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A specimen of the 
Spitzbergen variety of 
Apus glacialis (X ro), 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 
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ek essere 


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oes 
SBUXNM ATURE SERIES 
Ve Zoal- 


Aiie APODIDA: 


Pew ORPHOLOGICAL STUDY 


BY 


foe RY MEYNERS BERNARD 


M.A.- CANTAB. 


WITH SEVENTY-ONE ILLUSTRATIONS 


Dondon 
Vee ert AN: AN bE Oo, 
AND NEW YORK 


1892 


The Right of Translation and Reproduction is Reserved 


RIcHARD CLAY AND Sons, LIMITED, 
LONDON AND BUNGAY. 


TO’ THE MEMORY (OF 


Bike Ari, bo REE) Wee aa 


TOO SOON LOST TO: SCIENCE 


AND 


EO; THE MANY FRIENDS WHO LOVED 


HIM AS A MAN 


TABLE 


INTRODUCTION 


PAR i 


. THE OUTER BODY FORM 


THE APPENDAGES ... 
THE MUSCULATURE .. 
THE NERVOUS SYSTEM 
THE SENSORY ORGANS. . 
THE ALIMENTARY CANAL . 


THE CIRCULATORY SYSTEM 


. OBJECT AND LINE OF ARGUMENT 


13-9729 


Ole CONEEDEES 


THE EXCRETORY AND OTHER GLANDS 


REPRODUCTION ..-.. 2065. % 


DEVELOPMENT—THE NAUPLIUS 


PAGE 


eT 


x TABLE OF CONTENES 


PAT Lh 

PAGE 

§ XII. RELATION OF APUS TO THE OTHER CRUSTACEA— 
TEMURUIS V5 5 Se a ie ae 8 eas 2 Soe Sa ee ae 
Sree el re MERTEOBETES 6.5. (0 tec bp Pelee bs) Roe Comm ee 
SOG -LHE: EUR VPERRIDA = Veo. 8 gs eet aia ec oie ey 
S XV. fHE NEw CLassipicATION PROPOSED™ 55. =z. 7y 8252 
SUNT, sPERIPATUSTAND: THE “TRACHEAPA (5 /<).) 0-24. fateh seu 
APPENDICES ?.; 3 oof ates as SS aes eee SOU 
LITERATURE 313 


[IN FRODUC TION 


AMONG the numerous interesting zoological 
specimens brought back by Professor Kiikenthal 
from East Spitzbergen on his return from the Bremen 
Expedition (1889),’ 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 
Leché, 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, was Professor 
Kiikenthal’s companion during this expedition. He died shortly after 
his return. 


Xi 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 iat 
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 Apodide 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 xlil 


of each organ and system of organs, to find out as far 
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! of preparing a comparative 
anatomy of the Apodide 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 Mature, reprinted in Appendix V. 


XIV 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. [ intended to 
content myself with an endeavour to show that in 
the Apodide 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 see, 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 XV 


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 
sudy. Whe appeal made in the. second, part to 
palzozoic 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 Xiphosuride 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 


XV1 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, [ 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 Apodidz 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 XVil 


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 
Kikenthal 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. 


HH. WEB. 


Streatham, 1892. 


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THE -APODID A. 


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OBJECT AND LINE OF ARGUMENT 


THE Apodide 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 
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2 = THE APODIDZS 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 Apodide 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 Apodidz 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” sie 
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. 


sei ie OBJECT AND LINE OF ARGUMENT 3 


Between the Ccelenterata 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 


4 Pie AOD Tp At 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 carnivorous 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 
Apodidz 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 
eut~for the seizing of -~prey: “Wé assume: that the 
Annelid from which Apus is derived, adopted a 
habit of browsing, which rendered this protrusible 


SECY.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 paleontologist abundant evidence that early 
Chzetopods 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 manille. 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 Chatopodan 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 Apodide,” 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- 


6 THE APODIDZ: PARE I 


stood, it is found to be very superficial. We shall be 
able to show, in the following pages, that the Apodidz 
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 bea 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 Apodide. 

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, 


srcren OBJECT AND LINE OF ARGUMENT a 


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 1s 
enough for our argument if we can show that such a 
deduction is posszble. It is not essential to our theory 
that we should show exactly ow 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 


5 THE APODIDZA PART I 


upon morphological, anatomical, and biological reason- 
ing, the Apodidz 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 paleontology 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 four 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 Trilobites. The mystery which surrounds 
these- primitive .Crustacea 1s 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.1. OBJECT AND LINE OF ARGUMENT 9 


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

After briefly discussing the Eurypteridz, 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 
Limulus, can be deduced from the Apodide. 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 Apodide 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 Apodidze are found, 
among the rich yield of Crustacean remains of the 
Silurian strata, admits of simple explanation. We say 
no ¢vue Apodide, for we shall find that such forms as 
Hymenocaris and Ceratiocaris, though perhaps some- 
what more specialised, were probably very closely 


ite) THE APODIDA: PART I 


related to the Apodide. Both these points will be 
discussed more in detail in Part IT. 

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: i! 


THE OUTER BODY FORM 


foe 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. 1 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 APODIDA PART I 


Owing to the bending on itself of the cylindrical 
Annelidan body, the original head must have been 
anteriorly almost completely hemispherical (Figs. 1 and 
46). This form of the original Crustacean-Annelid head 


6 


cd 
J 
\ 
\ 

\ 


Gers 


lic. 1.—Diagram showing the first six segments of a carnivorous Annelid ; the first 
five being bent round ; Pp, the prostomium with two pairs of eyes and a median 
cirrus. 1, the 1st segment carrying a pair of antenne, its under edge projecting 
backwards as the lower lip (2). 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 maxille, 
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 (rst free segment) with large dorsal parapodium carrying gill (g) and 
sensory cirrus (c). 


was, however,very clearly modified. In most Trilobites ~ 
traces of it arestill visible in the glabella (Fig. 47, p. 213). 
In Apus, the disguise is very complete, the whole head 


SECT. IT THEe-OUTER BODY. FORM 1g 


J 


being broad and flat. This form is due toa 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 believc that it was a 


Vv 
Coie =~C 
yr 
6 


Fic. 2.—Diagram of head and first trunk segment of Apus, for comparison with 
Fig. 1—the lettering the same. In addition: s, shell gland (the acicular gland 
of Fig. x 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,” z.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. 


14 THE APODIDA SPARE 

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 cesophagus, 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 backwardsas a sharp fold (see Fig. 1,7). 
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 maxilla 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.I1 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 of 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. 

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 Cf. pp. 39, 40, and 194, also Fig. 43, p. 188. 


16 THE APODIDAZ PART I 


ning round the front of the head, makes the latter 
broad and flat, obliterates all external marks of seg- 
mentation, and effectually ates 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. Inthe 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 larve 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 


SEGEs 11 fi OUTEROBODY- FORM 17 


~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 


Fic. 3.—Lepidurus Spitzbergensis, from nature. The left half of the shield removed 
to show the vermiform body. ‘The first r4 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, 4, brood pouch formed by the rith 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, ze. 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 APODIDAs PART I 


that it grows directly out of the Nauplius by the 
eradual 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 
midimentaty, sin -the Apodide is’ ta amatter sot 
considerable importance in estimating their true 
position. In all the other specialised Crustacean 
sroups the number of segments is, constant, (2 
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 Apodide. 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 OUPER 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 
Apodide varies between thirty-five and sixty-five. 
In the Entomostraca the number varies, but never 
reaches even the lowest number in the Apodide. 
That the number in the Apodide 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 Apodidze 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, z.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 Apodide, this 
segment is already fully developed; they are the 

C2 


20 THE APODIDA 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 
(f. pp. 85 and 274). The two cirri are stiffened for the 
greater part of their length by a thickened cuticle 
covered with sete, and showing slight rings of thinner 
skin. The tips of the cirri are quite thin-skinned, and 
seem to function as tactile papille. 

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


THE CUTICLE AND EXOSKELETON. 


The generally thin and flexible Annelidan cuticle of 
the Apodide 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 Apodide having this characteristic have been classed by Leach as a 
separate genus, Lefidurus. But Dr. Alfred Walter, to whose memory 
this essay is dedicated, discovered a form in a desert well in Trans- 
caspian Russia, A pus Heckelii (Walter), which makes it doubtful whether 


this division can be sustained. (Bulletin de la Société Impériale des 
Naturalistes de Moscow, 1887.) 


SECT. 11 fn Ob TER BODY FORM 2K 


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, z.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 maxilla, between which two pairs of limbs they 
are placed. 

(2) We find local thickenings 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 


29 THE APODIDA: 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. (0.) There are thick- 
ened strips to resist the longitudinal pulls of muscles, 
as along the shafts of the limbs, where, but for such 


Fic. 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 23 


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 


Sean =—=-e) 


Fic. 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 sameas in Fig. 6. On the endites are seen the denticulate 
setze 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 Mok gers RONBB Ise, 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, z.e. who threw them off altogether. 

But still more important consequences of the stiff 
cuticle are to be traced in the zzuer 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 CULER BODY FORM 25 


Apodide 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 Apodidz, thus visible 
in the form of the body (z.e. of both head and trunk), 
will be even more clearly seen when we come to 
consider the appendages. 


SEC ELON: (Ei 
THE APPENDAGES 


THE appendages of the Apodide 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 


SRC E III 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 maxilla, 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 APODIDZAE 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 ithe (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 
Apodide 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? 111 Ite 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 lone “mim,. however, the ‘typical 
Crustacean formula for mandibles and maxilla has, 
except in Limulus and in the Ostracods, which have 
different masticatory formule, 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. 

(1) 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- 
Gimee. 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 APODIDA PART I 


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 are 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 ofa 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 antennz (see Figs. 1 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 antenna, 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, 2e, to the use 
of the appendages as limbs whose functions were 
primitively all directed towards the middle line. 
The position of the antennz is especially interest- 
ing. In the Annelid, these antenne 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. II THE APPENDAGES 3i 


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 
Polychetan 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. (f~ 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 


32 TEE APF ODp TD As 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 4," is 
sufficient to enable us to homologise the shaft and its 


Fic. 6.—Diagram to compare an Annelid parapodium (A)1 witha limb of Apus (4). 
g, gill; cy, sensory cirrus of the dorsal parapodium 3 ca, ditto of the ventral para- 
podium ; v, ventral parapodium ; ¢, 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- 
wiirmer,” Leipzig, 1864-68. 


SECT. III THE APPENDAGES 33 


sensory cirrus of the dorsal parapodium of the 
Annelids. 

The sensory cirrus of the ventral parapodium (c,) 
entirely disappears in Apus, but zs retained tn Limulus 
(see Fig. 44; p. 192). Both the dorsal and ventral para- 
podia in Apus carry highly developed sete, as in the 
Polycheta, and again, in both, the gill is entirely free 
from sete, which would hinder the free exchange 
of the respiratory medium. It may be noted that 
many Polycheta have appendages on their parapodia 
quite as complicated as those on the limbs of Apus 
known as the endites (ZB. 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 Antenne (Fig. 7A)—This limb 
has retained its original position at the side of the 
Annelidan prostomium or upper lip of the Apodide. 
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 setze are modified into sensory 
hairs, homologous with the olfactory hairs of the 
higher Crustacea. 

Morphologically, the first antenna must be regarded 

D 


34 THE APODIDA 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. 1 will make it 


B 


Fic. 7.—A. First antenna (Z SAztzbergensis) with 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. £&. Second antenna (ZL. Spztsberxgensts) showing the rudiments 
of the endopodite, z.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. 1 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. 


