head and all,
be deduced from the folding down of the lateral

FIG. 61. — Limnetis brachyurus, 9 O. F. Miiller (from Bronn's Kiassen und
Ordnungeii), to show the position of the head as transition stage between the
Cladocera and the Estheridae (Fig. 62).

halves of a dorsal shield ? It fortunately happens
that we have a series of forms which make the point
quite clear.

In the Cladocera, we have the shell folded down
against the sides of the animal, leaving the head quite
distinctly marked off (Fig. 60). In Limnetis we find
the lateral folds of the shell extending more ante-
riorly so as partially to enclose the head, the change

272 THE APODID^: PART n

being chiefly due to the bending down of the head in
order to bring it within the shells (Fig. 61).

Limnadia and Estheria show the process com-
pleted, i.e. the head bent down to such an extent as
to be entirely enclosed between the bivalve shells
(Fig. 62). The position of the head in these animals,
bent ventrally downwards, is in striking contrast to
that of the Ostracoda, which is situated far back
in the shell and looks forwards.

We must here leave this interesting subject in the

FIG. 62. — Estheria Donaciiormis Baird 9 > to shovy the completion of the process of
bending the head into the bivalve shell.

hope that some one may be induced to attempt to
build up a natural order of the Phyllopoda, and
endeavour when possible to show how, and under
what biological laws, the different forms have arisen
from Apus.

THE MALACOSTRACA.

We come lastly to the most highly developed
group of the modern Crustacea — the Malacostraca.
We need not say much about these. By deducing

SECT, xv CLASSIFICATION OF CRUSTACEA 273

the Apodidae from a bent Annelid, we have endea-
voured to establish them as the racial form of the
majority of modern Crustacea. We at first thought
Apus might actually be the primitive Crustacean,
but further investigation and comparison with such
forms as the Trilobites have shown us that these
also claim the same origin as Apus from a bent
Annelid. These other groups have for the most part
died out. Apus remains, having been isolated through
many geological periods in freshwater pools. While,
however, Apus itself was not able to hold its own in
the struggle for existence in the open sea, modifica-
tions of Apus succeeded in surviving, and in producing
the rich Crustacean fauna of modern seas. We have
already deduced some of the natural groups from
their Apus ancestors, and we have now the chief
group of all to trace back to Apus.

The Malacostraca have, by general consent, been
traced back to Packard's Phyllocaridae, the only living
representative of which is Nebalia, which, according to
Packard, combines Phyllopodan and Decapodan cha-
racteristics. It has been placed by Claus in a special
order — the Leptostraca — as a transition form between
the Entomostraca and Malacostraca.

Going back to the earlier members of this group,
we find in palaeozoic times the remains of large
Crustacea, which appear to be true Nebalidae. The
most important of these are the two forms Hymeno-
caris and Ceratiocaris (Figs. 63 and 64). At the
first sight of these fossils we are at once reminded
of Apus, and this is exactly what our theory demands.

T

274 THE APODID^: PART II

No one can study the beautiful plates in Jones and
Woodward's monograph of Palaeozoic Phyllopods
without being convinced that the forms represented
were nearly related to the Apodidae. This first
impression is fully borne out when we come to examine
the forms more closely. We find several striking
characteristics of the Apodidae, which convince us that
we really have here to do with animals at least closely
related to and easily derivable from Apus.

FIG. 63.— Hymenocaris vermicauda Salter. Upper Cambrian. To be compared
with Apus (from Zittel).

Hymenocaris has a simple flat shield and a terminal
segment carrying a long caudal plate, and three
visible anal cirri. From the arrangement of these cirri
we may safely conclude that there was a fourth
hidden behind the caudal plate. It will be remem-
bered that we found it necessary to assume that the
original Crustacean-Annelid had four anal cirri, two
of which were preserved in Apus, while the two others
became rudimentary. This assumption certainly re-
ceives some support from the fossil under discussion.

SECT, xv CLASSIFICATION OF CRUSTACEA 275

Anteriorly, we find that the shell has been cut off,
an arrangement which the next form, Ceratiocaris,
fully explains.

Ceratiocaris differs somewhat from Hymenocaris,
but shows even closer resemblance to the Apodidae.
We have the caudal plate and two anal cirri, which
are clearly, as in Apus, the ventral pair. If the fossil
were well enough preserved, we might perhaps find,

FIG. 64. — Ceratiocaris papilio Salter. Upper Silurian. Showing the rostrum, the
first pair of antennae, and the mandibles (from Zittel).

as in Apus, the rudiments of the dorsal pair. Im-
pressions of the mandibles are clearly visible, and
bear the closest possible resemblance to those of
Apus. Traces of branchial limbs have been found
on the abdominal segments of Ceratiocaris Stygia.
Anteriorly, however, we find the same piece of the
shield cut out as in Hymenocaris, with remains of a
rostrum and anterior antennae. The antennae bear a
close resemblance to those of Apus (see Fig. /A, p. 34)

T 2

276 THE APODID^: PART n

although apparently larger in proportion to the size
of the body. The rostrum is, however, clearly a new
structure. How can we explain its origin, at the
anterior edge of an Apus-like head ?

According to our theory the anterior antennae once
pointed backwards, as do those of Apus. In
Ceratiocaris, however, we find them almost at the
anterior end of the head. It is not difficult to show
that this migration would almost necessitate the
formation of a rostrum.

One variation on the primitive Apus type would
certainly be a species using their antennae forwards as
organs of sense. . Just as, in Apus, the eyes travelled
forwards, so, in process of time, the antennse might
tend to move forwards, but, by way of protection for
these, at first, delicate organs, we may suppose them
to have moved forwards in slight grooves on each
side of the median line. As they moved forwards
they may have become more and more developed,
not only as sensory organs, but as appendages, until
they projected freely from the front (as typical
Crustacean antennae). The rostrum is the remains
of the middle wall between the two grooves. It is
clear that such grooves could not exist on the under
surface of the head of an Apus without forming
a primitive rostrum. According to this view, the
rostrum was originally a necessary accompaniment
of the migration of the antennae from the sides of
the labrum to the front of the head. The articulation
of this rostrum was a secondary acquirement not in
itself difficult to imagine.

SECT, xv CLASSIFICATION OF CRUSTACEA 277

This view explains the morphological significance
of the rostrum, as the protective point for the more or
less delicate antennae, arising, not per se, but as the
further development of the tip of the middle piece
between the two depressions along which the antennae
travelled forwards.

From all that remains then of these primitive
Nebalidae we see a sufficient resemblance to the
Apodidae to form a very striking confirmation of
our theory. We see in them true transition forms
between Apus and the higher Crustacea ; the fossils
showing very clearly one of the first steps in this
transformation, and one of the most needful for
success in the struggle for existence, i.e. the gradual
migration of the antennae to a frontal position near
the eyes.

The many points of likeness between Apus and the
Macrura will already have struck every reader of the
first part of this book. The detailed deduction of
Astacus from Apus on the lines here laid down
would be a most interesting and profitable study.

Starting, then, from our theory that Apus, owing
to its likeness to an Annelid, must be one of the racial
forms of the whole group, we have been able, with
varying success, to show that all ancient Crustaceans
are clearly related to Apus, and that all the chief
groups of the modern Crustacea, with the probable
exception of some of the Ostracoda, can be more or
less clearly deduced from Apus. An attempt to derive
the modern forms from the Apodidae in detail would

278 THE APODID^E PART n

be the work of a life-time and would fill many
volumes, but we believe we have established our
theory beyond question, and have shown for the first
time how a natural system of the Crustacea may be
built up by taking Apus as the key to the original
Crustacean form.

SECTION XVI

PERIPATUS AND THE TRACHEATA

BEFORE closing this essay, in which we have
endeavoured to prove that Apus is an almost ideal
transition form between the carnivorous Annelids
and one large division of the Arthropoda, viz. the
Crustacea, it is but fit that we should briefly refer to
Peripatus, which is acknowledged to be a transition
form between the Annelids and the other division of
the Arthropoda, viz. the Tracheata, in which we
include the Myriapoda, Hexapoda, and Arachnida.
It cannot but add to the interest of this book if we
dwell upon this point for a short time.

The accepted fact that both divisions of the Arthro-
poda are derived from Chaetopods, the chief cause of the
transformation being the same in both, viz. : the use of
the parapodia as appendages for mastication and loco-
motion, accounts for the resemblances in the organ-
isations of the Crustacea and Tracheata which have
led to their being placed side by side as Arthropods.
There are, however, striking differences in their

280 THE APODID^E PART n

morphology which stand obstinately in the way of
attempts to establish a close relation between them.

Has not our derivation of Apus and the Crustacea
from a bent Annelid supplied us with the clue as to
the essential morphological difference between the
Crustacea and the Tracheata, leaving out of sight for
the moment the tracheae and the Malpighian tubules
which are confined to the latter ?

The Annelid which gave rise to the Tracheata
started, as did the Crustacean-Annelid, by using its
anterior parapodia as mouth parts, but, unlike the
latter, it did not bend round its anterior segments to
browse in the manner described in the opening sen-
tences of this essay, but remained straight. The fusion
of segments to form the head was, in the Tracheatan-
Annelid, axial, the mouth remaining at the anterior
end of the body.

In such an axial fusing there is nothing to fix the
number of segments to form a head common to all
the Tracheata, whereas in the Crustacea the bending
round of the five segments marked off this region of
the body as the head for all time.

The difference between the number of the cephalic
appendages of the Crustacea and the Tracheata is to
be referred to the fact that with the mouth at the
anterior end of the Annelidan body it was impossible
to bring so many pairs of parapodia into the region
of the mouth to function as mouth parts as in the
Crustacea, where its ventral position allows of the
arranging on each side of a large number of para-
podia as jaws.

SECT, xvi PERIPATUS AND THE TRACHEATA 281

The common derivation of the two divisions of the
Arthropoda from Annelids modified to use the para-
podia as jaws, &c., in the one case round a mouth
at the anterior end of the body, and in the other
round a mouth bent under so as to face posteriorly,
makes it possible, we think, for the first time
clearly to homologise the head regions of the two
divisions.

The Annelidan prostomium became in both cases
the labrum. In both groups the Annelidan antennae
were retained as sensory organs, having disappeared
only in the Arachnida. The first pair of parapodia,
the antennal parapodia of the Annelids, became
differently modified on account of the different posi-
tion of the mouth. In the Crustacea the mouth was
carried round ventrally to between the parapodia of
the third and fourth segments, which thus, in the
typical Crustacean head, became the chief jaws, leav-
ing the antennal parapodia as a rule free to continue
to function as sensory organs. In the Tracheata, on
the other hand, the anterior position of the mouth
almost necessitated the formation of the chief jaws
out of the first pair of parapodia. In Peripatus these
alone function as jaws. In the Myriapoda and
Hexapoda they are the chief jaws, but are assisted by
the two following pairs as first and second maxillae.
In the Arachnida they form the powerful and vari-
ously modified chelicerae which develop so largely as
to displace and lead to the degeneration of the pro-
stomium and antennae. These formidable jaws are
assisted by the second pair of parapodia as accessory

282 THE APODID^: PART n

jaws, supplied with long feelers, or as powerful chelate
limbs.

The second pair of parapodia, which in the typical
Crustacean head become the chief mandibles, form, in
Peripatus, the oral papillae ; the slime glands opening
at their tips being perhaps homologous with the
acicular glands of the Annelidan parapodia. In the
Myriapoda and Hexapoda they become the anterior
maxillae ; and in the Arachnida they form the pedi-
palps or their homologues.

The third pair of parapodia, which in the Crustacea
form typically the first pair of maxillae, in Peripatus
and the Arachnida function as the first pair of feet.
In the Myriapoda and Hexapoda they form the
posterior maxillae.