SHCT. III THE APPENDAGES a5 


Il. The second pair of Antenne (Fig. 7 B)—The 
second antenna is so much reduced in the Apodide 
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. 1, 2). Its great reduction in the Apodidz 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. 188). 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 8) 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 

De 2 


35 THE APODIDA 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 maxilla 
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 /) our explanation 
of the parts of the second antenna. The correctness 
of this explanation naturally depends on a right 
understanding of the same limbinthe 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 gf Fic.:7 2, the biramose limb of the 
Nauplius consists of the dorsal parapodium, the distal 
portion of which forms the endopodite, the sensory 


SECT. 111 THE APPENDAGES 37 


cirrus forming the exopodite. As the larva grows, 
the endopodite 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). 

Ill. Phe Mandibles—These are the first limbs which 
admit of undoubted comparison with parapodia. We 
find, however, that while the dorsal branch has entirely 


Fic. 8.—A, diagram of mandible; s, shell-fold. At @ the dorsal parapodium has 
disappeared, but is indicated by the musculature. wv, ventral parapodium form- 
ing the mandible itself, the musculature coming from the sternal plate (sf). 4, 
diagram of ist maxilla. Atd, a larger rudiment of the dorsal parapodium is 
retained than in the mandible. 


disappeared, the ventral has grown cnormously in all 
directions to form the powerful masticatory limbs 
which are such a striking feature in the Apodide. 
That these mandibles are 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 APODIDA 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 
en  Musculature,-p. 52)... [he redevelopment. (im 
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 sce 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 sete are very 
sinall and -crouped in tufts round-the teeth, Tints 
limb has no hinge on which it works; it probably 
moves round its upper dorsal end as axis. 

The homologous limb in Limulus, z.2., the third, is 
not so specialised as it is in Apus; its masticatory 
process is not more developed than that of the second 
antenne, or of the two pairs of maxilla, 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. III THE APPENDAGES 39 


mandibles.t 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. 

The 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. 1, 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 maxilla. 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, 


oe TAFE APODUTEDA PART 1 


It is clear that this division of the under lip must have 
been: a svery. 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 
maxilla 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 (see 
Fig. 49, p. 220), which offers a curious parallel to the 
divided under lip of Apus and Limulus, but must 
naturally be due to other causes (see however note 
pre): 

IV. Zhe First Maxilla—Unlike the mandibles, 
the first maxilla has retained a small rudiment of 
the dorsal parapodium in the form of a fold. That 
{his is, im. fact, the reduced dorsal ‘branchis ‘elear 
from a comparison of the musculature with that of the 
other limbs (see Fig. 8 2). In the higher Crustacea, 
this limb also may redevelop its dorsal branch as 
maxillary feeler. The maxilla 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: 111 THE APPENDAGES AI 


in addition to the masticatory ridge or ventral para- 
podium. In the Eurypterida, the ventral masticatory 
part seems to have almost disappeared in Pterygotus, 
(see Fig. 155,.-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 
ise saquiecezed (Gee Figt 2 and Frontispiece), ~ Phe 
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 
Piece i, p12) The openine of the shell sland 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 
limit (seer Pigs. ba..p.'50;;and: 15,/p: 60). 

The five limbs just described are the typical five 


42 THE-APODIDA 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 mawillae, and less 
on the first than on the second maxilla. 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, z.c. the two pairs of antenne. 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- 
teridz, 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. III THE APPENDAGES 43 


rule, that the more the forehead is pressed against the 
eround, the antennz 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 antennze would tend to 
point forwards as sensory organs, and would then 
undergo further development as such. This we see in 
some Eurypteride, Branchipus, and Nebalia. 

The Apodide 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 antennz) 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 
Kurypteride, and are tabulated p. 250. 


44 THE APODIDZ PART I 


THE LIMBS OF ‘THE TRUNK, 


Passing from the second maxillez 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- 
surida, the Eurypteride, 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, z.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’ sepment,. 2.2. the first seament 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 (see Fig. 1). 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 mawillipedes, z.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- 


Sie MIT THE AP PEN DAGES 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 Eurypteride, 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, z.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 APODIDZ: PART I 


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 sete! on their flat surfaces, and 
tufts of sensory hairs along their edges. Besides 
helping in swimming, these sensory cndites 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 


see Fig. 5. 


SHCT, III 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, 


Fic. 9.—First trunk limb (Z. SAztzbergensis). Lettering as in former figures. 
é, sensory endites. 


and perhaps killed, by the ventral parapodia, which 
are provided with sharp thorn-like setz, 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 APODIDA PART I 


their muscles (see Fig. 12, p. 55) that what the 
enathobases 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 


tay 
a. UU nen,, 
‘ 
a, SO IGOre Uv 
lS ea (eee / 
& , 
7 
/ 
/ 
ti 
4 
——J <J 
~ 
js 
A \ 
Wa / / \ 
‘ 
wg vy VA , \ 
/ 4 
/ ¢ \ 
, / “ 
/ Z SS % 
/ 7 NS \ 
/ ; ~ \ 
‘ ~ ‘\ 
4 4 eens \ \ 
, 7 Se So 
4 ~ 
/ y = ay 
/ ¥ SSS SSS Se ees 
tf. / é 
/ rhe 
/ 
Cr 


Fic. 10.—The last rudimentary trunk limb (Z, Aroductus), 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. II Thi APPENDAGES re 


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 APODIDA: 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. [:conomy, 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. Til THE APPENDAGES Sa 


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. 


SECTION: IV. 
THE MUSCULATURE 


THE musculature of the Apodidz 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, ze., 
the bending of the head, that were there no other 
resemblance between the Apodidz 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 Apodidz 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 sesments.. Fig. 11 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 
gsrouping 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. 

ine the: sposterior end- ‘of the®. body »;wherey the 
parapodia are less developed, we might expect that 
the muscle bands would gradually spread out to 
form a more and more complete dermo-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 


54 THE APODTDA: 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 


S& 

6, 

a \ Reber 
QD CSS 

9 Wy ae 


df 


Fic. 11.—Transverse section through the trunk of a carnivorous Anneiid, diagram- 
matic (from Lang’s 7ext-book of Comparative Anatomy). g, gill; d, dorsal ; 
v, ventral parapodium ; cy, cirrus of the dorsal parapodium ; ¢e, ditto of the 
ventral parapodium 3 cz, circular musculature ; ¢7z, longitudinal musculature 5 
tim, transverse musculature; ac, aciculum; 7, nephridium ; ¢, 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, ze. those which 
diverge in the transverse plane. 

Let us now compare this sketch with the muscula- 
ture of the Apodide (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 


{ 1 ‘ 
i Og 
Wie KY 


Fic. 12.—Section through a specimen of Apus cancriformis, 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. @z, dorsal; vz, ventral muscle bands; s, shell ; 
e, eye 3 Z, under lip 3 #1, 79, rst and 2nd maxille ; v, ventral parapodia of trunk 
limbs ; 6 ditto of tst 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 cesophagus 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. 13. We see here not only the dorsal 
bands themselves lengthened to pass round the curve 


56 LHE APODIDZ 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. 


\ 
\ \ ‘ 
i Vi \ 6 
/ LE Gad} 
ma 2 


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


The ventral muscle bands of the Apodidz, 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, 
7.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 APODIDA: PART I 


much more weakly developed than the longitudinal. 
This ‘is also the case in the Apodidz, 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 dermo-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. 11, é7z). In 
the Apodidz, 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 


ae 


i 
ra wisn Bz5 - ! 
pene 
e 
: SoA 
n eo 


: 
as eee 
ones 


Fic. 14.—Transverse section through Apus cancriformis, to show the distribution of 
the musculature. /, heart; dr, dorsal; vz, ventral muscle bands 3 ¢7z, circu- 
lar musculature (as shown in Fig. 15, A); dv, dorso-ventral musculature (cf 
Fig. 11, 7772); #2, membrane enclosing the intestinal and genital sinus ; z, intes- 
tine; g, genital glands; ¢, eggs; a and 4 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 


60 THE APODID/® 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 


Cc 


Fic. 15.—Tangential sections through three segments passing between the long:- 
tudinal muscle bands and the lateral body wall, diagrammatic, the plane of 
the section passes through the points a and 6 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, ze, 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 


Y 
I 


_ 


Fic. 16.—Diagram to explain the courses of the muscle ees in B and C, Fig. 15. 
1 represents an Annelidan parapodium in its original horizontal position with 
three longitudinal muscle bands running into it. 2represents 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, 4) 
are probably elements of the circular muscle layer ; 
their position close under the lateral integument 
favours this derivation (Fig. 14, cm. ; of also Fig. 11). 
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 APODIDA: PART I 


of the septa (see Fig. 15, &), 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. 16. 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, C1 

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 Apodide is, with slight 
differences, the same as that found in the majority of 
modern Crustacea, viz., one pair of mandibles and 
two pairs of maxillz ; although in Apus, the second 
maxilla 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 


64 THE APODIDA 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, z.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, 4) 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, d, p. 37). 
The longitudinal muscles, attached to the integu- 
mental folds between the limbs (see Fig. 15, 4), 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 maxilla, 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 maxilla 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 maxilla to 


SECT. IV THe MUSeuULATLURE 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 maxillz 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 APODIDZ: PART I 


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: 
‘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.1 We find the brain in the 


1 See however p. 80. 


68 THE APODIDZ: 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 antenne diverge; they may perhaps have 
been united for some distance with the cesopha- 
geal commissures. It is even possible that their 


Fic. 17.—Diagram of the first five segments of a carnivorous Annelid to show the 
arrangement of the nervous system, from above. 4, brain; é9, anterior pair of 
eyes on the prostomium 3 ¢}, posterior ditto : aj, first antennz 3 a, 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 antennz moved forwards till they appear to 
be projecting from the posterior edge of the prostomium. 
The second antenne, 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 


° 
‘ 
1 
\ 


Fic. 18.—Anterior end of the same, bent as in Fig. 1 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 APODIDA 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 (@,) longitudinally, thus producing two 
cesophageal commissures, one (@,) in its original 
position, innervating the cesophagus and the upper lip, 
and the other (@,) carrying the brain and the eycs. 

This origin of the two cesophageal commissures in 
Apus is especially interesting because it explains ¢he 
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 


SECTS V. THE NERVOUS SYSTEM 71 


would be affected by this splitting of the cesophageal 
commissure owing to the travelling backwards! 
of the brain. Taking first the case illustrated in Fig. 
19, z.€., assuming that the nerves for the first antenne 
branched, in the original Annelid, from the first 
ventral ganglia, we tried to answer this question 
theoretically. Our answer, however, was not quite 
correct. Weassumed 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 antenne of Apus near the prostomium. These 
mistakes were very natural. For the second antennae, 
however, our answer was correct. Werightly 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 
antennz as branching off from the brain commissures, 
this position agreeing best with the position of the 
second antennz 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 antennz which 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. 


72 THE APODIDA: 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 


"1 


@37-~~ 


Hf 


Yen 
Uy cs C2 


ey 


Yj -_-"~ b 


Fic, 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 antenne of the original Annelid branched from the first ventral 
ganglion. 1, 2, 3, 4, 5, ganglia of the 1st, and, 3rd, 4th, and sth segments. 4, 
brain of Annelid; B, of Apus ; @), cesophageal commissures of Annelid ; @9, @3, 
the two cesophageal commissures of Apus derived from @  } a1, @g, nerves to the 
1st and 2nd antenne of the Annelid; aj, ag, ditto of Apus; ej, ég, nerves to the 
eyes of the Annelid; 7, ng, ditto to those of Apus. 


should be compared with Figs. 1 and 2, which show 
the positions of the antenne. 

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 


? 


a Le, | 


\----Lo 


- 
Aw 


ey 
Fic. 20.—Diagram showing the same as Fig. 19, drawn, however, on the assump- 
tion that the nerves for the first antennz in the original Annelid came from the 


’ posterior end of the brain. Lettering the same as in Fig. 19. shows the 
position of the ganglia of the first antennz 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 APODIDAL 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 antennz had already travelled up the 
cesophageal commissures to near the brain, then we 
have to suppose that these ganglia split away with 
the brain-cescphageal 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 antennz, and he supports 
this claim by the fact that the nerves branch back- 
wards, as shown in Figs. 20 and 21.!_ If this reason- 
ing is correct, then we may assume either (1) 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 hematoxylin 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-cesophageal ganglion. 


SECT. V THE NERVOUS SYSTEM 7 


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 antenne 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 antennz and the antenne 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. “proxiinity of the antenna: 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 antennz in Apus, and 
(0) the distance of the eyes from the antenne, and the 
difficulty of correlating their respective sensations, the 
eyes pointing forwards and upwards, the antenne 
backwards and downwards. 

In the higher Malacostraca, with well-developed 
antennz 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 APODIRA PART I 


Annelid, We may perhaps find some support for 
this view in the tact that +the frst vantenna~ 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 antennz 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 antenne. 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 
Pelsencer, because we considered this ring as no 
special nerve branching from the cord, but simply as 
the remains of the original cesophageal commissures. 


Fic. 21.—Central nervous system of Apus. Lettering as in Figs. 17, 18, and 19. 
ent, nerve to the eye muscles ; ¢2, the anterior pair of eyes transformed into the 
unpaired ‘‘ eye ” (cf § on the sensory organs); * the position of the ganglia of 
the 1st pair of antennz 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 APODIDA 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 
eanglia of the second antenne, would be, according to 
this view, the fused transverse commissures of the 
ganglia of both pairs of antenne, 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 
cesophagus, reaching as far back as the mid-gut. 
The brain lies between the hepatic-diverticula. 

In. discussing the appendages, we saw how the 
antennz, 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 antennz 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 doth the pairs of antennz 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, z.e, in an early Annelidan, not in a 
Crustacean stage. 

Of the rest of the nervous system little need be 


8o THE APODIDA 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 
Chzetopod 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 
ganelia of the, ventral cord -which} in’ the zbent 
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. 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 ncither 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 
antenne. 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 (@,); this shows 
that the ganglia must have had a hypodermal posi- 
tion, z.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 APODIDA: 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 
antenne along the cesophageal commissures, or, if 
these were already near the brain, their final union 
with the same, added further complications. 

Lastly, when the antenne, 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.¢., 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 st7// 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 


SHeT. Vv TEE 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. 


SECTION Va 


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 antenne, as sensory organs, admit without 
difficulty of deduction from the antennz 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. 1) 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 aré covered 
with setz 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 papillz 
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. 

(1) 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. 2u). 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 APODIDA 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 setz on the limbs may 
perhaps be classified as follows. 

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

(8) Long feathered hairs on the gnathobases, occur- 
ring together with sharp tooth-like setz, which latter 
help to give the gnathobases the character of jaws 
(Gee Fic. 0; p. 47). > The) merges of : these jhichty, 
developed tactile hairs are easy to follow; the 
ganglia at their roots are compound (see Fig. 31, 
p: E31). 

(y) 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. V1 THE SENSORY ORGANS 87 


(6) Hooked or bent hairs on the first antennze 
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 Pig. 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 
sete. If any of the original Annelidan sete 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 


88 THE APODIDZA: PARTA 


nerve swells into a ganglion. In the feathered hairs, 
a fibre runs into each barb, and the ganglion is a 
reoular group of: cells: “(see> Pig: mp) ie). 1 as 
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 
cesophagus 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, z.¢., 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 SY 


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 
Nereida) two pairs of eyes on the prostomium, which 
we have called simple eye-spots. Von Graber has 
shown that these Chatopodan 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, z¢., under a thickening 
exoskeleton. 

When we come to ask what are the most character- 


go THE APODIDZ PART I 


istic elements in the Crustacean eye, we find them to 
be“()./the cnystal.-cones, and *(2) the: ‘retinule, ae 
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 (z.e., visual) 
cells ; we have simply to account for the “ retinula- 
tion” of these cells, as Lankester calls the grouping 
of them into retinule. 

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, z.¢., 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 particulanty, 
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 gI 


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.’ 

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


—— rs 


SS 


> Se 


, 
! 


Fic. 22.—Section through the eye of Limulus (after Lankester). c, cuticle which 
grows into conical papille directed inwards, and pushing down the hypodermis 
cells (Ay); at the tips of the cones are found the retinule imbedded in connective 
tissue. 


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 retinula having been pushed down even 

1 Grenacher 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. 


g2 THE APODIDAL PART I 


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


CC 
poe 
‘ 
tf 
é/ 
/ 
a 
/ 
“ ° =~; 
; 7 yak 
i ale a S / 
uh 0 : ine Oe 
el --------> 
© a7 @ 
= Cc =e 7 f° 
ot 0O\ ASS, 
ie 2) BAS 00) (e 090 | 
ee Tie 5S . POO one 
C7 C Sa93 IQotoa Or, 
f CO OC GC 
SO) ) Lig? E , Le) Fee ex 
p - 0950: Yeas 


SS eee, 
CS Cae 
ona) 
a 


HO 
eG 
we 
4 | if 
at Wel 
— ———/ 
te Hie 


Fic. 23.—Diagram of two single eyes of the paired eyes of Apus. “Ay, original 
undifferentiated hypodermis cells secreting outer cuticular membrane. cc, 
crystal cone cells = differentiated hypodermis cells secreting the cones (cv); cd, 
layer of connective tissue fibres =subhypodermal connective tissue layer; 7”, 
retinule, z.e., groups of sensory (retinal) cells (7¢), with their rhabdoms (72), 
and pigment cells (#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 toseek. The rounding off of the distal ends 


1 See Appendix II. on the pigment in these cells. 


oe 


Go 


SECT. VI THE SENSORY ORGANS 9 


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 in 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 APODIDA. PART 


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 eyée* “Our “object here has been te 


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: 

1. 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 APODTDZs 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, z., about ds 
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 “s7the 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 jeyes of 2ipus: 
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 


Fic. 24.—Diagram to show the water-sacs over the eyes of Apus. s, water-sacs; 
c, canal leading into the same; , pore; é, paired eye ; é, brain; @, cesophagal 
commissure; og, optic ganglion; wz, eye-muscles. The eye is drawn in the 
section ; in reality a median section passing through the pore passes between the 
eyes ; é9, 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 APODTDYE: PART I 


above the unpaired “eye ” (sce Fig. 27, p. TO5).. 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 sims. If 1s not amprobabie® 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, z.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 with 
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 Tt 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: 
(1) 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. 


2 


t;, 


Tao THE APODIDA PART I 


eradual diminution of the shield and head, and the 
sinking anf: the sidés' lof the latter’ till -the eyes: 
with their cones of long muscle-bands (cf Fig. 24, 72), 
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 carryme the 
enormous second antenne and the stalked eyes. 


THE UNPAIRED “BYE.” 


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 
FuNS: into it is really*open at the end) tis 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 IOI 


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. 
heis, 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 (ze. 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 (1) 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 (see Fig. 25); posteriorly, it ends in a blunt 


102 THE APODIDA 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 retine. The 


Z 2 o 


Fic. 25.—Longitudinal median section through the unpaired ‘‘eye,” diagrammatic. 
vr, ventral ; dv, dorsal retina; , tangle of pigment cells; s, water-space over 
the eyes; c, canal of the same, giving off a branch (cj) into the interior of the 
organ; 8, brain. 


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 retinz 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. 


S 


Fic. 26.—Lateral view of the unpaired ‘‘eye,”’ diagrammatic. v7, ventral ; dr, dorsal ; 
Zr, lateral retina 5 é, brain; S, water-space; ¢, canal of the same; Z, 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 retine, 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 APODIDA: PART I 


pscudopodia without any apparcnt arrangement. 
They contain the same minute olive green pigment 
granules as the other pigment cells, which spread out 
in great numbers over the subhypodermal conncctive 
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 scen from the drawings that the retinal 
cells are not all of the same size. The lateral retine 
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 


Lettering the 


SS) 


, diagrammatic. 


nerve from the anterior part of the lateral retina (cf Fig. 26). 


Fig. 2 


in 


a 
2) 


CY / 


OY OPP ~ 
con 


Ss 
\ 
\ 
4 


Bh 
BAY 
{Mo 
<A 


sy 
EAN “NS INS EN 

me Yy 
“ \ or eet 


Os ( 
S QR 


‘A A Lieiges 
OK ) 4 N\ 
AX NY ty 


npaired “‘eye,’’ along the lines 1, 2, 


Fi 

ry Si 
s yg & 

s 
i S 
— oS 
. ~ 8 e 
\, Bs 
NC FB4) “NA QQ ae 

© L eM ON) yi) \ . NN a 
ae RSA h \ i SS o | 
sie EEN UO INES WW ae 
DR z 

: SAs7y RY ) oS ee 
SRS WY: §4 

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io) 


106 THE APODIDA: 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 
retine. 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 Apodidze 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 


Ra 


SS VE : 
\ - £6 VE 
. NN a, Pp 
. 4 
NS “ir 


Fic. 28.—Diagram to show the origin of the unpaired eye out of two eye-spots, the 
retin being drawn down to form the lateral retin of that organ. /, pigment 
cells. The pigment in theeyes 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 retina 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 


108 THE APODIDZA: PART I 


crowding between the retine. 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 cycs. Further investigation must decide 
whether the dorsal and ventral retina and the larger 
posterior cell groups of the lateral retina are later 
differentiations of the same two original eyes or new 
developments. In the inner ends of the retinal cells, 
Z.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 hypodermis, 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 retine 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 appéar-according te 
Packard, ox the 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 cye, 
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 retine 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 ofens 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, ze. of 
“epi: The end of the tube.as. shown. in: Fise 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. Butsuchan appreciation of changes of pressure 


IIO THE APODIDZ: PART I 


could also no doubt be equally well effected if the 
canal were open, the increase of pressure leading 
to arush 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 havea 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 Apodide 
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 LHe SENSORY ORGANS III 


with the rowing limbs, and long before the paired eyes 
are developed or needed. Itis 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.’ 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 
larve, 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 Grenacher, and that of Apus will be referred to in § XV. 


SECTION. Vil 
THE ALIMENTARY CANAL 


ALTHOUGH it is not, as a rule, possible to draw 
any conclusions as to the relationship between animal 
eroups from the similarity of their alimentary canals, 
yet the likeness between the digestive tract of the 
Apodide 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 cesophagus, 
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. 

ihe 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 
cesophagus would slope upwards and forwards. We 
find that it has this position in the Apodida, and 
from the Apodidez it has been handed on to the 


SECT. VII THE ALIMENTARY CANAL 13 


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 Anus 
to Limulus and to the Trilobites. 

The cesophagus 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 cesophagus 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 cesophagus is lined by a chitinous 
intima and provided with setz which project upwards 
so as to form a fish-trap apparatus. The paired glands 
which open on the under lip close to the mouth (sec 
Fig. 29) will be described in the section (IX.) on the 
excretory and other glands. 

The cesophagus 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 


114 THE APODIDZEZ 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 cesophagus projects somewhat into the mid- 


Fic. 29.—Diagram of the branched diverticula of the anterior end of the mid-gut (wz), 
on the left without the glandular (hepatic) branched ceeca, on the right a few of 
the latter are drawn. @, entrance to the cesophagus, 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. 


cut, 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 occu rin 
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, - Ity 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 elands. 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 


EZ 


116 THE APODIDA PART I 


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!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 cesophagus, 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. 


SEGMION VITt 
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 


118 THE APODIDAs PART I 


the development of the exoskeleton, 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 lacune, 
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 truc 
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 Anneclidan vessels ; 
these are the short vessels which supply the shell 
olands. (See. Fig. 30, p. 125.) They branch Jom 
from the dorsal vessel on each side, and descend 
towards the dorsal parapodia of the second pair 
of *maxillz, 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 moreand 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 Apodidze 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 descriked, it streams out 
ventrally through fenestrae in the membrane which 
forms the sinus.. (Gee Fig. 14,772, p. 59.) It,is thus 
enabled to stream over the ventral cord, and then 
outwards on each side along the ventral side of cach 
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.’ 