Just as we saw that all the Crustacean groups, how
ever aberrant, must have been derived from the same
bent Annelid, so we would deduce all the groups of
the Tracheata from the same Tracheatan-Annelid.
We find the same variety in the arrangement and
form of jaws, limbs, &c., and the same variety in the
number of segments. In both cases some of the
groups can be shown to have been differentiated direct
from the original Annelid, while others are only later
modifications of such groups. In the Crustacea we
think the Apodidae, and the Trilobites, were original
differentiations ; in the Tracheata, the Arachnida, the
Protracheata, and the Myriapoda.

Turning now to the important morphological
characteristics common to all the Tracheata, viz.
the tracheae and the Malpighian tubules, we shall

SECT, xvi PERIPATUS AND THE TRACHEATA 283

not, we think, be far wrong in assuming that these
were developed as adaptations to a life on land, and
appeared in the original Tracheatan-Annelid, in its
gradual passage from a purely aquatic to a terrestrial
life. It seems to be a strict biological law that, when
aquatic animals migrate to the land, external respira-
tory surfaces such as gills, which are morphologically
folds of the skin, give place to internal respiratory
surfaces. This requires no special comment. It is
probably, however, an equally strict biological law
that free movement on land necessitates such a place
of exit for the waste products as will not interfere with
such movement. Insects clean themselves from no
love of cleanliness. The disadvantages of discharging
the waste products in the cephalic or thoracic region,
as in the Crustacea, are avoided by means of the
Malpighian tubules which open into the hind-gut.
This is not the only advantage. Small land animals
have often to exercise the most rigid economy in their
supply of fluid. The discharge of the waste products
into the hind-gut permits the reabsorption of their
purely fluid constituents, which would thus be re-
tained within the body. These two advantages are of"
such importance that the gradual concentration of
excretion to the walls of the hind-gut (which we saw
in Apus to be highly glandular) until special excretory
caeca, the Malpighian tubules, were developed, pre-
sents no difficulty.

\Ye have already referred to the able attempt of
several distinguished zoologists, Kingsley in America,
and Ray Lankester in England, to connect the Arach-

284 THE APODID^ PART ii

nida with the Xiphosuridae and Eurypteridae — taking
the two latter out of the division of the Crustacea.

If there is any truth in our general argument as to
the derivation of the primitive Crustacea from a bent
Annelid, and of the Tracheata from an Annelid not
so bent, there is no need for any such alteration
in the formerly accepted classification. The resem-
blances in inner and outer organisation between
the Xiphosuridse and the Scorpionidae, striking as
they undoubtedly are, we believe to be simply
due to the fact that they are both descended from
Annelids. The agreement in the number of segments
and cephalothoracic limbs is by far the most important
argument in favour of the new classification.

But now it seems to us that it is by no means im-
probable that two groups of animals descended from
many-segmented Annelids should possess the same
number of segments, especially when we find that
somewhere about the same number of segments seems
to have best suited many other groups belonging to
both divisions. The Malacostraca have twenty, the
free-swimming Copepoda about fifteen, the Hexapoda
sixteen, and many genera of the Myriapoda from
fifteen to thirty.

The resemblance between the limbs of Limulus and
Scorpio does not seem to us so great as it is often
assumed to be. The five pairs of jaws ranged round the
ventral mouth of Limulus, whether our theory of their
origin from Annelidan parapodia is true or not, form
a feature which has no counterpart in the limbs of
Scorpio. This is, to our mind, a most important

SECT, xvi PERIPATUS AND THE TRACHEATA 285

point, for in most other respects all Arthropodan legs
strongly resemble one another, and the presence of
chelae on a certain number of anterior limbs is a com-
mon occurrence. Again, is there anything in Limulus,
or in any Crustacean, which resembles the two chiti-
nous hooked-claws at the ends of the legs of Scorpio,
which the latter possess in common with all other
Tracheata ? Nor do we find in the Scorpionidae any
special development of the sixth pair of limbs such as
we have shown to be characteristic not only of the
Apodidae but of the Trilobitae, the Xiphosuridae, and
the Eurypteridae, and which is especially marked
in the last, although this is claimed as a transition
form between the Xiphosuridae and the Arachnida.

We do not, then, admit that very much weight can
be laid upon this agreement in number of segments
and in number and form of limbs. It certainly cannot
outweigh, for purposes of classification, the tracheae
and the Malpighian vessels, the presence of which in
the Scorpionidae and other Arachnida classes them
unmistakably with the Tracheata.

Even if we admit the possibility of the concurrent
development of tracheae and Malpighian tubules for a
second time, the improbability of such an occurrence
is so great that we should require much stronger
evidence than any which has been adduced before we
could accept it. It is, further, very improbable that such
a highly specialised animal as a species of Eurypterus
should develop exactly the same respiratory and excre-
tory adaptations to a land life as the more generalised
Annelidan ancestor of the other Tracheata.

286 THE APODID^E PART n

The origin of the book-leaf tracheae from the gills
of the Xiphosuridae, fascinating as it is, breaks down
when carried into detail. It is easier to believe that
the lung-books are only a specially concentrated
arrangement of the tracheal tubes, no more extra-
ordinary than the other extreme, viz. the diffuse
arrangement found in the Hexapoda. We find
almost every form of tracheal arrangement between
these two extremes within the division of the
Tracheata, and further both tubular and book-leaf
tracheae within the Arachnida. We think that the
evidence in favour of the new classification, to be
drawn from the form of the tracheae, is not con-
vincing.

The most probable origin of the tracheae appears
to us to be that which refers them back to dermal
glands. The original Tracheatan-Annelid on first
migrating on to the land probably respired through
the whole skin. The increase of surface afforded by
the ducts of the dermal1 glands would very naturally
be taken advantage of. The walls of these ducts being
internal, their surfaces would be selected and special-
ised until they undertook the whole respiration. That
this was the origin of the tracheae is rendered very pro-
bable by the fact that the openings of the tracheal tubes
in Peripatus are, in some species at least, scattered
irregularly over the whole body. This derivation of the
tracheae from dermal glands receives some support also

1 If these include the coxal glands, it may throw light upon the
developmental relations between the book-leaf tracheae of the Arachnida
and their rudimentary abdominal limbs.

SECT, xvi PERIPATUS AND THE TRACHEATA 287

from the fact that it is accompanied by the development
of the Malpighian tubules, except in Peripatus, where
the nephridia are retained. The loss of the dermal
excretion necessitates the further development of
other excretory surfaces. The advantages of the
Malpighian tubules, or glandular caeca of the hind-gut,
over excretory organs in any other part of the body
have been already dwelt upon. This physiological
connection between tracheae and Malpighian vesicles
which lessens the improbability of their concurrent
development twice, cannot however be taken advantage
of in the special case under discussion, because the
tracheae are not supposed to have been dermal glands
but imbedded external gills.

The early differentiation of the Arachnida from the
original Tracheatan-Annelids accounts for the high
specialisation of their tracheal gills. The same may
be said of the Myriapoda, while Peripatus has remained
in this respect, as in so many others, almost entirely
undifferentiated.

In addition to these arguments we have to refer on
the one hand to those brought forward in this essay to
show that Limulus is a Crustacean, and on the other
hand to the discovery of rudimentary antennae in the
embryo of a spider,1 which removes the only difficulty
in the way of homologising the limbs of the Arach-
nida with those of the Antennata.

It seems to us that we find evidence of the early
specialisation of the Arachnida, not only in the loss
of the antennae, in the form of the limbs and trachese,

1 Trochosa Singoriensis Laxm. See Zool. Anzeiger, May u, 1891.

288 THE APODID^: PART 11

and in their general organisation, but in the posses-
sion of a sternal plate or entosternite. The same
explanation given in this essay of this sternal plate
in Apus and Limulus must be applied here. It is due
to a massing together of the ventral longitudinal
muscle bands of a certain number of anterior seg-
ments, so that their muscular elements disappear,
leaving the sinewy elements for the attachment of
transverse muscles. In the primitive Crustacea,
the longitudinal muscles of these segments were
rendered useless by the bending of the body. In the
Arachnida, however, they were rendered useless by
the axial fusing of the segments ; while the muscular
elements degenerated, the sinewy elements were
retained to form the entosternite. This seems to
show that the Arachnida were differentiated from
the Tracheatan-Annelid at a stage when the
Annelidan segments were still of the typical form,
i.e. before the ventral longitudinal muscle bands had
become specialised in adaptation to new modes of
life.

In conclusion, it may be interesting to see how
Peripatus compares with Apus as a transition form.
The Annelidan characteristics of Peripatus are
certainly more visible than are those of Apus, where
they are all more or less disguised or transformed.
On the other hand, Peripatus is a development by
itself, and can hardly be shown to have given rise to
any group of the Tracheata. It is indirectly a most
remarkable transition form, having preserved so many
characteristics of the common racial Tracheatan-

SECT, xvi PERIPATUS AND THE TRACHEATA 289

Annelid. We think that a little more may be
claimed for Apus, and that in its organisation it takes
a distinct place in the direct line of descent of many
of the modern Crustacea from the original Crustacean-
Annelid.

U

APPENDIX I

ON comparing the East Spitzbergen species found by
Professor Kiikenthal with the West Spitzbergen species
found by Professor Nathorst, we concluded that they are
identical, but that L. Spitzbergensis differs considerably
from L. glacialis in size and in the shape of the caudal
plate. We were at first disposed to consider it a new
species, especially on account of its possessing second
antennae which were said to be wanting in Lepidurus
glacialis. .Closer examination, however, showed it to be a
small variety of L. glacialis, most probably derived from
the latter by being obliged to ripen at an earlier stage of
development, in adaptation to the shortness of the more
northerly summer.

That this view is correct seems probable from the fol-
lowing considerations :

(1) The possession of second antennae does not dis-
tinguish it from L. glacialis, for we have succeeded in
finding these appendages on the latter.

(2) The position of the sperm-forming centre (see § on
reproduction) is identical in the two.

(3) The genital tube is very much simpler, the diverticula
showing hardly any traces of branching, therein exhibiting
a more larval condition.

(4) The same may be said of the smaller size of the
caudal plate, which develops gradually, as Brauer has shown
in his paper on the development of L. productus.

U 2

292 APPENDIX I

(5) The small size of the whole animal also agrees with
the supposition.

It is interesting to find that Packard's measurements for
L. glacialis (from Cape Krustenstern ?) make it even smaller
than the Spitzbergen variety. From this, however, it is
difficult to draw any certain conclusions, as his drawings
give a fully-developed tail-plate (see Monograph of the
North American Phyllopoda). It thus appears that L.
glacialis may be much stunted by unfavourable surroundings.

That the specimens from Spitzbergen were not young
specimens follows from the facts that they (several hundred)
were nearly all the same size, and that they were caught in
the end of August, a week or so before the close of the short
summer, while the freshwater pools were still unfrozen. Pro-
fessor Kiikenthal informs me that this season in the latitude
in which they were found lasts about ten weeks.

Packard's measurements for a fully developed L. glacialis
make it doubtful whether we are to look upon this variety
as permanent. It is possible that in favourable summers
they may further develop (without any great increase of
size) into stunted L. glacialis. This question, however, can
only be certainly answered by cultivating specimens further
south, in an aquarium, to see whether they develop into
L. glacialis. In the meantime it will be useful to call the
animal L. glacialis var. Spitzbergensis, or, for shortness,
L. Spitzbergensis.