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 APODIDA: PARTI 


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 
(where 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 Pigs 14, p. 50; "77 also Fis.-06; sp. 207) 
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 dorso-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, asa 


SECT. VIII THE*CIRCULATORY SYSTEM I2I 


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 ccelom 
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 
bandss (See Pigs: 06,'p: 207,,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 


E33 THE APODIDZ: PART I 


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 degencration 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 hamolymph between 
the integument and the ccelom epithelium might 
very easily lead to the loosening of the latter from 
the integumental musculature, excepting along 


SECT. VIII THe GriRevULATORY SYSEEM 123 


the intersegmental folds, across which the dermal 
blood-vessels of the original Annelid did not run (?). 
These two modifications, (1) 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 Apodidz does not therefore 
stand in the way of the acceptance of our theory of 
their close relationship. Apart from the well-known 
fact that) amone,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 IIT.) 


SECTION LX 
EXCRETORY AND OTHER GLANDS 


The Shell gland—THE most conspicuous glands 
of the Apodidz, 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. 1 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 


ui 
1 


STAT 
erate ‘ 
ee 
co 

Ga 

ass 

soe 

Gav's 

rete 


eae 


AEN 


NIN phi SONS 
ROS TARR, 
URIS 


RNY \ 
CESS ATWO LNT 
SAID oa Sea FAENUONS 


Ve. 

Se rR 

ie ee rt A 
OEE WY 


Fic. 30.—Diagram of the shell gland. 4%, the heart 3 ac, the aorta cephalica. The 
blood is distributed to the gland through a special vessel on each side. Zs, 
terminal saccule}; #c, urinary canal; 4, chitin-lined bladder in the shaft of the 
dorsal parapodium of the 2nd maxilla (7), 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 APODIDA PART I 


served. The mandibles and first maxilla have lost 
their dorsal parapodia almost entirely, and with them 
all traces of setiparous glands. The antenne, 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 
ef. the esophagus. (Fis; 29, p.1i4), ~ We thints 
that these belons almost ‘certainly to the first 
antenne ;! 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 antenne, 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 maxillz 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 antennz, 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 
antennee which had redeveloped their acicula as teeth. 

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


eri 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 
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 
antenne ; 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 1 and 2, we think the last view is the most probable, 
although we do not reject the second alternative. 


128 THE APODID 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, 
11 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 antennz (in Apus at least), the antennal 
gland, and the shell gland of the second maxille. 

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

(1) 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.* 
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. ‘We 
should expect the shell gland in Apus to be a 
transition form between the Annclidan 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 Annclida, 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. 


Kk 


130 THE APODIDAI PART I 


panereas ofthe higher Crustacea” But this: as 
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. 

Setiparous glands on the trunk segments are hardly 
to be expected ; the dorsal parapodia are developed 
into complicated limbs covered with sete, and the 
ventral parapodia are also thickly beset with sete of 
different kinds. We have succeeded, however, in 
finding two such glands on the same limb in one 
specimen of Apus cancriformis (see Vig. 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 are 


SECT. 1x EXCRETORY AND OTHER GLANDS Ist 


inclined to consider them as occasional abnormal 
returns to the Annelidan method of developing the 
sete (sce 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. 


Fic. 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. 2, 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 sete, establishes beyond 
all contradiction the usual homology of the leg or 
coxal glands of the Crustacea with the setiparous 
glands of the Annelida. 


132 THE APODIDA: 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 cstablish 
the close relationship which we maintain exists 
between Apus and the Annelida, to discuss the 
typical excretory organs “of the latter (72, °etre 
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 Vatural 
FHlistory of Creation,’ characterises the Crustacea as 
segmented animals without nephridia, and the worms 
as segmented animals with segmental 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 cither 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 homologise 
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 Annelidan 

L Ed. vil: 1801,)p-" 576: 


Stem ix EXCKE TORY 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 postcrior 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 antcrior 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 Anncelids the nephridia are often the ducts for 
the transmission of the sexual products, we naturally 
Eied 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 PART I 


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 zzzo 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 
ere. 

All the segmental genital tubes on each side are 
connected together bya 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 


Shei. EXOCRETORY 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, 
mpem invthe body ‘furdi(cee: Pie 11, 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.) Isnot 
this epithelium homologous with the Annclidan 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: cofonmr 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 orzg7nal cwelom epithelium 
which covered the 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 


136 THE APODIDA 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 nephridial apertures except the 
one between the tenth and elevetith 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 
elands themselves, as one would naturally have ex- 
pected. This epithelium, then, isa 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. 11, p. 54). 


sue Ix EXCRE TORY 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, z., 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. 

We 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 APODIDZ PARP 


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 ne- 
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 either to remove all difficulties so long as 
they are not positive contradictions, or to state exactly 
how such and such a transformation came about, but 
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 Annclidan 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 EXCRETGRY 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. 1. It appears to be an island of dermal 
elandular cells, the last remains of the Annelidan 
dermal glands, which the thickening exoskeleton 
probably rendered impracticable. This organ will 
be desenbed im detan in Appendix IV. Tt eters 
indirect support to our theory of the Annelidan origin 
of Apus. 


SEGTION “X 
REPRODUCTION 


THE carnivorous Annelids are mostly sexually 
separate. The same was originaily the case with 
the Apodide, which are 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. II, p. 
54), and then, falling off, ripen in the body fluid. 
They are 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 
Apodide. 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 saidabove. The genital glands 


SECT. X REPRODUCTION I4I 


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 im Es slacialis 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. 


Fic. 32.—Diagram of a somewhat simplified genital tube of Z. Spztzsbergensis or 
gtacialis, the dorso-ventral segmental tubes only slightly branched. a, aperture 
between the roth and 11th segments; ¢, the rudimentary part, ze. 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 APODIDZ PART I 


any other Crustacean, and is in itself evidence of 
the primitive or Annelidan character of Apus. The 
eradual 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 
coelom 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 Polychztan 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 


SHOT. X REPRODWU CTION 143 


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 
eranules. The former is the nucleus of the future 
eae, the “latter aré 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. 33,7). 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 APODIDA: 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 
ese M5 fipe, these nuclei ‘come to the Suriaceand 
eradually disappear. 


Fic. 33.—Eggs at different stages. In stage 1 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. 7, testes as 
occasionally found (e.g. in A. cancriformis); 2, 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 roth and 11th 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, z.c. 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). 
While 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 Apodidz 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. 33,2). In some species ! 
the whole of the epithelium at the extreme posterior 
end of the genital tube breaks up into sperm cells. 

The sperm cells of the Polychzta 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 LZ. glactalis and L. Spitzbergensis, see Fig. 32. 


146 THE APODIDE: PART 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 cax 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- 
celland three nutritive cells. As the nutritive cells are 
probably modified egg-cells, the eggs of the original 
Crustacean-Annelid may have developed out of its 


SECT. X REPRODUCTION 147 


ccelom 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, 7), but in this case the organs which 
this wall originally covered, z.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. 


SEC DI@ine C1 
DEVELOPMENT 


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. 
ihe -seneral: likeness -of “the “adult sApus 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- 
tonine (see -Faes.. 134,30, .and qi). ~~ Iaus “the 
Nauplius larva of other Crustaceans is simply the 
Apus-stage in their development. We repcat 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, z.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 APODIDA PART I 


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 a7 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. 
Ws *to the case)/in’ which! the stage (is. passed 
through in the egg (eg. among the Malacostraca) 
it is, important to mote that) this. is mot the ‘ease 
in alZ Malacostraca, a Nauplius stage occurring, 
for example, in the development of Penzus 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 T51 


beyond the Trochophora stage. A comparison of 
the Nauplius of Apus just hatched from the egg 
with the figure of a Polychzetan 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, ze. the homo- 
logues of the parapodia of the first #4vee Annelidan 
segments, traces of the dorsal fold which belongs 
to the f7ti segment, and further slight indications 
of ve trink segments (see Fig. 34), in all ex 
Annelidan segments. A larva so far developed 
cannot be compared with the simple ciliated Tro- 
chophora, which when hatched probably represents 
only ¢wo, z.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 suchs a bent, intestine (Gee Fic. 37)... We. will 
take these points in turn. 

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


152 THE APODIDZ 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 frce-swimming larval life. Asa 
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 


fil 


Fic. 34.—Nauplius of Asus 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, z.c. 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, ze. unless the slight bulging, 


which. 1sseen..at) its base: in dA pus (see Pig. 7a 


p..34):.eepresents. the: wemains./ of such, a, dorcal 
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. 

The second limb is biramose, z.e. besides the sensory 
cirrus, the parapodium on which it stands is also re- 


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


tained. Repeating the homologies 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 
likea small branch. Here clearly the exopodite is the 
sensory cirrus, the endopodite the tip of the dorsal 


154 THE APODIDA: PART I 


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 Nauplius of L, 
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 asensory organ, one of its branches, that represent- 
ing the tip of the original dorsal parapodium, dege- 
nerates, leaving the other, the sensory cirrus, z.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 homologue of the ventral para- 
podium, we may perhaps sce 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, 


Shei Xi ; DEVELOPMENT 155 


as we shall seé in Part I]. 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 Ist and 2nd 
maxilla, 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 (z.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 APODIDZ: é PART I 


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 witli the abdominal 
limbs of the macrurous Decapoda. 

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 assumcd to be. 

2. The Unpatred “ Eye.’—The presence of powerful 
rowing limbs in the larva necessitated some more per- 
fect sensory organ than any possessed by the Annc- 
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 


SECE XI DEVELOPMENT 157 


the original Annelidan prostomium has been discussed. 
We 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 cyes paired and unpaired 
far more anteriorly placed in the Nauplius than they 
are later in the adult, ze. mzdiway between the ventral 
position in the bent Crustacean-Annelid and the 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. 108 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 morc highly developed larve 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, 
Zoza larve. 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 asa 
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#@ PART I 


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 
flabeila are far more capable of appreciating and 
responding to sound waves than is a plexus of pig- 


\ ~ 


\\ 
\ \ 


\\ 


\ 
\ 

\ 

\ 
\\ 


\ 
\ 
\ 
\ 
\ 
\ 


Fic. 36.—Nauplius of ZL. productus from the side (after Brauer), showing the position 
of the eyes at the frontal end, 7.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 


Fic. 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. 4, 
large upper lip; s, dorsal shield; @, dorsal parapodium of the second antenne ; 
¢, 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 


160 THE APODIDZA 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 
Gee Fig: 30,°/) on) each ‘side of the unpaired seve: 
They disappear throughout nearly the whole class 
in the course of development. They may perhaps 


Fic. 38.—Nauplius of A. cancriformis just hatched, dorsal view (after Claus). 
S, posterior edge of the shield ; JV, the large larval excretory organ, the dorsal 
or neck organ. 


be supposed to represent a pair of feelers rising on 
the prostomium of the original Annelids, suchas those 
found, for example, on the prostomium of the Eunicide. 
In asection 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. 


SHOT. XI DEVELOPMENT 161 


3. The Dorsal Shield.—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 Ostracoda!) can it be said with truth 
that the racial form possessed a dorsal shield, at least 
asa fokdeot the hith 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 im 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 Estheridz, which have a large dorsal 
shield, are more primitive than the Apodide 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 larvee 
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 be 
discussed in a special section of Part II. 