APPENDIX II
THE EYE-PIGMENT OF APUS

IT was very difficult to decide whether the cells marked
/ in the diagram (Fig. 43) of the eye of Apus were really
cells, as there drawn, or only collections of very minute
pigment cells. [Grenacher, in his drawings of the single
eyes of Apus, leaves the matter rather indefinite. He
indicates rather than draws the pigment cells with nuclei.
His drawing leaves the impression that he took it for
granted that they were large pigment cells, without actually
ascertaining the facts.] We were at first inclined to take
the latter view, having found that under a very high power, l
the granules themselves were not easy to distinguish from
cells. Each one consists of a stainable nucleus surrounded
by a pigment crust, the whole being enclosed in a layer
of some hyaline substance. These " cells J> were of all
sizes (from 1-2 //,), and were found in all stages of fission
(see Fig. 65). There are thus two ways of regarding
these pigment masses in the eye of Apus. Either the
whole is a kind of loose syncytium of minute pigment
cells, as we at first thought, or these pigment gran-
ules are formed inside a large cell around stainable
protoplasmic granules, as starch is formed round the
leucoplasts. This we now think to be the case.

1 Zeiss apochromatic 2 mm. homogeneous immersion, 1.40 n.a.,
eye-piece No. 12, giving 1500 diam.

294 APPENDIX II

Although we cannot be certain that we have seen the
nuclei of the large pigment cells as shown in the dia-
gram (Fig. 23, p. 139), we concluded that there must be
such nuclei, and that the pigment masses were real cells
and not syncytia. We were chiefly led to this conclusion
by noticing the long regular lines of granules running down
the nerves towards the optic ganglion, as shown in the
diagram. It seemed to us that these rows of single
granules would not be so straight and even, unless enclosed
within a long pseudopodium-like process of the pigment
cells. Were the granules semi-independent cells, their
arrangement could hardly be so straight and regular. We

FIG. 65. — Pigment granules (? cells) from the eye of Apus, X ca. 3000, showing a
stainable nucleus, surrounded by a thin crust of brown pigment, the whole
enclosed within a hyaline substance.

were further induced to take this view from finding that, in
some specimens, the pigment in the unpaired " eye " was
composed of similar eye-pigment granules, also arranged in
long pseudopodium-like strands. In most of the specimens
examined, the pigment in the unpaired "eye"' was similar
to that in the pigment cells of the rest of the body, i.e. it
was in the form of very minute olive green granules. The
occasional finding of eye-pigment in the unpaired " eye "
was especially interesting in reference to the origin we
attributed to that organ out of an anterior pair of
Annelidan eyes.

Around the paired eyes, the green pigment reaches up to
their very rim, and indeed stretches over the outer edges of

APPENDIX II 295

the eye itself, but there it changes into the black brown
granules above described.

These " eye-pigment " granules certainly appear to be
very primitive formations. The utilisation of excretory
matter as pigment is at once suggested, the incrustation of
brown stuff round the nucleus reminding one forcibly of
the incrustation of excretory matter round the blood cor-
puscles under the dorsal organ (see Appendix IV.). In the
pigment granules, however, . it was quite regular, whereas
it was irregularly massed around the blood • corpuscles.
These corpuscles, again, are very much larger than the
pigment-forming granules, and moreover fairly uniform in
size, whereas the latter are of all sizes.

APPENDIX III

CIRCULATION

As far as we know, since Zaddach's time no detailed
account of the circulation of Apus has been given. Ger-
staecker adopts and incorporates Zaddach's description in
Bronn's Klassen und Ordnungen, vol. v. Zaddach's ob-
servations seem to have been made on living transparent
animals. All who have tried this method know how diffi-
cult it is to make out the details, however visible some of
the main streams may be. Thus Zaddach's plan of the
circulation requires considerable amendment.

As already pointed out in the text, the system is a
lacunar system through which the blood is propelled by a
contractile dorsal vessel or heart. On the expansion of the
heart the blood is drawn out of the cardial sinus to be
propelled forwards through (i) the anterior aorta to supply
the head and liver, and (2) the two lateral vessels which
dip down under the shell gland to convey blood into the
shield.

The heart is composed of striated circular muscle fibres
crossing each other diagonally — the muscle-cells being
turned inwards, and forming a kind of syncitial lining
to the tube. The heart is suspended by an exquisite
arrangement of connective-tissue fibres, which, seen together
under a low power, take the form of triangular wings.
These connective tissue alae are not flat and membranous,

APPENDIX III

297

but composed of a number of fibres attached to the walls
of the heart over a considerable area around the ostia.
They may either be contractile and serve to expand the
heart, or, more probably, elastic and restore the heart to its
expanded condition after each contraction.

The blood, after circulating through the head, runs
ventrally backwards through the intestinal sinus, towards

FIG. 66.— Diagram to illustrate the plan of the circulation in the anterior part of the
body. The blood propelled by the heart (h) through the head, is returned
through the intestinal sinus (is) from which in each segment it escapes ventrally
into the limbs. Its course is indicated diagrammatically by the arrows (/). From
the limbs it returns through the dermo-muscular sinus in each segment into the
cardial sinus (cs). z", intestine; ac, aorta cephalica; p, points of attachment of
the dorso-ventral muscle bands.

the posterior end of the body. Near each pair of limbs,
the membrane forming this sinus appears to be fenestrated,
the openings being regulated by special muscles (?). Through
these windows the blood streams down over the ventral
cord and into the limbs on each side ; it runs along the
ventral face of the limbs, returning along the dorsal. On
its way back it is guided into the gills, and thence back

298

APPENDIX III

into the body, where it flows up between the membrane of
the intestinal sinus and the body wall, bathing the muscula-
ture in its course. It is interesting to note that a special
separate stream flows from each limb into the cardial sinus,
there being membranous dissepiments, corresponding with

ao

FIG. 67. — Diagram of the circulation from above. '7i, heart, expanded by the fine
connective tissue ala; («) ; s, segmental septa forming the transverse walls of
the dermo-muscular sinuses (Is) ; ac, cephalic aorta ; g, lateral artery to shell
and shell gland ; /, points of attachment of dorso-ventral muscle bands to dorsal
wall.

the segmental constrictions, stretched between the body wall
and the membrane of the intestinal sinus. The dissepi-
ments, which have already been described in the text, are
only formed in the first ten to eleven segments, i.e. as
far as the heart extends. Thus while part of the blood,
flowing through the intestinal sinus, passes down through

APPENDIX III 299

the openings above the ventral cord into the limbs, to
find its way back to the heart after passing through the
gills, the rest continues posteriorly, bathing the intestinal
canal (the hind-gut) to find its way, by some means which
is not yet clear, into the dermal sinus which, in the
posterior part of the body, is continued right round the
body, there being no dissepiments and no separate cardial
sinus. It then flows forwards till the first lateral dissepi-
ment confines it to the dorsal channel which contains the
heart and forms the cardial sinus. How the blood finds
its way from the intestinal to the dermal sinus in this
posterior part of the body we have not been able to ascer-
tain. A longitudinal dissepiment runs right along each
cercopod or anal cirrus, which shows that the blood flows
along one side and back by the other. We may also per-
haps assume that under the two rudimentary cirri openings
occur corresponding with the communication which once
existed at the tips of the cirri which they represent. These
are however probably not the only openings between the
two sinuses. We have not been able to make out the
relation of the circulation to the rudimentary limbs ; sec-
tions of the rudimentary gills seem to show that they are
functional as such.

We have seen that special muscles probably regulate the
flow of the blood out of the intestinal sinus into the neural
sinus (if it can be so called), on its way to the limbs. There
can be no doubt that the dorso-ventral muscle-bands play a
part in propelling the blood through the sinus. The intes-
tinal musculature, except in the hind-gut, is too weakly
developed to assist much. But it is easy to see how, in
such a tubular sinus, the movements of the intestinal canal
running along its centre could materially help the circula-
tion of the fluid between them.

We had no means of following the course of the blood

300 APPENDIX III

from the dorsal shield after bathing the shell glands. As
far, however, as anatomical researches enable us to judge,
we cannot confirm Zaddach's statement that it returns to
the heart through special venae branchiales. We think that
when this point comes to be further investigated it will
probably be found that the blood, which enters the shield
laterally, circulates round through it, and returns to re-enter
the body anteriorly and dorsally ; not, however, to enter the
cardial sinus, but rather to descend on each side of the
cephalic aorta, bathing the mandibular muscles on its
way to join the main stream from the head into the
intestinal sinus.

APPENDIX IV

EXCRETION

The Shell Gland. — The shell gland falls easily into
three typical sections (Fig. 30, p. 125); the terminal
saccule, the urinary canal, and the urinary duct with
the bladder. The terminal saccule is branched and
irregular, and lies in the blood stream between the
central coils of the urinary canal. As far as we could
judge from our material it seemed to be lined by large
flat granulated cells, resting upon a fine basal membrane.
It is difficult to say if the large vacuoles to be seen in many
of them are natural.

The urinary canal shows the structure depicted in Fig. 68,
which is characteristic of the antennal gland of the other
Crustacea. Grobben described the striped appearance as
being due to protoplasmic strands arranged vertically on the
basal membrane owing to the active streaming from without
into the lumen of the tube. Tangential sections of the mem-
brane, however, show it to be an independent spongy struc-
ture, like hardened foam (see Fig. 68, to the right), forming
a strong but very porous support to the large flat epithelial
cells. Grobben figures the nuclei as imbedded in this
striped membrane. In Apus, however, they lie with their
surrounding protoplasm on the membrane, here and there
heaped up, or even artificially torn away, in both cases
leaving the membrane intact, which would hardly have

302 APPENDIX IV

been the case if the striation were really due to strands of
protoplasm free in the cells. There is no trace in Apus of
a chitinous cuticle lining the canal.

The nuclei of the epithelium are very large and oval, the
longest diameter being 40-45^ (in a specimen of A. cancri-
formis), i.e. slightly larger than the nuclei of the nutritive
cells of the eggs, a sign of their great physiological activity
in the economy of the animal.

The urinary canal shows a slight widening as it bends

n

FIG. 68. — Part of a section of the urinary canal (shell gland of Apus). bin, basal
membrane on which rests the harder supporting framework, seen in tangential
section (at is) to have a spongy structure;^, inner layer of protoplasm; n,
nuclei with numerous clear round nucleoli.

down towards the ventral side. We at first thought that
this might be the bladder, but there is no change in the
character of the epithelium. At the base of the second
maxilla, this wider portion leads through a very narrow
chitinous canal into the true bladder, which is a chitin-lined
sac in the shaft of the limb. The chitinous lining of the
bladder, which distinguishes it from the urinary canal, makes
it a suitable reservoir for excretory fluids.

The opening of the duct at the tip of the second maxilla
is shown in Zaddach's drawing as a point which he, however,

APPENDIX IV

303

did not understand. Glaus recognised it as an opening of
the shell gland. This, as already shown, we have been
able to confirm by following the gland through series of
sections.

p

FIG. 69.— Sketch of the region about the eyes of an adult specimen of L. productus,
showing the relative position and sizes of the dorsal organ, the paired eyes and
the pore (/>) leading into the water-sacs. At the anterior end of the dorsal organ
is seen the fine pore mentioned in the text.

The Dorsal or Neck-Gland. — One more gland in Apus,
the neck- or dorsal gland, remains to be mentioned.
It shows as an oval spot behind the eyes, and is

304 APPENDIX IV

visible in all the Apodidse. We originally thought that
this spot was the remains of a frontal cirrus which
travelled back with the eyes, but which, being a hindrance
to swimming and burrowing, had become quite rudimentary.
Gerstaecker suggests the homology of it with the frontal
cirrus (Stirnzapfen) of the Ostracoda. From lack of any
detailed study of its finer structure it has, in fact, been
very generally claimed as a sensory organ of some kind.
A close microscopic study of it, however, shows very clearly
that it is an excretory organ.

Fig. 70 shows the surface view, and Fig. 71 shows it in
longitudinal section —both are taken from adult specimens
of L. productus. Fig. 38, p. 160, shows it in the Nauplius of
Apus cancriformis. From this last figure we may perhaps get
a hint as to its real origin and significance, viz. that it was
the larval excretory organ.

First, however, as to its structure. A longitudinal section
shows us a number of fine connective-tissue strands stretched
between the thin cuticle of the organ and the connective
tissue belonging to the longitudinal muscle-bands, which
bend round over the mid-gut to be attached close to the
prostomium. These fibres lie in the full blood stream issu-
ing from the aorta cephalica, and form a net-work to arrest
the blood corpuscles.1 This net-work does not, however,
stretch right across, but, as the animal always swims on its
back, it forms, as in the drawing, a ground-net to catch those
particles which sink, and roll along the bottom. Hence,
while the ordinary blood corpuscles shoot through the open
part as indicated by the arrow, those laden with excretory
matter are caught in the net spread across their path as they
roll heavily along. The whole action is purely mechanical ;

1 This interesting use of the connective-tissue fibres is well illustrated
in many parts of Apus, particularly around the large reserve or fat cells.
It has already been noticed by Grobben in another connection.

APPENDIX IV

305

the corpuscles laden with waste stuff are too large to pass
the meshes ; they seem to stick on to the connective tissue
fibres and gradually find their way down to the hypodermis,
where they either break up or else, after discharging their
burdens, return into the blood stream. We are inclined to
think the latter to be the correct account, for the connective-

FIG. 70. — Diagram of a section of the neck gland or dorsal organ of Apus, drawn
upside down (i.e. as the animal swims) so as the better to illustrate the catching
of the laden blood corpuscles in the connective tissue net. m, epithelium of
the mid-gut ; c, connective tissue belonging to the longitudinal musculature ; en
blood stream from the aorta cephalica ; b, blood corpuscles laden with excretory
matter ; at, cuticle (the small arrow indicates the position of the pore shown in
the last figure). The glandular epithelium is shaded dark.

tissue strands were covered with excretory matter, and the
corpuscles near the hypodermis were not nearly so heavily
laden as those newly arrived. The hypodermis cells them,
selves were very different from those of the ordinary cuticle :
they were much larger, with larger nuclei, each nucleus
containing two or three nucleoli, whereas the ordinary hypo-
dermis cells have but one nucleolus. They also stain badly-

X

306 APPENDIX IV

having a muddy look, doubtless due to the excretory matter
absorbed. How this matter is discharged we cannot see.
The cuticle is extremely thin and perhaps allows of passage
through it. If so, what is the use of the single fine pore at
one end, which by itself could apparently only relieve one or
two of the hypodermis cells ? We have thus not been able
to ascertain the mechanism of discharge, but that the whole
organ is essentially glandular, no one who has studied it can
doubt.

We can say nothing certain as to the origin of this organ.
From its relatively enormous size in the Nauplius,1 it
is clearly the principal larval excretory organ, and under-
takes the discharge of waste products before the shell gland
appears. It may perhaps be a sort of island of Annelidan
hypodermal glandular cells left by the developing exo-
skeleton, taking the place of the head nephridia of the
Annelidan larva. That these latter should not be developed,
owing to the bending double of the front segments of the
Crustacean-Annelid, was to be expected. We may perhaps
therefore find in this neck organ a group of dermal glandular
cells, derived from the Annelidan dermal glands, and
serving for excretion until the typical Crustacean glands
are developed. Its singular position in the larva may
perhaps be considered as protective, since an excretory
gland might well serve as a protective organ on the exposed
dorso-frontal surface.

A comparative study of this organ, which also plays an
important part as an excretory organ in most Crustacean
embryos or larvae, is much to be desired. According to
Bullar, in some Isopodan embryos it forms as an invagina-

1 In the Nauplius, figured p. 160, it measures about 0.25 mm., whereas
that of the adult L. productus (Fig. 69) measures only 0.5 mm.
Brauer, curiously enough, shows no traces of it in the Nauplius of L.
productus (Fig. 35).

APPENDIX IV 307

tion of the ectoderm. As a starting point for such a com-
parative study we should like here to emphasise the fact
that, if our theory is correct, the primitive structure of the
organ is most probably to be found in the larva of Apus,
and that its form in the other Crustacea must have been
derived from that. We do not therefore see any reason
to modify our suggestion as to its origin because of the
fact that in the higher Crustacea it first appears as a more
complicated, and even sometimes as a paired, organ.

It is this organ which, in the Daphnidae, functions as a
sucker for fixing the little animals to stationary objects. The
glandular nature of the organ might easily be supposed to
assist this action by supplying a slimy secretion.

X 2

APPENDIX V
REPRODUCTION

A. HERMAPHRODITISM OF THE APODID^E. l

(From Naitire, vol. xliii. p. 343.)

THE reproduction of Apus cancriformis has been a much
discussed subject. Although the animal has been well
known since the middle of last century, it was not till 1833
that a male was reported to have been found, and not till
1856 that the occasional presence of males in small numbers
was certainly established by Kozubowski. On the other
hand, the fact that several generations of "females" could
be produced without the presence of a male, was established
as long ago as 1755 by Schaeffer, who concluded that the
animals were hermaphrodite. Since that time authors have
been divided in opinion between hermaphroditism and
parthenogenesis (not to mention v. Siebold's theory of
Thelytoky) ; the latter view has lately prevailed.2

1 The letter here reprinted was written before the author had
recognised the Annelidan character of Apus which led to the writing
of this book ; hence its point of view is not altogether the same as that
of the foregoing pages.

2 For the history of this subject see Bronn's Klassen und Ordnungen
des Thierreichs, vol. v. On p. 810 the following words occur : —
" Untersuchungen iiber die Gattungen Apus und Daphnia, welche
offenbar in dem bis zu voller Evidenz gefuhrten Nachweis der partheno-
genetischen Fortpflanzung beider gipfeln." See also Lang's Lchrbnch
der Vergleichendc Anatomic, p. 393.

APPENDIX V 309

The animals, however, prove after all to be hermaphro-
dites. Since the last careful study of Apus cancriformis, as a
whole, by Zaddach in 1841 (the works of Ray Lankester
and others deal only with special points), new methods of
research have been introduced into our laboratories which
reveal details not easily discoverable by the older methods.
Zaddach's figures of the ovaries and testes of Apus are thus
naturally somewhat deficient — as deficient, indeed, as the
best work we can do to-day will, we hope, be found to be
fifty years hence.

*•***.****

In my preliminary notice (Jenaische Zeitschrift fur
Naturwissenschaft) Band xxv., N.F. xviii.) announcing
the hermaphroditism of L. Spitzbergensis, knowing how
much the reproduction of the Apodidae had been dis-
cussed, I ventured to assert that in all probability the
other species of the genus would also prove on closer
examination to be hermaphrodite. As above stated, I
found the sperm-forming centres in L. glacialis in identically
the same position as in the Spitzbergen variety. By
the kindness of Professor Mobius, the Director of the new
Berlin Museum, and of the Rev. Canon Norman, I have
also been able to examine Apus cancriformis and Lepidurus
productus. In both these the sperm-forming centres
were found scattered here and there among the rich
branchings of the segmental diverticula of the genital tube.
They occur either at the tips of such branches, where the
eggs ordinarily develop, or as slight lateral bulgings of the
same. In all cases the spermatogenesis is the same, the
epithelium breaking up into sperm-cells ; these escape into
the lumen of the tube, and are found in considerable
numbers near the genital aperture, where the epithelial
lining of the tube is hardly demonstrable, the walls of the
tube consisting of a fibrous membrane, in the folds of which

3io APPENDIX V

the sperm-cells lurk. The eggs are then fertilised as they
stretch this membrane in passing out into the egg pouch.
The whole richly-branched reproductive organ, with the eggs
developing at the tips of the branches, and with here and
there a testis, strongly reminds one of a monoecious plant,
self-contained, and able to dispense with pollen from
without.

I reserve the drawings and the more detailed description
of the reproductive organs of the different species for a short
comparative study of the Apodidae which I hope soon to
have ready for the press.1 By way of caution, however, I
should here add that small yellowish sacs filled with minute
cells occur here and there among the developing eggs.
These must not be mistaken for the testes. They are the
loci of discharged eggs, and the minute cells are the epithe-
lium cells dislodged by the shrinking of the membrane of
the genital tube, which is stretched some loo-fold by the
ripening eggs.

The origin of this secondary hermaphroditism is not far
to seek ; it is clearly a protection against isolation, as in the
case of the Cirripedia and certain parasitic Isopoda. The
manner of life of all these animals is such that they are
always in danger of being cut off from their kind ; they would
thus die out unless able to reproduce either parthenogeneti-
cally or by means of self-fertilisation.

Some species of Cirripedia, as is well known, have dwarf
males, the last remains of the original separation of the
sexes. As already mentioned, small males of Apus cancri-
formis are sometimes found. Twelve finds of A. cancriformis
and L. productus recorded by Gerstaecker, give 4,458
"females" (i.e. hermaphrodites) to 378 males; while sixteen

1 As stated in the Preface, this intended work gave way before the
more ambitious task of trying to prove Apus to be but a modified
carnivorous Annelid.

APPENDIX V 311

finds, numbering 10,000 individuals, did not contain a single
male. I have found no record of a male L. glacialis, and
none of the twenty odd specimens of the Spitzbergen
variety I have as yet examined have been males. It is
probable that throughout the whole genus self-fertilisation
is taking the place of cross-fertilisation, but that some
species have gone further than others in dispensing with
males. Two species, for instance, L. couesii, Packard, and
L. macrurus, Lilljeborg, are reported to have more males
than " females " (?), but the finds in these cases seem hardly
large enough to allow us to judge ; it may have been purely
accidental that more males than " females " were caught.

The males of the Apodidae, with the doubtful exception
of L. productus, seem to be smaller than the hermaphrodites,
otherwise there is no very pronounced sexual dimorphism,
as there is among the Cirripedia. We are perhaps justified
in concluding from this that the hermaphroditism of the
Cirripedia is of much older date than that of the Apodidae.
No comparison is here, however, possible, since the two
have nothing further in common beyond the fact that they
are both hermaphrodite, and that this hermaphroditism is in
both cases an adaptation against extermination through too
wide dispersion of the individuals.

B. ON THE FORMATION OF THE EGGS.

The regular formation of the eggs out of four cells, of
which three are nutritive and one the definitive egg-cell,
gives opportunity for many interesting observations. The
general method of growth is shown in Fig. 33, p. 144,
where we see the egg in different stages.

The originally round group of cells as a rule soon
becomes oval, in consequence of the more active growth

312 APPENDIX V

of the three nutritive cells. Traces of this activity can be
seen in the different staining of the protoplasm. That
round the definitive egg nucleus remains a rose pink when
stained with carmine, while that round the other nuclei has
a coarse red colour, and a high magnifying power reveals
very clearly the meshes of the spongioplasm of this part
widened out, and dotted with small lumps of nuclein which
are evidently the lecithoblasts.

On examining the small disk-like grains of yolk with a
very high power, and repeatedly changing the focus, the
small stained lecithoblast in its centre is found not to be
a nucleus surrounded with yolk, but a thread passing
through the disk, which is thus like a flat bead threaded on
the filaments of the chromatin spongioplasm. It is not,
however, a smooth thread which passes through the yolk
disk, but it has irregularities consisting of sometimes one,
sometimes two of the minute lumps of nuclein, these being
apparently nothing but slight thickenings of the chromatin
fibres. The yolk disks are, therefore, not nutritive masses
floating freely in the protoplasm of the cells (like starch
grains?), but they remain in organic connection with the
nucleus.

There are many further points of great interest which
we have not yet succeeded in following ; for instance, the
nature of the membrane dividing the cells, its relation to
the spongioplasm, its gradual disappearance so that the
spongioplasm of all the four cells becomes one continuous
whole. We especially wished to find out whether the
threads of spongioplasm of the different cells ran into
one another through the membrane or not. The former
seemed to us to be probable ; if not, we should have to
assume that, on the disappearance of the nuclei of the
nutritive cells, and of the dividing membranes, the threads
of their spongioplasm joined those of the definitive egg-

LITERATURE 313

cell, in order that the ripe egg should constitute an organic
whole with yolk disks threaded on its chromatin filaments.
Sedgwick's observations on the development of the Cape
species of Peripatus l make it probable that these dividing
membranes are but differentiations of the spongework itself.
The disappearance of the membranes would then be nothing
more than the rearrangement of their substance as a
spongework, which must necessarily connect the spongeworks
of the neighbouring cells.