M 


162 THE APODIDA 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. 215, 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, z.e., of the two pairs of maxille. 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, 
iideed, if teaches ansenormous “size (see Pia) 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 Annclid, 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. X1 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 
Nauplius 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 
antenne 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 antenne branch 
off from the cesophageal commissures, there can be 
no doubt that the ganglia are infra-cesophageal (cf 
Section Vi.): 

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

M 2 


164 THE APODIDA: 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 antenne 
and the second pair of rowing antenne, 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 appcar 
more iin their delinitive form, “The two “pairs” on 
maxilla have no function to perform in the larva and 
are only developed later. Jt 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 cye, 


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 ( Figs. 34, :30, 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 
erowth 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 Apodide, z.e., the great number 
and peculiar arrangement of the limbs, is in reality 
one of the strongest proofs of the undifferentiated 


166 THE APODIDZ PART I 


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 


Fic. 39.—Second larval stage of Apus cancriformis (Claus), showing the gradual 
development of Apus out of the Nauplius, the liver as diverticula of 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 cither absolutely or very nearly con- 
stant. The type is fixed. In the Apodide, 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 
Apodide, 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 Anneclids we have a large 
and variable number of segments, in the higher 


Sees 
I fe = or 


EE 
ie 


ma! 


Lip 
nae, 


Fic. 40.—Second larval stage of Lepidurus productus (after Brauer). 


Crustacea a comparatively small, and for each group 
a fixed, number of segments. Between these two the 
Apodide 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 


168 THE APODIDA PARTI 


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


Fic. 41.—Fourth larval stage of Apus (Claus), the diverticula of the mid-gut com- 
mencing to form the glandular invaginations (2). 

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 


SECTAXI DEVELOPMENT 169 


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

We have had to limit our remarks to the larval 
history of Apus, as observations on its embryological 
development are unfortunately wanting. We may, 
however, here mention one or two facts in the embry- 
ology of the Crustacea which bear upon our theory. It 
is stated,! 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.? The significance of 
these two facts saken 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 ccenogenetic. 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. 
We find then this acquired rearrangement of the meso- 
derm shifted back to the earliest embryonic stages. 

1 Grobben, ‘‘ Die Entwickelungsgeschichte der Moina rectirostris. ” 


2 Korschelt und Heider, ‘‘ Lehrbuch der vergleichenden Entwicke- 
lungsgeschichte der Wirbellosen Thiere.” 


170 THE APODIDA PART I 


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


In pre-Cambrian times, of which there are now 
no fossil remains, a browsing carnivorous Annelid ac- 
quired the habit of keeping its “head,” z¢., 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 antennz, antennal para- 
podia, and parapodia developed gradually into Crus- 
tacean antenne, mandibles, maxilla, 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. 


ae ee 


4 Ww 


an 
Ly 


cee bas a 
id : Oe ig #3 i y ; 
M i ie 


PARLE IL 
Sb C7 ONS 211 


RELATION OF APUS TO THE OTHER CRUSTACEA 


IN 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 Apodida 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 sce 
whether they, like Apus, are capable of being deduced 


174 THE APODIDA: PART II 


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 Annclids into Crus- 
tacea did not result in only one form of primitive 
Crustacean, butin 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 Trilobites, 
that is, if we may judge from the extraordinary num- 
bers and varied development of these carly Crus- 
tacea in palezozoic 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 Eurypteride and Xiphosurida, to 
which latter the still living king-crab belongs. 

All these groups, however well adapted to their 
palzozoic 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 Apodida 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 


sreieown RELATIONSON APS £O CRUSTACEA 175 


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

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 
iozm Of a Special eroup, is in réality 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 Eurypteride, 
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 go 
into many details, especially in our comparison of the 
Apodide with the many living Crustacean forms. It 
will, we think, be granted, that a successful grouping 
of the Apodidz with the Xiphosurida, the Trilobites, 
and other early forms as common derivatives from a 


176 THE APODIDA: PART II 


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 Apodidz, 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 forthe 
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 agrce- 


Shere ait. RELATION OF APUS TO CRUSTACEA 177 


ment 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;! and the able attempt of Lan- 
kester 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. (1) 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) 
Whcereas the Xiphosura bear markedly the character 
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 
paleontology, 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 Nauplius stage in the embryological development 
of Limulus has met with no favour. 

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


1 **Tes Limules ne sont pas des Crustacés—ils n’ont rien de commun 
avec les Phyllopodes.” Journal de Zoologie, par P, Gervais, vol. 1. p. 42. 
Paris,: 1872. 

N 


178 THE APODIDZ PART Il 


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, z¢., as a sur- 
vival from early geological periods, its likeness to 
Limulus takes on at once a new meaning. ‘The diff- 
culties above mentioned deserve to be re-examined ; 
fortunately they are not insuperable. Before going 
into a detailed comparison of Limulus, let us briefly 
indicate the way these three difficulties may be met. 

I. Inthe 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 


sEecT. X11 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 
pus: - 

Further, while Apus has, as we have scen, retained 
the more primitive form of limb, not far removed from 
the Annclidan 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 


NO 


180 THE APODIDA: PART II 


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- 
t10OnS: = 

(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 Seldon? presefved;/in va. fossil, ‘state, Ib was 
probably first shut off from the ocean in brackish 
lagoons, and was gradually driven by the struggle for 


SECYr.-xiIr 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, whatiare 
the probabilities of its leaving any fossil remains ? 
The marine carnivorous Annelids of palaozoic 
times have left only their hard, chitinous teeth, 
so that the Apodidz 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 Apodidzx 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 Apodidx 
recorded from Paleozoic strata, yet there are abun- 


182 THE APODIDA PART II 


dant remains of Phyllopods, many of which show 
such a striking resemblance to the Apodide 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 deficicncy 
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 oar 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 dent Annelid. 


Fic. 42.—Section of Limulus rotundicauda 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. 4, brain; /, heart; 2, openings of the hepatic ducts in the mid-gut : sf, 
the sternal plate. From Bronn’s A7vassen und Ordnungen des Thierreiches. 


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 zs found in Limulus, in 


184 THE: APODIDAZ PART II 


essentially the same place as in Apus, and is known 
as the sternal plate. If we have piven: the) risks 
explanation of these two pcints, 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 dent 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- 
patison. with..that. of Agus, the. vefy “difierences 
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 homologise 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 are many 
points of resemblance in external organisation which 
should be mentioned. It will also be useful to 


SECIT XI 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.! 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 Limuius 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, of: 
the sections of a Trilobite Fig. 54, p.230. 


186 THE APODIDZ PART II 


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 Xiphosuride, 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 
eroup. Apus developed, comparatively speaking, a 
large number (50-60), Limulus a small numbcr 
(Ga, 16). | 

The metamerism : of imulus: as probably-to “be 
reckoned as follows : 

Cephalothorax.—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. 


Src xr RELATION OF APUS TO.CRUSTACEA 187 


Abdomen and Caudal Spine—These probably re- 
present nine! 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. z.¢., of those Apodide in which the anal 
segment is produced posteriorly into a flat plate. 

According to its erternal 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 
foot, so as to allow the? limbs, wc. 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. 


158 THE APODIDA: PART II 


THE LIMBS. 


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


Fic. 43.—Ventral surface of Limulus moluccanus 9 (after Van der Hoeven), 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 first 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 


Snort RELATION-OF APUS TO CRUSTACEA «180 


_ of the Phyllopoda 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 
Partal. 

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 antenne. In general 
form these limbs do not differ much in the two animals, 
as may be seen by comparing Fig. 43 and Frontis- 
piece: hes chief difference) isithat in; Limulus; the 
bends have developed hinges, owing to the greater 
development of the exoskeleton, and there are chelz 
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 
Araneidz), asa simple feeler ; in another (the Scorpio- 
nidz), it develops powerful chelz; 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 


i tere) THE APODIDA PART I] 


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 
antenna of A‘pus: “In the first place “the “position 
aprees very well (ci Fig. 43 with Frontispiece), 
This was not, however, the real reason for our change 
of opinion, which was duc to a comparison of the sixth 
limb in Apus with the sixth hmb in Limulus, and, to 
anticipate somewhat, with the sixth limb in the Eury- 
pteridz 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 was the 
first free segment, and its parapodium was thus free 
to develop into a limb for locomotion or for some 
other function. /Phus, Haking the ‘sixth lmb yer 
Limulus to represent the first trunk limb, the full 
number (five) of head limbs were left to be homologised 
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, 


SHOR KREATION OF PUSS; TO CRUSTACEA -i91 


also develop powerful seizing hooks. The homology 
of the second pair of limbs of Apus with the second 
antennee of the Apodide 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 samc 
form as the third, and are homologous with the two 
pairs of maxille 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 ecxopodite) 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, the 


192 THE APSDIDAL PART II 


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 homologised 
with the first trunk limbs of Apus and the other Crus- 


Fic. 44.—Basal part of one of the 2nd pairs of limbs of Lzmulus moluccanus (from 
Bronn’s Klassen und Ordnungen des Thierreiches) to show the well-developed 
ventral parapodium, retaining the remains of its sensory cirrus (c); @, 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 feclers 


°9 


Speman REE ATION Or APs. LO 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 
chelz. This whole limb seems to combine loco- 
motory with protective functions. Its tip les 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, 
7.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 legis abrupt ; there are no transition forms 
asin 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, ze. 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, ze. the last four head limbs and the first 

O 


194 THE APODIDZ: PART II 


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, 
z.c. 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. Sit 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, 
Z.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 (ze. 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 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 integumental folds, the dorsal (z.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 


a ih) 
[J ig 


i Y. 
‘ 


‘ AN 
We 
PNA 
OVE G 


tye \ 


eee wi 
y Sg Eero ey ee 
SSS 
an 
S 


tet, 


iil 


Fic. 45.—One of the abdominal limbs of Zzyz2dis for comparison with a Phyllopodan 
limb. w, ventral parapodia fused together; d@, dorsal parapodia (endopodites) ; 


ff, 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, X11 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 deducible, 
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 APODIDZ PART Il 


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

The 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 im’ 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. Ju 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 (shown 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 Ray Lankester pointed out, the brains of 
Apus and Limulus are alike in constitution, both 


SECE. XU RELATION‘ OR 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 antenne ; in Apus, however, according to Pelse- 
neer, the ganglia for the first antennz 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. We 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 
EMe hemes tomtie, Unpaired. “ eye; “Or 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 retinze with the 
ocelli, the nerves from the postero-dorsal and ventral 


200 THE APODIDA PART II 


retina 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 
meting: from: those “of the. lateral. retinee:Seemy fo 
indicate that they must have been derived from some 
other sensory crgans. 

Owing to the backward prolongation of the mouth 
and the cesophagus, and the arrangement of the 
limbs round the former, the anterior nervous system 
is. very concentrated ; the nerves forthe anterior 
antennz and the five pairs of limbs branch out 
radially from the thickened cesophageal commissures. 
It is as if the cesophagus 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 cesophagus. 

Between the nerves to the fifth pair of limbs and 
those to the operculum is a pair of nerves to the 
chilaria or cunder lip, If ithe ‘homology of “he 
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. 


Seelam RELATION VOR 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 occupicd 
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 thickenine 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 APODIDZ: PART TH 


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 cn 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, z.e. from between the optic nerves. 

Judging from the lateral retinee 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 retinula. 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- 
nule, 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 formed 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 retinula, being» no 
longer grouped round crystal cones, are, as reti- 
nule, comparatively uscless. We turned, therefore, 
with great interest to Lankester and Bourne’s 
account of these retinule, and found what we 
expected, that they are by no means so definite as 


204 THE APODIDA PART Il 


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 are 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 are 
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 cesophagus of Apus. Its 
anterior portion is widened out to form the so-called 
pro-ventriculus, 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 p@sterior send of this ‘projects sliexa 
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. x11 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 diverticula 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 Claus’ 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 


206 THE APODIDA& PART II 


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 Annelidan 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. 

The 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 Apusis what we should 
expect from the compression of the whole body, and 
the consequent diminution of the body cavity. 


Siemon REEATION OF APUS FO 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, ze. 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. 


208 THE APODIDZ: PART II 


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, z.e. 
either dorsal folds like the Apodide, 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. 