LITERATURE

Besides the well-known text-books of Balfour (Em-
bryology), Claus (Zoology), Gegenbaur, Huxley, and Lang
(Comparative Anatomy), Gerstaecker (Crustacea, Vol V. of
Bronn's Klassen und Ordnungen des Thierreichs), Haeckel
Natiirliche Schopfungsgeschichte), Rollston (forms of Animal
Life), and Zittel, Nicholson, and Lyddeker (Paleontology),
the following are the chief works which have been
consulted : —
BAIRD. — A Monograph of the Family Apodidse. Proceedings

of Zool. Soc., London, 1853.
BARRANDE. — Systeme Silurien de la Boheme. Vol. I. and

Suppt.
BEDDARD.— On the possible Origin of the Malpighian

Tubules in the Arthropoda. Ann. N.ff. (6) IV.
BOURNE (see Lankester).
BRADY. — A Monograph of recent British Ostracoda. Tr.

Linn, Soc. Vol. XXVI.
BRAUER.— Ueber die Entwickelung des Lepidurus pro-

ductus, in Site. Ber. d. K. Acad. d. Ulssensch.

Wien, Vol. LXIX., Pt. I., 1874.

1 Q.J.M.S. Vol. XXVI.

314 LITERATURE

BRAUER. — Beitrage zur Kenntniss der Phyllopoden. In the

same, Vol. LXV., Pt. L, 1872, and further in

1878.
BUCHANAN. — Respiratory Organs in Decapodous Crustacea.

QJ.M.S. Vol. XXIX.
BURMEISTER. — Die Organisation der Trilobiten aus ihren

lebenden Verwandten entwickelt. Berlin, 1843.
CARRIERS. — Die Sehorgane der Thiere. Miinchen und

Leipzig, 1885.
CLAPAREDE. — Les Annelides Chaetopodes du golfe de

Naples. Geneve, 1868, supplement, 1870.
CLAUS. — Zur Kenntniss des Baues und der Entwickelung

von Branchipus stagnalis und Apus cancriformis.

Gottingen, 1873.
,, Untersuchungen zur Erforschungen des Crusta-

ceen Systems. Wien, 1876.
„ Organisation und Entwickelung von Branchipus

und Artemia. Arb. Zool. Inst. Wien, 1886.
,, Neue Beitrage zur Morphologic der Crustaceen.

Arb. Zool. Inst. Wien, 1886.
,, Neue Beobachtungen liber Cyprididen. Zeitsch.

fur Wiss. Zool. Vol. XXIII.
„ Organisation of Nebalia, and Systematic Position

of the Leptostraca. Ann. N.H. (6) III.
DOHRN. — Geschichte des Krebs-Stammes nach embryolo-

gischen, anatomischen und palseontologischen

Quellen. Jenaische Zeitsch. Vol. VI., 1871.
EHLERS. — Die Borstenwiirmer. Leipzig, 1864-68.
FERNALD. — The Relationships of Arthropoda. Studies from

Biol. Lab. Johns Hopkins University. Vol. IV.,

1890.
V. GRABER. — Morphologischen Untersuchungen iiber die

Augen der frei-lebenden marinen Borstenwiirmer.

Bonn, 1879.

LITERATURE 315

GRENACHER. — Untersuchungen iiber das Sehorgan der

Arthropoden. Gottingen, 1879.
GROBBEN. — Die Antennendriise der Crustaceen. Arb. Zool.

Inst. Wien, 1880.
„ Entwickelungsgeschichte der Moina rectirostris.

Arb. Zool. Inst. Wien, 1879.
GRUBE. — Bemerkungen iiber die Phyllopoden. Archiv fur

Naturwiss. Vol. XIX., 1853.
HATSCHEK. — Lehrbuch der Zoologie.
HERRICK — Metamorphosis and Morphology of certain

Phyllopod Crustacea. Bulletin, Denison University,

1885.
HUXLEY. — The Crayfish. International Science Series. —

4th Ed., 1884.
JONES & WOODWARD. — Monograph of the British Palaeozoic

Phyllopoda, 1888.
KINGSLEY. — Notes on the Embryology of Limulus.

QJ.M.S. Vol. XXV.

KORSCHELT UND HEiDER. — Vcrglcichende Entwickelungs-
geschichte der wirbellosen Thiere. Jena, 1891.
KROYER. — Apus glacialis. Naturhist. Tidskrift. 2 Rak.,

1847.
LANKESTER. — The Appendages and Nervous System of Apus

Cancriformis. Q.J.M.Sc. Vol. XXI.
„ Limulus an Arachnid. QJ.M.S., XXI. And

further Vol. XXV.
LANKESTER AND BOURNE.— The minute Structure of the

lateral and central Eyes of Scorpio and Limulus.

QJ.M.S. Vol. XXIII.
LILLJEBORG. — Synopsis Crustaceorum Svecicorum. Reg.

Soc. Sc. Upsaliensi. Tradita die vii. April. 1877.
LUBBOCK. — Notes on some New and Little Known Species

of Freshwater Entomostraca. Trans. Linnean Soc.,

1863.

316 LITERATURE

LUBEOCK. — Origin and Metamorphosis of Insects. Nature

Series, 1890.
MILNE EDWARDS. — Recherches sur 1' Anatomic des Limulcs.

A^ialSc. Nat. 5th Se'rie. T. XVII. Paris, 1873.
MULLER, FRITZ. — Facts for Darwin. Translated by Dallas.
PACKARD. — Monograph of the North American Phyllopoda,

Twelfth Annual' Report of the U.S. Geological

Survey. Washington, 1883.
., Development of Limulus Polyphemus. Soc. Nat.

Hist., Boston, Vol. XI.
„ Anatomy, Histology, and Embryology of Limulus

Polyphemus. Boston, 1880.

And other shorter papers by the same Author.
PELSENEER. — Nervous System of Apus. Quart. Journ.

Micro. Sc. Vol. XXV.
WALCOTT. — The Trilobite. Old and New Evidence relating

to its Organization. Bull. Mus. Comp. Zool.

Vol. VIII. Camb., Mass., U.S., 1 880-81.
WATASE. — On the Morphology of the Compound Eyes of

Arthropods. Studies from Biol. Lab. Johns

Hopkins University. Vol. IV., 1890.
WOODWARD. — British Fossil Crustacea. 1872.
ZADDACH. — De Apodis Cancriformis. Bonn, 1841.
ZENKER. — Monographic der Ostracoden. 1854.
&c., &c.

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INDEX.

PAGE

ABBEY (E. A.) . . .37

PAGE

ATTWELL (H.) . . 20

.BERNARD (J.H.)

PAGE

• 25

ABBOT (F.E.) .

33

AUSTIN (Alfred) . 14

BERNARD (M.) .

. 12

ABBOTT(RCV. E.) 3,13,30,31,
AcLAND(SirH.W.).

33

22

AUTENRIETH (Georg) . 7
AWDRY (F.) . .38

BERNERS (J.) .
BESANT (W.) .

II

4

ADAMS (Sir F. O.) .

28

BACON (Francis) . 19, 20

BETHUNE-BAKER (J. F.)

• 33

ADAMS (Herbert B.).

28

BAINES (Rev. E.) . . 33

BETTANY (G. T.) .

. 6

ADDISON ... 4,

2O

BAKER (Sir S. W.) 28, 30, 37, 38

BICKERTON (T. H.) .

22

AGASSIZ (L.)

3

BALCH (Elizabeth) . . 12

BlGELOW(M. M.) .

. 12

AiNGER(Rev. A.) 4, 16, 20,

33

BALDWIN (Prof. J. M.) . 26

BlKELAS (D.) .

• !7

AlNSLIE(A. D.).

J4

BALFOUR (Rt. Hon. A. J.) 25

BINNIE(RCV. W.) .

• 33

AIRY (Sir G. B.) . 2,

2 7

BALFOUR (F. M.) . . 5, 6

BIRKS (T. R.) . 6, 25,

3°j 33

AITKEN (Mary C.) .
AITKEN (Sir W.) .

20

23

BALFOUR (J. B.) . .6
BALL(V.). ... 38

BjORNSON (B.) .

BLACK (W.) .

• 17
4

ALBEMARLE (Earl of)

3

BALL (W. Platt) . . 6

BLACKBURNE (E.) .

• 3

ALDRICH (T. B.)

J4

BALL(W.W. R.) . . 22

BLACKIE ( J. S.) . 9,

14, 19

ALEXANDER (C. F.) .

20

BALLANCE (C. A.) . . 22

BLAKE (J. F.) .

2

ALEXANDER (T.)

8

BARKER (Lady) . 2, 8, 37

BLAKE (W.) .

3

ALEXANDER (Bishop)

33

BARNARD (C.) . . 27

BLAKISTON (J. R.) .

. 8

ALLBUTT (T. C.)

22

BARNES (W.) ... 3

BLANFORD(H. F.) .

9» 27

ALLEN (G.)

6

BARRY (Bishop). . . 33

BLANFORD (W. T.) .

9, 24

Al.LINGHAM (W.) .

20

BARTHOLOMEW (J. G.) . 3

BLOMFIELD(R.)

• 9

AMIEL(H.F-) .

3

BARTLETT (J.) ... 7

BLYTH (A. W.) .

. ii

ANDERSON (A.).

J4

BARWELL (R.) . . .22

BOHM-BAWERK (Prof.)

. 28

ANDERSON (Dr. McCall) .

22

BASTABLE (Prof. C. F.) . 28

BOISSEVAIN (G. M.) .

. 28

ANDREWS (Dr. Thomas) .

26

BASTIAN (H. C.) . 6, 22

BOLDREWOOD (Rolf ).

• i7

APPLETON (T. G.) .

37

BATESON (W.) ... 6

BONAR (J.)

. 28

ARCHER-HIND (R.D.) .

36

BATH (Marquis of) . . 28

BOND (Rev. J.).

• 3i

ARNOLD, M. 8,14,19,20,21,

30

BATHER (Archdeacon) . 33

BOOLE (G.)

. 26

ARNOLD (Dr. T.)

9

BAXTER (L.) ... 3

BOUGHTON (G. H.) .

• 37

ARNOLD (W. T.)

9

BEESLY (Mrs.) ... 9

BOUTMY (E.) .

. 12

ASHLEY (W. J.).
ATKINSON (J. B.) .
ATKINSON (Rev. J. C.) i,

3

2

38

BENHAM (Rev. W.) . 5, 20, 32
BENSON (Archbishop) 32, 33
BERLIOZ (H. . . 3

BOWEN(H.C).

BOWER (F. O.) .
BRIDGES (J. A.).

• 25
. 19

INDEX.