SEGCLION XI 
PAE ERI ORL PEs 


IN 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 Apodide with the Trilobites and the 
Eurypteride. 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 Xiphosuride, the Trilobites, and the 
Apodidz for the first time in a natural system. 

It is important to bear in mind that the Trilobites 

EP 


210 THE APODIDA& PART II 


are the earliest known Crustacean forms. A special 
interest therefore attaches to our endeavour to prove 
that they were nearly related to the Apodide. 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 Apodide 
as related forms, it Seems to US that eur scase ais 
established. Such concurrent testimony from deve- 
lopmental history and from paleontology is almost 
without parallel. 

The relationship of the Apodidez and the Trilobites 
has already been assumed by the earlier zoologists. 
Burmeister,! 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 paleozoic seas. Although 
Burmeister’s reconstruction was not correct, yet his 
assumption of a relationship between the two was 
justified. The fact that the Apodide have rowing 
limbs does not in any way oblige us to assume that 
if the Trilobites were-related to the Apodidz they 
must have had similar limbs. As a matter of fact we 


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


SHCTe Mitt TEE TRtLOBITES 201 


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 Annelidan 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. 

(I1.) 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 tetained: ‘only in the Trilobites. In.Apus it is 
completely disguised by the growing together of the 


ee 


— 


212 THE APODID£ PART II 


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 Fis a7iand 57, p. 257). The: orem 
of this ridge is probably to be sought in the folds 


Fic. 46.—Longitudinal section through Ceraurus pleurexanthemus (after Walcott), 
showing the intestinal canal and ventral membrane, and the bend in the head. 
Cf, Figs. x 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 antenne both in Apus and in Limulus, where 


SHeD. SIT PHE ERILOBITES 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 


Fic. 47.—Dionide formosa (Barr), showing the glabella 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 body, all the shapes of the Trilobite 


Zid THE} APODIDZA: PART II 


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. Asa 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. XIII 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 
Fig. 57, p. 257. When this crescent is large, owing 


Fic. 48.—Acidaspis Dufrenoyi (Barr), Upper Silurian (after Barrande, from Zittel’s 
Handbuch). 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 APODIDAZ PART II 


two origins are, 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-shicld 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 we 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 
inthe 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, 


NI 


SECT. XII GEE LREEOBITES FeAl 


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

(II) The 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, ze. a continuation of the gla- 
bella alone, as was no doubt originally the case. The 
eradual 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 APODIDA PART II 


The variation in the number of trunk segments is 
also a point of no small interest. While some authors 
have tried to classify the Trilobites 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 Apodidz, where the 
number of segments varies greatly: from 60-65 in 
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 Apodide 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 Apodidz 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. XIII iE TRIEOBITES 219 


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

We 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 /used 
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 APODiTD A: PART II 


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


Fic. 49.—Asaphos megistos Hall (after Walcott), showing the well-developed 
ventral parapodia, and the gradual simplification of the limbs from before back- 


wards, asin Apus. Cf Frontispiece ; 4, pygidium. 


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

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. XT ELE TE RILOBELFES 


iS) 
S) 
ial 


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. 


The Trilobite Limbs.—I\n spite of the great progress 
which has been made in our knowledge of the limbs of 


N 
N 
tN 


THE APODIDA PART II 


the Trilobites, chiefly through the patient researches of 
Walcott, they are still shrouded ina 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 (ze. of antennz) in front 
of the mouth.. This, however, from our point of view 
is no real difficulty. In reality the antenne 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 antennez 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 homologise these large locomotory limbs with the 
sixth pair of typical Crustacean limbs, ze. 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 maxilla of the typical 
Crustacean head. In front. of these and behind the 
labrum, we have, in Walcott’s restoration (Fig. 50), a 


SECT. XIII THE TRILOBITES 223 


space in which we think the antennz 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 antenne as here de- 


Fic. 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 antennz 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. 

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


224 THE APODIDZ: PART II 


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 antennz as we have 
described, at least they seem to show that there were 
appendages of some sort starting out sideways from 
each side ‘or the labrum, “just asin Apus:” Peas 
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, 


Fic. 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 antenne 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 antennee, because: 

(2.), We. thiak that, if the place: assigned) by 
Walcott to the three posterior head limbs is correct, 
some form of antenne must have been present, if not as 
antenne 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. XIII THE TRIEOBITES 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. Walcott’s restora- 
tion, given in Fig. 50, is therefore so far incomplete. 
The mouth, which is covered by the large labrum, mast 
have had some kind of appendages bordering it on 
cach side. When therefore we find clear traces of such 


FG. 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 antenne 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 antenne. ‘This homology depends, it is 
true, upon the truth of our main argument that the 
Trilobites, like Apus, were originally bent Anncelids, 


() 


1. 


226 THE APODIDEA: PART II 


and further, upon our homology of the large loco- 
motory limbs with the first trunk limbs; to this 
latter: point we. “shall seturn, "an: 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 
antennze like the Apodidze and the typical Crus- 
tacca. 


I 


PR DY 


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


aS s 


When now we come to ask how these antenne 
were developed, we can only conjecture that in some 
way or other they must have supplied the opening of 
the cesophagus with jaws, or perhaps with simpler 
instruments for pushing in food. It is improbable 
that the first antennz 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. XIII THES TRILOBITES Dy 


first antennz being simply sensory cirri ; and secondly 
because in no other group of early Crustaceans do 
the anterior antenne 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 antenna 
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 antenn& 
on each side of the labrum, and a pair of posterior 
antennz, 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 
cesophagus ; 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 Eurypteride, highly developed 
to function as chief mandibles. The crushed food 
would have to be forwarded towards the opening 
of the cesophagus, 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 


Or 2 
= 


228 THE -APODIDG PART II 


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 Eurypteride we 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 antenne be to function as antenne ; they 
might either degenerate as they have done in Apus, 
and, according to Walcott’s restoration, in the Trilo- 
bites, or they might function as seizing organs or mouth 
parts, asin 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 antennz 
as sensory organs, but also to the travelling of the 
antennz themselves towards the anterioreend of the 
body, an advantage for the animals which has enabled 


SECT. XIII ibe TRReOBTTES 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 antenne. 

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. 
We have already seen, further, that Limulus—and we 
shall see that the Eurypteridz, with some exceptions— 
not only possessed the two pairs of antennz, but also 
the large locomotory limbs as the sixth pair, zz. 
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 APODIDA PART 1] 


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, 


Fic. 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 
enathobase or accessory jaw for the holding and 
forwarding of food to the mouth, ze. on a certain 


SECR. cial te LT RTEOBILES 23% 


number of Jimbs 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 APODID7E PART II 


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 ina 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 Trilobite may be supposed to have arisen 
as follows, A‘<sirip, 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 rolled-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. 


SHGAS SeLLy ee, ERECOBILTES 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 Eurypteridze 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-seé -to -have™ beem the-case also, in the: Eury- 
pteridz ; the disadvantages of this arrangement will, 
however, be pointed out later. 

We have alrcady 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 APODTDAs PART II 


highly developed. The use of: such a specialised 
limb in the Trilobites, however, is difficult at first 
sisht,"to. see. In ~Apusi we find 1podeveloped vacwa 
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 
simpliication of the limbs irom. front” to pback 
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. 

The 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. 


Ww 
Oo 
ea 


SECT. XAII THE TRILOBITES 


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, ze. 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). 


We 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 primt- 
tive Crustacean known to the palzontologist should 
show so many points in its organisation directly 
deducible from the Annelids, z.e. deducible after the 
Apodidez 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 APODIDA PART il 


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 Apodide (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 oriam, and, more or Tess 
modified, the browsing habit of life. 


SiO TION. AAV 


THE EURYPTERIDA 


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 Xiphosuride 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 
Apodidz 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 Xiphosuride. 


238 THE APODEDAL “PART 


In the Eurypteridz, 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- 
uridz, 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, ze. 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 Xiphosuride. 

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 
Apodide, 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 EURYPTERIDAS 239 


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


ont 
ae 
a pw 


Fic. s5.—Pterygotus Osiliensis, upper Silurian, after F. Schmidt (from Zittell), show- 
ing five pairs of cephalic limbs, the enormously developed first pair of antennz, and 
first trunk limbs in which the dorsal and ventral parapodia are greatly developed 
as locomotory and masticatory limbs respectively. The second pair of cephalic 


limbs sometimes disappear, asin 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 APODIDA PART II 


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 Apodide and Trilobites. 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- 
urida 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- 


~ 


Shea, XIV THE ERY TERIDA 24i 


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 
ievCaile in as lime with it*) Ay natural concomitant 
change in the under lips would also take place; 
they would develop into the large metastomata found 
in the Eurypteride, 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 three small pairs of posterior cephalic limbs 
may have assisted in this latter process. 


242 THE APODIDA PART II 


comparative smallness 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 are 
analogous to the mandibles, while those of the four 
posterior head limbs probably function as maxilla, 
their dorsal parapodia doubtless helping in the catch- 
ing and holding of prey. And lastly, the first antenne 
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 antc- 
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 


SHCLARIV THe TUR YPTERID A: 24 


Os 


highly sensitive antenne, 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 Xiphosuride as early Trilobites 
specialised for slow deliberate browsing ; the Eury- 
pteridz 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.! 

This’ deseription: of the manner of life of: these 
animals (the Eurypteridz) 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 antenne 
developed into long chelate seizing feet, like the first 
antenne of Limulus, but much more highly developed. 
The analogy of the Scorpionide will at once suggest 
itself, where for the same purpose the palps have 
developed in the same way. These chelz 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 APODIDA PART II 


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 
maxilla 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 chele 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 maxillz, 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, ze. 
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 EURYPTERIDAL 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 


Fic. 56.—Eurypterus Fischeri Eichw. : Upper Silurian, natural size, after F. Schmidt 
(from Zittel’s Handbuch der Palwontologie). 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 antenne 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 APODIDA: PART II 


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

This degeneration of the anterior antenne in 
Eurypterus is hardly what we should have expected 
theoretically. The rapid forward movement for feed- 
ing would seem to require highly developed antennz 
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 ina limb should be 
immediately followed by an act of seizing, by means 
of its masticatory ridges. The anterior antenne 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 Kurypterus, are the sensory 
cirri alone of the original parapodia. We see no reason 
why this should not be the case. Nature seems to 
delight in every possible variation, and indeed in the 


SECT. KIV THE EURYPTERLD A: 247 


limbs of the modern Crustacea we have almost every 
possible combination of the parts of an Annelid para- 
podium. We have, for example, the sensory cirrus 
alone in the antenne, 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 upa 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 Eurypteride 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 APODPIDA: PART II 


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 was 
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, 
which in point of size reached the highest develop- 
ment of all the Crustacean descendants of our car- 
nivorous Annelids. The exact relationship of the 
eroup to the Trilobites and the Xiphosuride, 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 Xiphosuridz and Eurypteride 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 Pe DURY PT ERID A: 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 
cesophagus as possible, otherwise the greater part of 
the juices of the crushed animal would be lost before 
it could reach its destination within the cesophagus of 
its devourer. The enormous metastoma or under lip 
of the Eurypteride 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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SGAVANMOG CVE dd, OL, SNNOTOO, AHL 


SECT. XIV THE RURYP PERIDAL 251 


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 
Tracheata. 


SECTION. 2oy 


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 4s;sall 
extant, and to the palzozoic Trilobites and Eury- 
pteride. 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 palazozoic Crustacea which we have 
so far mentioned, viz. the Trilobites, Xiphosuride, 


sHem xv- CLASSIFICATION OF CRUSTACEA 253 


Eurypteride, 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 fo: 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 palzozoic forms. It is, however, as 
completely out of the sphere of this book as it is 


254 THE APODIDA: PARTY Il 


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 
Apodide, 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 proposc 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 Eurypterida and the Xiphosuride. 