PAGE"

PAGE

PAGE

BRIGHT (H. A.). . . 9

CLARKE (C. B.). . 9, 28

DlLLWYN (E. A.) . . 17

BRIGHT (John) . . 28, 29

CLAUSIUS (R.) ... 27

DOBSON (A.) ... 4

BRIMLEY(G.) . . 19

CLIFFORD (Ed.) . . 3

DONALDSON (J.) . . 33

BRODIE (Sir B. C.) . .7

CLIFFORD (W. K.) . 19, 26

DONISTHORPE (W.) . . 29

BRODRIBB (W. J.) . 13,37

CLIFFORD (Mrs. W. K.) . 38

DOWDEN (E.) . . 4, 13, 15

BROOKE (Sir J.) . . 3

CLOUGH (A. H.) . 14, 19

DOYLE (Sir F. H.) . . 14

BROOKE (S. A.) 13, 14, 21, 33

COBDEN (R.) . . -29

DOYLE (J. A.) . . . 10

BROOKS (Bishop) . . 33

COHEN (J. B.) ... 7

DRAKE (B.) . .36

BROWN (A. C.) - . .26

COLENSO (J- W.) . . 32

DRUMMOND(Prof. J.) . 34

BROWN (J. A.) . . . i

COLERIDGE (S. T.) . . 14

DRYDEN . . . .20

BROWN (Dr. James) . . 4

COLLIER (Hon. John) . 2

Du CANE (E. F.) . «Q

BROWN (T. E. . . .14

COLLINS (J. Churton) . 19

DuFF(Sir M.E.Grant) 20,29,37

BROWNE (J. H. B.) . . n

COLQUHOUN (F. S.) . . 14

DUNSMUIR (A.). . . 17

BROWNE (Sir T.) . .20

COLVIN (Sidney) . 4, 20

DUNTZER (H.) . . . 4, 5

BROWNE (W. R.) . . 27

COMBE (G.) ... 8

DUPRE(A.) ... 7

BRUNTON(Dr.T.Lauder) 22, 33

CONGREVE (Rev. J.). . 33

DYER(L.). . . . i

BRYCE (James) . . 9, 28, 37

CONWAY (Hugh) . . 17

EADIE (J.). . . 4, 30, 31

BUCHHEIM (C. A.) . . 20

COOK(E.T.) ... 2

EASTLAKE (Lady) . . 3*

BUCKLAND (A.). . . 5

COOKE (C. Kinloch) . . 24

EBERS(G.) ... 17

BUCKLEY (A. B ) . . 9

COOKE (J. P.) . . 7, 34

EDGEWORTH (Prof. F. Y.). 28

BUCKNILL (Dr. J. C.) . 22

CORBETT (J.) . . 4, 17, 38

EDMUNDS (Dr. W.) . . 22

BUCK-TON (G. B.) . . 40

CORFIELD (W. H.) . . II

EDWARDS-MOSS (Sir J. E.) 30

BUNYAN . . .4, 19, 20

CORRY (T. H.) . . .6

EiMER(G.H.T.) . . 6

BURGON(J.W.) . . 14

BURKE (E.) ... 28

COTTERILL(J.H.) . . 8

COTTON (Bishop) . . 34

ELDERTON (W.A.) . . 9
ELLERTON (Rev. J.) . . 34

BURN (R.). ... i

COTTON (C.) . . .12

ELLIOT (Hon. A.) . . 29

BURNETT (F. Hodgson) . 17

COTTON (J-S.) . . . 29

ELLIS (T.). ... 2

BURNS ... 14, 20

COUES (E.) . . . 40

EMERSON (R. W.) . 4, 20

BURY(J. B.) ... 9

COUR'1.IOFE(W.J.) . . 4

EVANS (S.) . . .14

BUTCHER (Prof. S. H.) 13,19,36

COWELL(G.) . . .23

EVERETT (J. D.) . .26

BUTLER (A. J. . . -37

COWPER . . . .20

FALCONER (Lanoe) . . 17

BUTLER (Rev. G.) . . 33

Cox(G.V.) ... 9

FARRAR (Archdeacon) 5, 30, 34

BUTLER (Samuel) . . 14

CRAiK(Mrs.)i4, 17, 19, 20, 37, 38

FARRER(SirT. H.) . . 29

BUTLER (W. Archer) . 33

CRAIK (H.) . . 8, 29

FAULKNER (F.). . . 7

BUTLER (Sir W. F.) . . 4

CRANE (Lucy) • • 2, 39

FAWCETT (Prof. H.) . 28, 29

BYRON . . . .20

CRANE (Walter) . 39

FAWCETT (M. G.) . 5, 28

CAIRNES (J- E.) • • 29

CRAVEN (Mrs. D.) . . 8

FAY (Amy) . . • . 24

CALDECOTT (R.) .12, 38, 39

CRAWFORD (F. M.) . . 17

FEARNLEY(W.) . . 27

CALDERWOOD (Prof. H.)

CREIGHTOX (Bishop M.) 4, 10

FEARON (D. R.) . .8

8> 25. 26, 33

CRICHTON-BROWNE(SirJ.) 8

FERREL(W.) ... 27

CALVERT (Rev. A.) . . 31
CAMERON (V. L.) . . 37

CROSS (J. A.) . .30
CROSSLEY(E.) ... 2

FERRERS (N. M.) . . 27
FESSENDEN (C.) . . 26

CAMPBELL (J. F.) . . 37
CAMPBELL (Dr. J. M.) . 33

CROSSLEY (H.) . . .37
GUMMING (L.) . . . 26

FiNCK(H.T.) ... i
FISHER (Rev. O.) . 26, 27

CAMPBELL (Prof. Lewis) 5, 13

CUNNINGHAM (C.) . . 28

FISKE (J-)- 6, 10, 25, 29, 34

CAPES (W.W.). . . 13

CUNNINGHAM (Sir H. S.) . 17

FISON(L.). ... i

CARLES (W. R.) . . 37

CUNNINGHAM (Rev. J.) . 31

FITCH G. G.) ... 8

CARLYLE(T.) ... 3

CUNNINGHAM (Rev. W)3i, 33, 34.

FITZ GERALD (Caroline) . 14

CARMARTHEN (Lady) . 17

CUNYNGHAME (Sir A. T.) . ' 24

FITZGERALD (Ed ward) 14,20

CARNARVON (Earl of) . 36

CURTEIS (Rev. G. H.) 32, 34

FITZMAURICE (Lord E.) . 5

CARNOT (N. L. G.) . . *7

DAHN (F.) . . . 17

FLEAY(F. G.) ... 13

CARPENTER (Bishop) . 33

DAKYNS (H. G.) . . 37

FLEISCHER (E.). . . 7

CARR(J.C.) ... a

DALE (A. W. W.) . . 31

FLEMING (G.) . . .17

CARROLL (Lewis) . 26, 38

DALTON (Rev. J. N.) . 37

FLOWER (Prof. W. H.) . 39

GARTER (R. Brudenell) . 23

DANTE . . .3, 13, 37

FLUCKIGER(F.A.) . . 23

CASSEL(Dr. D.) . .9
CAUTLEY(G. S.) . . 14

DAVIES (Rev. J. LI.). 20,31,34
DAVIES(W.) ... 5

FORBES (A.) . . 4, 37
FORBES (Prof. G.) . . 3

CAZENOVE (J. G.) . . 33

DAWKINS(W. B.) . . i

FORBES (Rev. G. H.) . 34

CHALMERS (J. B.) . .8
CHALMERS (M. D.) . . tg
CHAPMAN (Elizabeth R.) . 14
CHASSERESSE (Diana) . 30

DAWSON (G. M.) . . 9
DAWsoN(SirJ.W.) . . 9
DAWSON (J.) ... i
DAY(L. B.) ... 17

FOSTER (Prof. M.) . 6, *7

FOTHERGILL (Dr. J. M.) 8,23

FOWLE (Rev. T. W.). 29, 34
FOWLER (Rev. T.) . 4, 25

CHERRY (R. R.) . . ia

DAY(R. E.) ... 26

FOWLER (W.W.) . . 24

CHEYXE (C. H. H.) . .a

DEFOE (D.) . . 4, 20

Fox (Dr. Wilson) . . 23

CHEYWE (T. K.) . . 30

DEIGHTON (K.). . . 15

FOXWELL (Prof. H. S) . 28

CHRISTIE (J.) ... 23
CHRISTIE (W. D.) . . 20

DELAMOTTE (P. H.). . 2
DELL(£.C.) ... 12

FRAMJI (D.) . .10
FRANKLAND (P. F.) . i

CHURCH (Prof. A. H.) . 6

DE MORGAN (M.) : . 39

FRASER (Bishop) . . 34

CHURCH (Rev. A. J.) 4,30,37

DE VERB (A.) . . .20

FRASER-TYTLER (C. C.) . 14

CHURCH (F. J.). . 20, 37

DICEY (A. V.) . . 12, 29

FRAZER (J. G.) . . . i

CHURCH (Dean) 3,4,13,19,31,33

DICKENS (C.) . . 5, 17

FREDERICK (Mrs.) . . 8

O.ARK (J. W.) ... 20

DIGGLE (Rev. J. W.). . 34

FREEMAN (Prof. E. A.)

CLARK (L.) ... 2

DILKE (Ashton W.) . . 19

2, 4, 10, 29, 32

CLARK (S.) . . . 3

DILKE (Sir Charles W.) . 29

FRENCH (G. R.) . . 13

42

INDEX.

PAGE

PAGE

PAGE

FRIEDMANN (P.) . . 3

HARRISON (Miss J.) . . i

JONES (F.). ... 7

FROST (A. B.) . . . 38

HARTE (Bret) ... 17

KANT .... 25

FROUDE (J. A.)- • • 4

HARTIG (Dr. R.) . . 6

KARI . . . -39

FUI.LERTON (W. M.) . 37

HARTLEY (Prof. W. N.) . 7

K.AVANAGH(Rt.Hn.A.M.) 4

FURNISS (Harry) . . 38

HARWOOD(G.). .21,29,32

KAY (Rev. W.). . . 31

FURNIVALL (F. J.) . . 14

HAYES (A.) ... 14

KEARY (Annie). 10,18,39

FYFFE (C. A.) . . . 10

HEADLAM(W.). . . 36

KEARY (Eliza) ... 39

FYFE(H. H.) ... 9

HELPS (Sir A.) . . .21

KEATS . . .4, 20, 21

GAIRDNER (J.) ... 4
GALTON (F.) . . i, 27

HEMPEL (Dr. W.) . . 7
HERODOTUS . . . 3?

KELLNER (Dr. L.) . . 25
KELLOGG (Rev. S. H.) . 34

GAMGEE (Arthur) . . 27

HERRICK . . . .20

KEMPE(A. B.) . . . 26-

GARDNER (Percy) . . i

HERTEL(Dr.) ... 8

KENNEDY (Prof. A. B. W.)

GARNETT (R.) . . .14

HILL (F. Davenport). . 29

KENNEDY (B. H.) . . 3t

GARNETT (W.) ... 5

HILL (O.) .... 29

KEYNES (J. N.). . 26, 28

GASKELL (Mrs.) . . 12

HIORNS(A. H.) . . 23

KlEPERT(H.) ... 9

GASKOIN (Mrs. H.) . . 30

HOBART (Lord) . . 21

KILLEN (W. D.) . . 32

GEDDES(W. D.) . 13,37

HOBDAY (E.) ... 9

KINGSLEY (Charles) . 4, 8, 10,

GEE (W. H.) . . 26, 27
GEIKIE (Sir A.). . 4, 9, 27

HODGSON (Rev. J. T.) . 4
HOFFDING (Prof. H.) . 26

11,12,13,15,18,21,24,32,37,39
KINGSLEY (Henry) . 20, 38

GENNADIUS (J.) . . 17

HOFMANN (A. W.) . . 7

KIPLING (J. L.). . . 38

GiBBiNs(H.deB.) . . 10

HOLE (Rev. C.). . 7, ™

KIPLING (Rudyard) . . 18

GIBBON (Charles) . . 3

HOLIDAY (Henry) . . 38

KIRKPATRICK (Prof.) . 34

GILCHRIST (A.). . . 3

HOLLAND (T. E.) . 12,29

KLEIN (Dr. E.). . 6,23

GILES (P.). ... 25

HOLLWAY-CALTHROP(H.) 38

KNIGHT (W.) ... 14

OILMAN (N. P.) . . 28

HOLMES (O. W., junr.) . 12

KUENEN (Prof. A.) . . 30

GILMORE (Rev. J.) . . 13
GLADSTONE (Dr. J. H.) 7, 8
GLADSTONE (W. E.) . . 13

HOMER ... 13, 36
HOOKER (Sir J. D.) . 6, 37
HOOLE (C. H.) . . . 30

KYNASTON (Rev. H.) 34, 37
LABBERTON (R. H.). . 3
LAFARGUE(P.). . . 18

GLAISTER (E.) . . . 2, 8

HOOPER (G.) ... 4

LAMB. . . .4, 20, 21

GODFRAY(H.) ... 3

HOOPER (W. H.) . . 2

LANCIANI (Prof. R.). . 2

GODKIN (G. S.). . . 5

HopE(F.J.) ... 9

LANDAUER (J.). . . 7

GOETHE . . . 4, 14

HOPKINS (E.) ... 14

LANDOR . . . 4, 20

GOLDSMITH 4, 12, 14, 20, 21

HOPPUS (M. A. M.) . . 18

LANE-POOLE (S.) . . 20

GOODALE (Prof. G. L.) . 6

HORACE ... 13, 20

LANFREY (P.) ... 5

GOODFELLOW (J.) . .11

HORT (Prof. F. J. A.). 30, 32

LANG (Andrew). 2,12,21,36

GORDON (General C. G.) . 4

HORTON (Hon. S. D.) . 28

LA^JG (Prof. Arnold) . . 39

GORDON (Lady Duff) . 37
GOSCHEN (Rt. Hon. G. J.)- 28

HOVENDEN (R. M.) . . 37
HOWELL (George) . . 28

LANGLEY (J. N.) . . 27
LANKESTER (Prof. Ray) 6, 21

GOSSE (Edmund) . 4, 13

HOWES (G. B.) . . . 40

LASLETT(T.) ... 6

Gow(J.) .... i

HOWITT(A. W.) . . I

LEAF (W.). . . 13, 36

GRAHAM (D.) . -14

HOWSON (Very Rev. J. S.) 32

LEAHY (Sergeant) . . 30

GRAHAM (J.W.) . . 17

HOZIER (Col. H. M.). . 24

LEA(M.) . . . . 18

GRAND'HOMME (E.) . . 8

HUBNER (Baron) . . 37

LEE (S.) ... 20, 37

GRAY (Prof. Andrew) . 26

HUGHES (T.) 4, 15, 18, 20, 37

LEEPER (A.) . . -37

GRAY (Asa) ... 6

HULL(E.). . . . 2, 9

LEGGE (A. 0.) . . 10, 34

GRAY ... 4, 14, 21

HULLAH (J.) . . 2, 20, 24

LEMON (Mark) . .20

GREEN (J. R.) . 9, 10, 12, 20
GREEN (Mrs. J. R.) . 4, 9, 10

HUME(D.) ... 4
HuMPHRY(Prof.SirG.M.) 28,39

LESLIE (A.) ... 38
LETHBR^DGE (Sir Roper) . 10

GREEN (W.S.). . . 37

HUNT(W.) ... 10

LEVY (Amy) . .18

GREENHILL (W. A.) . . 20

HUNT(W.M-). . . 2

LEWIS (R.) ... 13

GREENWOOD (J. E.) . . 39

HUTTON (R. H.) . 4. 21

LiGHTFOOT(Bp.)2i,3o,3i,33,34

GRIFFITHS (W. H.) . . 23

HUXLEY (T.) 4, 21, 27,28,29,40

LlGHTWOOD (J. M.) . . 12

GRIMM .... 39

IDDINGS(J. P.). . . 9

LINDSAY (Dr. J. A.) . . 23

GROVE (Sir G.). . 9, 24

ILLINGWORTH (Rev. J. R.) 34

LOCKYER (J. N.) . 3, 7, 27

GUEST (E.) . 10

INGRAM (T. D.) . . 10

LODGE (Prof. O.J.) . 21,27

GUEST (M. J.) . . . 10
GUILLEMIN (A.) . 26, 27
GUIZOT (F. P. G.) . . 5

IRVING (J.) ... 9
IRVING (Washington) . 12
JACKSON (Helen) . . 18

LOEWY(B.) . . . -2-6

LOFTIE (Mrs. W. J.). . 2
LONGFELLOW (H. W.) . ao

GUNTON (G.) . . . 28

; ACOB (Rev. J. A.) . . 34

LONSDALE (J.) . . 20, 37

HALES (J. W.) . . 16, 20

AMES (Henry). . 4, iS, 21

LOWE (W. H.) . . . 30

HALLWARD (R. F.) . . 12

' AMES (Rev. H. A.) . . 34

LOWELL (J. R.). . 15, 21

HAMERTON (P. G.) . 2, 21

" AMES (Prof. W.) . . 26

LuBBOCK(Sir J.) 6, 8, 21, 22, 40

HAMILTON (Prof. D. J.) . 23

' ARDINE (Rev. R.) . . 26

LUCAS (F.) . . 15

HAMILTON (J.). . . 34

] BANS (Rev. G. E.) . 34, 37

LUPTON (S.) . . 7

HANBURY (D.) . . 6, 23

' EBB (Prof. R. C.) . 4, 10, 13

LYALL (Sir Alfred) . 4

HANNAY (David) . . 4
HARDWICK (Archd. C.) 31, 34

ELLETT (Rev. J. H.) . 34
; ENKS (Prof. Ed.) . . 29

LYTE(H. C.M.) . 10
LYTTON (Earl of) . 18

HARDY (A. S.) ... 17

ENNINGS'(A. C.) . 10,30

MACALISTER (D.) . 23

HARDY (T.) . . .17

EVONS (W. S.). 4, 26, 28, 29

MACARTHUR (M.) . 10

HARE (A. W.) . . . 20

EX-BLAKE (Sophia). . 8

MACAULAY (G. C.) . 36

HARE(J.C.) . . 20,34
HARPER (Father Thos.) 25,34

OHNSON (Amy) . . 27
OHNSON (Samuel) . . 13

MACCOLL (Norman) . 14
M'CosH (Dr. J.) . 25, 26

HARRIS (Rev. G. C.). . 34

ONES (H. Arthur) . . 15

MACDONALD (G.) . 16

HARRISON (F.). . 4, 5, 21

ONES (Prof. D. E.) . . 27

MACDONELL (J.) . 29

INDEX.

43

PAGE

PAGE

PAGE

MACKAIL(J. W.) . . 37

MOULTON (L. C.) . 15

POOLE (R. L.) . . . n

MACKENZIE (Sir Morell) . 23

MUDIE (C. E.) . . 15

POPE . . . . 4, 20

MACLAGAN (Dr. T.). . 23

MuiR(M. M.P.) . 7

POSTE (E.) . . 27, 36

MACLAREN (Rev. Alex.) . 34

MULLER (H.) . . 6

POTTER (L.) . .22

MACLAREN (Archibald) . 39

MULLINGER (J. B.) . II

POTTER (R.) . .35

MACLEAN (W. C.) . .23

MURPHY (J.J.). . 26

PRESTON (T.) ... 27

MACLEAR(Rev.Dr.G.F.) 30,32

MURRAY (D.Christie) . 18

PRICE (L. L. F. R.) . . 28

M'LENNAN (J. F.) . i

MURRAY (E. C. G.) . . 38

PRICKARD (A. O.) . . 22

M'LENNAN (Malcolm) . 18

MYERS (E.) . . 15, 36

PRINCE ALBERT VICTOR . 37

MACMILLAN(RCV. H.) 22,35,38

MYERS (F. W. H.) . 4, 15, 22

PRINCE GEORGE . . 37

MACMILLAN (Michael) 5, 15

MYLNE (Bishop) . . 35

PROCTER (F.) ... 32

MACNAMARA (C.) . 23

NADAL (E. S.) . . . 22

PROPERT (J. L.) . . 2

MACQUOID (K. S.) . . 18

NETTLESHIP (H.). . . 13

RADCLIFFE(C. B.) . . 3

MADOC(F.) . 18

NEWCASTLE (Duke and

RAMSAY (W.) ... 7

MAGUIRE(J. F.) . . 39

Duchess) . . .20

RANSOME(C.) ... 13

MAHAFFY(Prof. J. P.)

NE\VCOMB(S.) ... 3

RATHKONE (W.) . . 8

2, II, I^. 22, 25, 35, 38

NEWTON (Sir C. T.) . . 2

RAWLINSON (W.G.). . 2

MAITLAND (F. W.) . 12, 29

NlCHOL(J.) . . 4, 13

RAWNSLEY (H. D.) . . 15

MALET(L.) . 18

NOEL (Lady A.) . . 18

RAY(P.K.) ... 26

MALORY (Sir T.) . .20

NORDENSKIOLD (A. E.) . 38

RAYLEIGH (Lord) . . 27

MANSFIELD (C. B.) . .7

NORGATE (Kate) . .11

REICHEL (Bishop) . . 35

MARKHAM (C. R.) . . 4

NORRIS (W. E.) . . 18

REID(J. S.) ... 37

MARRIOTT (J. A. R.). . 5

NORTON (Charles Eliot) 3, 37

REMSEN (I.) ... 7

MARSHALL (Prof. A.) . 28

NORTON (Hon. Mrs.) 15,18

RENDALL(RCV. F.) . 31,35

MARSHALL (M. P) . . 28

OLIPHANT(MrS.M.O.W.)

RENDU(M. leC.) . . 9

MARTEL(C.) ... 24
MARTIN (Frances) . 3, 39

4, ii, 13, 19, 20, 39
OLIPHANT (T. L. K.) 22, 25

REYNOLDS (H. R.) . .35
REYNOLDS (J. R.) . .23

MARTIN (Frederick)-. . 28

OLIVER (Prof. D.) . .6

REYNOLDS (O.). . .11

MARTIN (H. X.) . . 40

OLIVER (Capt. S. P.). . 38

RICHARDSON (B. W.) n, 23

MARTINEAU (H.) . ^ . 5

OMAN(C.W.) ... 4

RICKEY (A. G.). . . 12

MARTINEAU (J.) . " . 5

OSTWALD (Prof.) . . 7

ROBINSON (Preb. H. G.) . 35

MASSON(D.) 4,5,15,16,20,22,26

OTTE(E.C.) . u

ROBINSON (J. L.) . . 24

MASSON (G.) . . 7, 20
MASSON (R. O.) . . 16

PAGE(T.E.) ... 31
PALGRAVE (Sir F.) . .11

ROBINSON (Matthew) . 5
ROCHESTER (Bishop of) . 5

MATURIN (Rev. W.). . 35

PALGRAVE (F. T.)