Sper xy (CLASSI PICATION: 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 


Limulus Ostracoda Cirripedia Copepoda Apus Phyllopoda Malacostraca 


Nebalide 


S 
s 
2 
RS 
N 
& 


Ceratiocaris 


Hy menocaris 


2 larva adult 
Eurypteride Xephosuride Trilobite A podide 


The Crustacean 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 that they have 
had at least a twofold origin. 


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 APODEDTE PART Il 


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 fossiliferous 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~bejexplained : either” (1), 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. 
We may well suppose that some Trilobites adopted 
this method of protecting themselves, since, besides 


Fic. 57.—Harpes ungula Sternb. A, dorsal view; Z, rolled up in profile (from 
Bronn’s Adassen und 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 
S 


258 THE APODIDAL PART Il 


_ 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. 

(1) 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- 


me 
, ele 
» 


\\y uN yi \ Zag 
WARE 


Fic. 53.—Cypris fasciata (from Bronn’s Klassen und Ordnungen des Thierreichs) 
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. 574, 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 ridgeof 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 


Fic. 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. 7, intestine; 7, hepatic diverticula. 


diverticula of the mid-gut. In across section through 
an Ostracod the observer is at once struck by the fact 
that the space between the laminz 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. 
» 2 


260 THE APODIDZ: PART II 


(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. 574, 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” tubegcle 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 
LO. 


sreow wy  CRASsIPIzATION OF CRUSTACEA 261 


fs tosthe 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 
fadiaves.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 
larve 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 Trilobite. This 


262 THE APODIDA PART II 


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 larve 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 


seGuexy CLUASSIFICAELONVOFP CRUSTACEA 263 


the group in the light of what is known as to the 
origin of the whole class from a bent Annclid. 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 
sroups 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 frec- 
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 larve of early Apodide, the conditions, as 
far as we know them, would be fairly well satisfied. 
The Apodide 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 Apodide 
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 larve finds some support in the 


264 THE APODIDAS PART II 


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, ze. 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- 
didz into the land, and: the larval to remaim attire 
larval stage, would tend to differentiate the larve 
themselves, as the wind has differentiated the beetles 
in the island of Madeira. These insects are either 
strong flicrs 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, Oriein of Species, p. 109. 
? ro) ] 


Spem xy ‘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, z.e. 1, 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, ze. the 
endo- and exopodite of the typical Copepod limb. 

Grenachers 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 “retine” are 
developed, each consisting of comparatively few 
retinal cells. “Vhere is. no trace of crystal. cones 
or rhabdomeres, and the pigment is in the centre 
o® the  eroup.. Whe aerves. from. the. retinal. cells 
come-ftom ‘their immer 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/ PART II 


from a larval stage of Apus, dating back to the time 
when the Apodidz could no longer develop fully in 
the open sea, and only those larve which were acci- 
dentally shut off and isolated in lagoons were able to 
erow 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, ze. larval Apus, which sought 
safety in a stationary life. As larve 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 Apodide will afford some capable zoologist a 
field for research which cannot fail to be rich in 


Seem xv CLASSIFICATION OF CRUSTACEA 267 


biological observations of extreme interest. 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 
eut to be a group’ of eggs; thrust in as-—far as 
possible) under the mecki~ 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 Apodide 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 
eges 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 second, 
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 APODIDZ PART II 


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 Apodidz 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..crow up so as ito Close ‘the 
posterior opening (see 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 Cladocera, for instance in the 
well-known Daphnia pulex or water flea (seq Fig: 
60). 


Fic. 60.—Daphnia (after Claus), showing the brood cavity (4c) 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 Apodide, it is perhaps worth 
suggesting (1) that it would originally be only very 
young Apodidz, whose shells were specially.large in 


270 THE -APODILDA: PART If 


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 Apodidz 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 Cladecera,sand “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 Apodide to collect eggs under the loosening 
cuticle; flis ‘certainly “seems to, be “the .case, irem 
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 eggs under the shield would naturally lead to 
some special arrangement for times of ecdysis. Hence 
the ephippiurm 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. 


Sheet. xv CLASSIFICATION OF CRUSTACEA 27% 


THE ESTHERIDA. 


This is the only other group of the Phyllopoda 
about which we have a few words tosay. 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 


UAV eR 
ay a Se 


oS VY a RRs 
ZL 


Fic. 61.—Limnetis brachyurus, @ O. F. Miiller (from Bronn’s Avassen und 
Ordnungen), to show the position of the head as transition stage between the 
Cladocera and the Estheride (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 APODIDA PART II 


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, z.c. 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 


Fic. 62.—Estheria Donacitormis Baird 9 , to show 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 


shen avy CLASSIFICATION, OF CRUSTACEA 208 


the Apodidz 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 
croup of all to trace back to Apus. 

The Malacostraca have, by general consent, been 
traced back to Packard’s Phyllocaride, 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 palzozoic times the remains of large 
Crustacea, which appear to be true Nebalide. The 
most important of these are the two forms Hymeno- 
canis. and —Geratiocaris (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. 

ay 


274. THE “APODIDZA 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 Apodide. This first 
impression is fully borne out when we come to examine 
the forms more closely. We find several striking 
characteristics of the Apodidz, which convince us that 
we really have here to do with animals at least closely 
related to and easily derivable from Apus. 


Fic. 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 a5 


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 Apodide. 
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, 


Fic. 64.—Ceratiocaris papilio Salter. Upper Silurian. Showing the rostrum, the 
first pair of antenne, 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 antenna. The antenne bear a 
~ close resemblance to those of Apus (sce Fig. 7A, p. 34) 
tT 2 


276 THE (APODILDA: PART II 


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 antennz 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 antennz forwards as 
organs of sense. Just as, in Apus, the eyes travelled 
forwards, so, in process of time, the antennz 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 antennz). 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 antennze 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, 


ssem xv CLASSIFICATION: OF CRUSTACEA 277 


This view explains the morphological significance 
of the rostrum, as the protective point for the more or 
less delicate antenne, arising, not fer se, but as the 
further development of the tip of the middle piece 
between the two depressions along which the antennz 
travelled forwards. 

From all that remains then of these primitive 
Nebalidz we see a sufficient resemblance to the 
Apodide 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, z¢. the gradual 
migration of the antennz 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 
afe 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 Apodide in detail would 


278 THE APODIDA PART II 


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. 


Sic RLON 2XV1 
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 Chetopods, 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 PART II 


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 dent 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 trachee and the Malpighian tubules 
which are confined to the latter? 

The Annelid which gave rise to the Tracheéata 
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. 


Shemexvi 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 antennz 
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 maxille. 
In the Arachnida they form the powerful and vari- 
ously modified chelicerze which develop so largely as 
to displace and lead to the degeneration of the pro- 
stomium and antenne. These formidable jaws are 
assisted by the second pair of parapodia as accessory 


282 THE: APODIDA PART II 
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 papille ; 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 
maxille ; 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 maxilla, in Peripatus 
and the Arachnida function as the first pair of feet. 
In the Myriapoda and Hexapoda they form the 
posterior maxille. 

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 (seoments:: .Ine@both: cases: somesotuiue 
groups can be shown to have been differentiated direct 
from the original Annelid, while others are enly later 
modifications of such groups. In the Crustacea we 
think the Apodidez, 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 


SECi: 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 
ceca, the Malpighian tubules, were developed, pre- 
sents no difficulty. 

We 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 APODIDA PART II 


nida with the Xiphosuride and Eurypteride—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 Xiphosuride and the Scorpionidz, 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 


ERenevn: oti beet ANI ETB TRACHEATA /285 


point, for in most other respects all Arthropodan legs 
strongly resemble one another, and the presence of 
chelz ona 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 Scorpionide any 
special development of the sixth pair of limbs such as 
we have shown to be characteristic not only of the 
Apodide but of the Trilobite, the Xiphosuride, and 
the Eurypterida, and which is especially marked 
in the last, although this is claimed as a transition 
form between the Xiphosuridz 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 trachez 
and the Malpighian vessels, the presence of which in 
the Scorpionidze and other Arachnida classes them 
unmistakably with the Tracheata. 

Even if we admit the possibility of the concurrent 
development of tracheze 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: APODIDZA PART II 


The origin of the book-leaf tracheze from the gills 
of the Xiphosuride, 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 
tracheze within the Arachnida. We think that the 
evidence in favour of the new classification, to be 
drawn from the form of the trachez, -is not cori- 
vincing. 

The most probable origin of the tracheze 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 dermal 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 trachez 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 
trachez from dermal glands receives some support also 

1 Tf these include the coxal glands, it may throw light upon the 


developmental relations between the book-leaf trachez of the Arachnida 
and their rudimentary abdominal limbs. 


5 ow 


SEGE XV0 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 
excreviony mecessitates the further development: of 
other excretory surfaces. The advantages of the 
Malpighian tubules, or glandular ceca of the hind-gut, 
over excretory organs in any other part of the body 
have been already dwelt upon. This physiological 
connection between trachee and Malpighian vesicles 
which lessens the improbability of their concurrent | 
development ¢wzce, cannot however be taken advantage 
of in the special case under discussion, because the 
trachee 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 antennz in the 
embryo of a spider,! 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 antennz, in the form of the limbs and trachee, 


1 Trochosa Singoriensis Laxm. See Zool. Anzeiger, May 11, 1891. 


288 PHE APODID ZA: PART II 


and in their general organisation, but in the posses- 
sion of a sternal plate or entosternite. Phe 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 clements 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: ZTracheatan-Annelid at a ‘stage when ~ the 
Annelidan segments were still of the typical form, 
i.c. 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 zudzrectly a most 
remarkable transition form, having preserved so many 
characteristics of the common racial Tracheatan- 


Stem 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. 


Pub, NTS - 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 
antennz 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 antennz 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 


APPEN DIX. 1 


N 
Ne) 
N 


(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 11 
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, ! 
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” were of all 
sizes (from 1-2 p), 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, I.40 n.a., 
eye-piece No. 12, giving 1500 diam. 


204 APPENDIX 


Although we cannot be certain that we have seen the 
nuclei of the large pigment cells as shown in the dia- 
eram (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-lhke process of the pigment 
cells. Were the granules semi-independent cells, their 
arrangement could hardly be so straight and regular. We 


Fic. 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; z.enat 
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 
eranules 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. 


AU Ee MOTOR hy 
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 (1) 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 alow power, take the form of triangular wings. 
These connective tissue alz are not flat and membranous, 


APPENDIX Ii 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 


Fic. 66.— Diagram to illustrate the plan of the circulation in the anterior part of the 
body. The blood propelled by the heart (Z) through the head, is returned 
through the intestinal sinus (7s) from which in each segment it escapes ventrally 
into the limbs. Its course is indicated diagrammatically by the arrows(Z). From 
the limbs it returns through the dermo-muscular sinus in each segment into the 
cardial sinus (cs). 2, intestine; ac, aorta cephalica; Z, 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 


Fic. 67.—Diagram of the circulation from above. 4, heart, expanded by the fine 
connective tissue alz (a); s, segmental septa forming the transverse walls of 
the dermo-muscular sinuses (Zs); @c, cephalic aorta; g, lateral artery to shell 
ang shell gland ; 2, 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, Ze. as 
far as the heart extends. Thus while part of the blood, 
flowing through the intestinal sinus, passes down through 


APPENDIX III | 366 


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 imbs. 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 venze 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. 


APEeNDIX 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 nght), 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), z.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 


HY 
; 


Fic. 68.—Part of a section of the urinary canal (shell gland of Apus).° 677, basal 
membrane on which rests the harder supporting framework, seen in tangential 
section (at ¢s) to have a spongy structure; #4, inner layer of protoplasm 3 2, 
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. Claus 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. 