ROCKSTRO (W. S.) . . 4

MAUDSLEY (Dr. H.) . . 26

2, 15, 16, 20, 21, 33, 39

ROGERS (J. E. T.) .11, 28, 29

MAURICE (Fredk. Denison)

PALGRAVE (R. F. D.) . 29

ROMANES (G.J.) . . 6

8, 22, 25, 30, 31, 32, 35
MAURICE (Col. F.) . 5,24,29

PALGRAVE (R. H. Inglis) . 28
PALGRAVE (W. G.) 15, 29, 38

ROSCOE (Sir H. E.) . .7
ROSCOE (W. C.) . . 15

MAX MULLER (F.) . . 25

PALMER (Lady S.) . . 19

ROSEBERY (Earl of) . .4

MAYER (A.M.). . . 27

PARKER (T.J.). . 6,39

ROSENBUSCH(H.) . . 9

MAYOR (J. B.) . . . 31

PARKER (W. N.) . . 40

Ross (P.) .... 19

MAYOR (Prof. J. E. B.) . 3, 5

PARKINSON (S.) . . 27

ROSSETTI (C. G.) . 15, 39

MAZINI (L.) ... 39

PARKMAN (F.) . . .11

ROUTLEDGE (J.) . . 29

M'CORMICK(W.S.). . I3

PARSONS (Alfred) . . 12

Ro\VE(F.J.) . 16

MELDOLA (Prof. R.). 7, 26, 27

PASTEUR (L.) ... 7

RUCKER (Prof. A. W.) 7

MENDENHALL (T. C.) . 27

PATER (W. H.) . 2, 19, 22

RUM FORD (Count) . . 22

MERCIER (Dr. C.) . . 23

PATERSON (J.) . . .12

RUSHBROOKE (W. G.) . 31

MERCUR(Prof. J.) . .24

PATMORE (Coventry) 20, 39

RUSSELL (Dean) . . 35

MEREDITH (G.). . . 15

PATTESON (J. C.) . . 5

RUSSELL Sir Charles) . 29

MEREDITH (L. A.) . . 12

PATTISON (Mark) . 4, 5, 35

RUSSELL (W. Clark) . 4, 19

MEYER (E. von) . . 7

PAYNE(E.J-) . . 10, 29

RYLAND (F.) . 13

MlALL(A.) ... 5

PEABODY(C. H.) . 8, 27

RYLE (Prof. H. E.) . . 30

MlCHELET (M.) . . II

PEEL(E.). ... 15

ST. JOHNSTON (A.) .19, 38, 39

MiLL(H.R.) ... 9

PEILE(J.). ... 25

SADLER (H.) ... 2

MILLER (R. K.). . . 3

PELLISSIER (E.) . . 25

SAINTSBURY (G.) . 4, 13

MILLIGAN (Rev. W.). 31, 35

PENNELL(J.) ... 2

SALMON (Rev. G.) . . 35

MILTON . . 13, 15, 20

PENNINGTON (R.) . 9

SAXDFORD (M. E.) . .5

MINTO (Prof. W.) . 4, X8

PEN-ROSE (F. C.) . . i, 3

SANDYS (J. E.) . . . 38

MITFORD (A. B.) . . 18

PERRY (Prof. J.) . . *7

SAYCE(A.H.) . . . n

M iv ART (St. George). . 28

PETTIGREW (J. B.) . 6, 28, 40

SCHAFF (P.) . . -30

MlXTER(W.G-) . . 7

PHILLIMORE (J. G.) . . 12

SCHLIEMANN (Dr.) . . 2

MOHAMMAD . . .20

PHILLIPS (J. A.) . . 23

SCHORLEMMER (C.) . . 7

MOLESWORTH (Mrs.) . 39

PHILLIPS (W. C.) .2

SCOTT (D. H.) ... 6

MOLLOY (G.) . . .26

PICTON (J. A.) . . .22

SCOTT (Sir W.). . 15,20

MONAHAN (J. H.) . . 12

PlFFARD (H. G.) . . 23

SCRATCHLEY (Sir Peter) . 24

MONTELIUS (O.) . . I

PLATO . . . .20

SCUDDER (S. H.) . . 40

MOORE (C. H.). . . 2

PLUMPTRE (Dean) . . 35

SEATON (Dr. E. C.) . . 23

MOORHOUSE (Bishop) . 35

POLLARD (A. W.) .. . 37

SEELEY (J. R.). . . «

MORISON (J. C.) . . 3, 4

PoLLOCK(SirFk., 2nd Bart.) 5

SEILER (Dr. Carl) . 23, 28

MORLEY (John). 3, 4, 16, 22

PoLLOCK(Sir F.,Bart.) 12,22,29

SELBORNE Earl of) 12,20,32,33

MORRIS (Mowbray) . . 4

POLLOCK (Lady) . . 2

SELLERS (E.) . . . 2

MORRIS (R.) . . 20, 25

POLLOCK (W. H.) . . 2

SERVICE (J.) . . 32, 35

MORSHEAD (E. D. A.) . 36

POOLE (M. E.) . . . 22

SEWELL (E. M.) . . n

44

INDEX.

PAGE

PAGE

PAGE

SHAIRP (J. C.) . 4, 15

TANNER (H.) . . . i

WAI:D (A. W.) . . 4, 13, 20

SHAKESPEARE 13, 15, 20, 21

TAVERNIER (J. B.) . . 38

WARD (H. M.) . . .6

SHANN (G.) . 8, 27

TAYLOR (Franklin) . . 24

WARD(S.). . . . 16

SHARP (W.) . . 5

TAYLOR (Isaac). . 25, 35

WARD(T.H.) . 16

SHELLEY . . 15, 21

TAYLOR (Sedley) . 24, 27

WARD (Mrs. T. H.) . 19, 39

SHIRLEY (W. N ) . .35

TEGETMEIER (W. B.) . 8

WARD(W.) . . 5,32

SHORT-HOUSE (J. H.) . 19

TEMPLE (Bishop) . . 35

WARINGTON (G.) . . 36

SHORTLAND (Admiral) . 24

TEMPLE (Sir R.) .4

WATERS (C. A.) . . 28

SHUCHHARDT (Carl). . 2

TENNANT (Dorothy). . 38

WATERTON (Charles) 24, 38

SHUCKBURGH (E. S.) 11,36

TENNIEL . . . .38

WATSON (E.) ... 5

SHUFELDT(R. W.) . . 40

TENNYSON . 14, 16, 21

WATSON (R. S. . - 3?

SIBSON (Dr. F.) . . 23

TENNYSON (Frederick) . 16

WEBB (W. T.) . . .16'

SIDGWICK (Prof. H.) 26,28,29

TENNYSON (Hallam). 12, 39

WEBSTER (Mrs. A.) . . 39

SlME (J.) . . . 9, 10

THOMPSON (D 'A. W.) . 6

WELBY-GREGORY (Lady) . 32

SIMPSON (Rev. W.) . . 32

THOMPSON (E.). . . 10

WELLDON (Rev. J. E. C.) . 36

SKEAT (W.W.) . . 13

THOMPSON (S. P.) . . 27

WESTCOTT (Bp.) 30, 31, 32, 36

SKRINE (J. H.). . 5,15

THOMSON (A. W.) . . 8

WESTERMARCK (E.). . i

SLADE (J. H.) ... 8

THOMSON (Sir C. W.) . 40

WETHERELL (J.) . . 25

SLOMAN (Rev. A.) . . 31

THOMSON (Hugh) . . 12

WHEELER (J. T.) . .11

SMART (W.) . . .28
SMALLEY(G.W.) . . 22

THOMSON (Sir Wm.) 24, 26, 27
THORNE (Dr. Thorne) . 23

WHEWELL(W.). . . 5
WHITE (Gilbert) . . 24

SMETHAM (J. and S.) . 5

THORNTON (J.). . . 6

WHITE (Dr. W. Hale) . 23

SMITH (A.) . . .20

THORNTON (W. T.) 26, 29, 37

WHITE (W.) ... 27

SMITH (C.B.).. . . 16

THORPE (T. E.). . . 7

WHITHAM (J. M.) . . 8

SMITH (-Goldwin) . 4, 5, 29

THRING(E.) . . 8,22

WHITNEY (W. D.) . . 8

SMITH (H.) . 16

THRUPP (J. F.) . . . 30

WHITTIER(J. G.) . 16,22

SMITH (J.) ... 6

THUDICHUM (J. L. W.) . 7

WICKHAM (Rev. E. C.) . 36

SMITH (Rev. T.) . . 35

THURSFIELD (J. R.) . .4

WlCKSTEED (P. H.) . 28, 30

SMITH (W. G.) . . .6

TODHUNTER (I.) . • 5, 8

WlEDERSHEIM (R.) . . 40

SMITH (W. S.) ... 35

TORRENS (W. M.) . . 5

WlLBRAHAM (F. M.). . 32

SOMERVILLE (Prof. W.) . '6

TOURGENIEF (L S.) . . 19

WILK!NS (Prof. A. S.) 2, 13, 36

SOUTHEY .... 5

SPENDER (J. K.) . . 23

TouT(T. F.) ... ii
TOZER(H. F.) ... 9

WILKINSON (S.) . . 24
WILLIAMS (G. H.) . . 9

SPENSER . . . .20

TRAILL (H. D.). . 4, 29

WILLIAMS (Montagu) . 5

SPOTTISWOODE (W.). . 27

TRENCH (Capt. F.) . . 29

WILLIAMS (S. E.) . . 13

STANLEY (Dean) . . 35
STANLEY (Hon. Maude) . 29

TRENCH (Archbishop) . 35
TREVELYAN (Sir G. O.) . n

WlLLOUGHBY (F.) . . 39

WILLS (W. G.) . . . 16

STATHAM (R.) . . .29

TRIBE (A.). ... 7

WILSON (A. J.) . . . 29

STEBBING (W.) . . . 4

TRISTRAM (W. O.) . . 12

WILSON (Sir C.) . . 4

STEPHEN (C. E.) . . 8

TROLLOPE (A.) ... 4

WILSON (Sir D.) . i, 3, 13

STEPHEN (H.) . . .13

TRUMAN (J.) . . . 16

WILSON (Dr. G.) . 4, 5, 22

STEPHEN (Sir J. F.) n, 13, 22

TUCKER (T. G.) . . 36

WILSON (Archdeacon) . 36

STEPHEN (J. K.) . . 13

TULLOCH (Principal). . 35

WILSON (Mary). . . 13

STEPHEN (L.) ... 4

TURNER (C. Tennyson) . 16

WINGATE (Major F. R.) . 24

STEPHENS (J. B.) . . 16

TURNER (G.) . . . i

WlNKWORTH (C.) . . 5

STEVENSON (J. J.) . .2

TURNER (H. H.) . . 27

WOLSELEY (Gen. Viscount) 24

STEWART (A.) . . .39

TURNER (J. M.W.) . . 12

WOOD (A. G.) . . . 16

STEWART (Balfour) 26, 27, 35

TYLOR(E.B.) . . . i

WOOD (Rev. E. G.) . . 36

STEWART (S. A.) . . 6

TYRWHITT (R. St. J.) 2, 16

WOODS (Rev. F. H.). . i

STOKES (Sir G. G.) . . 27

VAUGHAN (C. J.) 3I,32,35,36

WOODS (Miss M. A.). 17, 33

STORY (R. H.) ... 3
STONE (W.H.). . . 27

VAUGHAN (Rev. D. J.) 20, 36
VAUGHAN (Rev. E. T.) . 36

WOODWARD (C. M.) . .8
WOOLNER (T.) . . . 16

STRACHEY (Sir E.) . . 20

VAUGHAN (Rev. R.). . 36

WORDSWORTH . 5, 14, 16, 21

STRACHEY (Gen. R.). . 9

VELEY(M.) ... 19

WORTHEY (Mrs.) . . 19

STRANGFORD(Viscountess) 38
STRETTELL (A.) . . 16

VENN (Rev. J.). . 26,36
VERNON (Hon. W. W.) . 13

WRIGHT (Rev. A.) . . 31
WRIGHT (C. E. G.) . . 8

STUBBS (Rev. C. W.). . 35

VERRALL (A. W.) . 13, 36

WRIGHT (J.) ... 21

STUBBS (Bishop) . . 31

VERRALL (Mrs.) . . i

WRIGHT (L.) . . .27

SUTHERLAND (A.) . . 9

WAIN (Louis) . . . 39

WRIGHT (W. Aldis) 8, 15, 20, 31

SYMONDS (J. A.) . .4

WALDSTEIN (C.) . . 2

WURTZ (Ad.) ... 7

SYMONDS (Mrs. J. A.) . 5

WALKER (Prof. F. A.) . 28

WYATT (SirM. D.) . . 2

SYMONS(A.) . 16

WALLACE (A. R.) .6, 24, 28

YONGE (C. M.) 5, 6, 8, 10, ii,

TAIT (Archbishop) . - .35

WALLACE (Sir D. M.) . 29

19, 21, 25, 30, 39

TAiT(C.W.A.) . . ii

WALPOLE (S.) . . .29

YOUNG (E.W.) . . 8

TAIT (Prof. P. G.) 26, 27, 35

WALTON (I.) ... 12

ZIEGLER (Dr. E.) . .23

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