1! Wh; ay Ui, I, yp 
mI ML, 
TROY i, 


AY 


Se if ifs 


PA ¢ 7 
INS. CHEB 
LK Ye SI laecand 3 ni 
ALLA RAHAT I IA 
MENEZ MRA 


af. ie ‘ ti “l ty G 
GU OP aE! 
Y LA . Malibu: 


CR RK 
Lorain 
Sai 


P 


Fic. 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 (f) 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 Apodide. 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.’ 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- 


cw 


Fic. 70.—Diagram of a section of the neck gland or dorsal organ of Apus, drawn 


upside down (z.e. as the animal swims) so as the better to illustrate the catching 
of the laden blood corpuscles in the connective tissue net. #, epithelium of 
the mid-gut ; c, connective tissue belonging to the longitudinal musculature ; ca 
blood stream from the aorta cephalica ; 4, blood corpuscles laden with excretory 
matter; cz, 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- 
Xx 


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,! 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 larve, 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 Daphnidze, 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. 


APP BES 


REPRODUCTION 


A. FERMAPHRODITISM OF THE APODID. ! 
(From Nature, vol. xlill. p. 343.) 


THE reproduction of Agus 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.? 

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 AZassen und Ordnungen 
des Thierretchs, 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 Lehrbuch 
der Vergleichende Anatomie, 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 (/enaische Zettschrift fir 
Naturwissenschaft, Band xxv., N.F. xvi.) announcing 
the hermaphroditism of L. Spitzbergensis, knowing how 
much the reproduction of the Apodidz 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 Rey. 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 


310 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 moneecious 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 Apodidz which I hope soon to 
have ready for the press.! 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- 
hum cells dislodged by the shrinking of the membrane of 
the genital tube, which is stretched some 100-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 ” (ze. 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 Apodidz, 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 Apodide. 
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-hke grains of yolk witha 
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! 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 ze-arrangement 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 (Zm- 
bryology), Claus (Zoology), Gegenbaur, Huxley, and Lang 
(Comparative Anatomy), Gerstaecker (Crustacea, Vol V. of 
Bronn’s KZassen und Ordnungen des Thierretchs), Haeckel 
Natirliche Schipfungsgeschichte), Rollston (Forms of Animal 
Life), and Zittel, Nicholson, and Lyddeker (Pa/gontology), 
the following are the chief works which have been 
consulted :— 

Bairp.—A Monograph of the Family Apodide. Proceedings 
of Zool. Soc., London, 1853. 

BARRANDE.—Ssystéme Silurien de la Bohéme. Vol. I. and 
Suppt. 

BEDDARD.—On the possible Origin of the Malpighian 
Tubules in the Arthropoda. Ann. V7. (6) IV. 

BouRNE (see Lankester). 

Brapy.—-A Monograph of recent British Ostracoda. 77. 
Ligne. see, Nol:| XXVT. 

BRAUER.—Ueber die Entwickelung des Lepidurus pro- 
ductus, in Siés. Ber. d. K. Acad. ad. Wassensch, 
Wien, Nol. LXEX.; Pt. 1.1374. 


1 0.4,M,S, Vol, XXVI. 


314 LITERATURE 


BRAUER.—Beitrage zur Kenntniss der Phyllopoden. In the 
same, Vol; (LXV.,"<Pt)..L.)\ 1872; vand sfurtherag 
1878. 
BUCHANAN.—Respiratory Organs in Decapodous Crustacea. 
OI MiS« iN Oo. 
BURMEISTER.—Die Organisation der Trilobiten aus ihren 
lebenden Verwandten entwickelt. Berlin, 1843. 
CARRIERE.—Die Sehorgane der Thiere. Miinchen und 
Leipzig, 1885. 
CLAPAREDE.—Les Annélides Chztopodes du golfe de 
Naples. Geneve, 1868, supplément, 1870. 
Ciaus.—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. 47d. Zool. Inst. Wien, 1886. 
, Neue Beitrage zur Morphologie der Crustaceen. 
Arb, Zool. Inst. Wien, 1886. 
,, Neue Beobachtungen tiber Cyprididen. Zed¢sch. 
fiir Wiss. Zool. Vol. XXIII. 
» Organisation of Nebalia, and Systematic Position 
of the Leptostraca. Ann. V.H. (6) ITI. 
Dourn.—Geschichte des Krebs-Stammes nach embryolo- 
gischen, anatomischen und _ palontologischen 
Quellen. /enazsche Zeitsch. Vol. VI., 1871. 

EnLers.—Die Borstenwiirmer. Leipzig, 1864-68. 

FERNALD.—The Relationships of Arthropoda. Studies from 
Biol. Lab. Johns Hopkins University. Vol. IV., 
1890. 

V. GRABER.—Morphologischen Untersuchungen uber die 
Augen der frei-lebenden marinen Borstenwurmer. 
Bonn, 1879. 


LITERATURE 315 


GRENACHER.— Untersuchungen uber das Sehorgan der 
Arthropoden. Gottingen, 1879. 

GROBBEN.—Die Antennendruse der Crustaceen. 47d. Zool. 
Inst. Wien, 1880. 

»  Entwickelungsgeschichte der Moina rectirostris. 

Arb. Zool. Inst. Wien, 1879. 

GruBE.—Bemerkungen tber die Phyllopoden. Archiv fiir 
Naturwiss. Vol. XIX., 1853. 

HatscHEek.—Lehrbuch der Zoologie. 

HeERRICK—Metamorphosis and Morphology of certain 
Phyllopod Crustacea. Bulletin, Denison University, 


1885. 
Huxiry.—The Crayfish. International Science Series.— 
Ath Ed., 1884. 


Jones & Woopwarpv.— Monograph of the British Palzeozoic 
Phyllopoda, 1888. 

KINGSLEY.—Notes on the Embryology of Limulus. 
ONT |S. NO XX. 

KORSCHELT UND HEIDER.—Vergleichende Entwickelungs- 
geschichte der wirbellosen Thiere. Jena, 189r. 

Kroyer.—Apus glacialis. Vaturhist. Tidskrift. 2 Rak., 
1847. 

LANKESTER.—The Appendagesand Nervous System of Apus 
Cancriformis. Q./.AZ.Se. Vol. XXI. 

a wleimmulus an Arachnid: O17: 47.S.,. XXI.° And 
further Vol. XXV. 

LANKESTER AND BouRNE.—The minute Structure of the 
lateral and central Eyes of Scorpio and Limulus. 
C707S: Vol. XXINT. 

LILLJEBORG.—Synopsis Crustaceorum Svecicorum. Reg. 
Soc. Sc. Upsaliensi. Tradita die vu. April. 1877. 

Lupsock.—Notes on some New and Little Known Species 
of Freshwater Entomostraca. 7vrans. Linnean Soc., 
1863. 


316 LITERATURE 


Lusrock.—Origin and Metamorphosis of Insects. Vature 
Series, 1890. 
MILNE Epwarps.-—Recherches sur l’Anatomie des Limules. 
Anal. Se. Nat. sth Série: T) XV" Paris es ye: 
MU.uer, 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. 
Micra Sc Vol, SOV. 
Watcotr.—The Trilobite. Old and New Evidence relating 
to its Organization. Bull. Mus. Comp. Zool. 
Vol. VIII. Camb., Mass., U.S., 1880-81. 
WatasE.—On the Morphology of the Compound Eyes of 
Arthropods. Studies from Biol. Lab. Johns 
Hopkins University. Vol. IV., 1890. 
Woopwarp.—British Fossil Crustacea. 1872. 
ZADDACH.—De Apodis Cancriformis. Bonn, 1841. 
ZENKER.—Monographie der Ostracoden. 1854. 
&e., &c. 


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BOOKS FOR THE YOUNG—ZOOLOGY. 39 
a SE eR LS 


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40 


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ALEXANDER Suishap) 33 
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ANDERSON (A.). I4 
ANDERSON (Dr. McCall) . 22 
AnpreEws (Dr. Thomas) . 26 


AppLETON (T.G.) vei 
ARCHER-Htnp (R. D. ) 
ARNOLD, M. 8,14, 195 20, 21, 30 
Arno. (Dr. T.) Ee 
ARNOLD (W. T. ) : mois) 
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WHITE (Gilbert). (See NaTuraL History.) 


INDEX. 


PAGE PAGE 
ATTWELL (H.). re 20 | BERNARD (J. H.) 25 
AusTIN (Alfred) ‘ 14 | BERNARD (M.). 12 
AUTENRIETH (Georg) . 7 | BERNERS(J.) II 
Awpry(F.) . : . 38 | Besant (W.) . 4 
Bacon (Francis) 5 195 20 | BETHUNE- ae J. F. yi 33 
BaInEs (Rev. E.) . Betrany (G. T.) 6 
Peete S. W.) 28, 30, 37; 33 BicKERTON (T. H. ). 22 
Batcu (Elizabeth) . . 12 | BriceLtow(M. M.) I2 
Batpwin (Prof. J.M.) . 26] Bixfias(D.) . 17 
BALFOUR ee Hon. A.J.) 25 | Binnre (Rev. W.) . Sage 
BatFrour(F.M.) . ~ 5,6 | BIRKS((DRS) i Ones peomee 
BaLFour (J. B.) ; . 6] ByOrNson (B.). ; Mkt, 
BALL (V.). . . 38) Back (W.) 4 
BALL ae Platt) : . 6 | BLacKBURNE (E. yay oo 
Batt (W.W.R.)  . > 22 | BLACKIE .S:) «10, 14; EO 
BALLANCE(C.A.) . . 22| BLaKE (J. F.) 4 Men c: 
BARKER (Lady) - 2, 8,37 | BLAKE (W.) ‘ AS ac 
BARNARD (C.) . . 27 | Braxiston(J.R.) . so RS 
Barnes (W.) . : - 3 | Branrorp(H.F.) . Q, 27 
Barry (Bishop). . 33 | BLanrorp (W.T.) . Q, 24 
BARTHOLOMEW (J.G.) . 3 | BLomFIELD (R.) ' Fleas, 
BARTLETT (J.). s 21097) Biyrai(Anws) E o TR 
BARWELL (R.) . 22 | BOumM-BawerkK (Prof.) . 28 
BASTABLE (Prof. CaN E) 28 | BorssEvAIN (G. M.). . 28 
BasTIAN (H. C.) 6, 22 | BoLDREWooD (Relt aie) 
BaTESON (W.) . - 6] Bonar (J.) . 28 
Batu (Marquis of) . 28 | Bonn (Rev. J. ye yee 
BATHER (Archdeacon) 33 Boots (G.) : . 26 
BaxtTER (L.) ; . 3 | Boucuton (G.H. ee - 37m 
BEEsLY (Mrs. e é “ Boutmy (E.) . - I2 
BENHAM (Rev. W. ) . 5, 20, 32 | BowEn (H.C.). . (2505 
Benson (Archbishop) 32, 33 | BowER(F.O.) . - ion 
BeEruioz (H. : - 3] Brivcgs (J. A.). 


INDEX. 41 
PAGE PAGE PAGE 
BricutT (H. A.). son CrARrice (Cz B-): 9, 28 | Dittwyn(E. A.) . LT, 
BriGHT (John) . . 28, 29 | CLausius(R.) . J . 27) Dopsson(A.) . : aka: 
BriMLeEy(G.) . 5 - 19 | CLIFFORD (Ed.) : DOoNALDSON (J.) 3 -/° 33 
Bropie (Sir B.C.) . - 7 | CiirForp (W.K.) . 19, 26 | DONISTHORPE (W.) . 25 
BroprisBp(W.J.) . 13, 37 | CLirForp (Mrs. W. K.) 38 | DowpEn (E.) . 25451135015 
BRooKE (Sir J.) 5 2) 34] Croucm (A. H:) Sl) 245704 Dover (Sir bE.) pc a! 
Brooke (S. A.) 13, 14, 21, 33 | CoBDEN(R.) . : $290)| Doyv7E (--A:)) *- A 5 tite) 
Brooxs (Bishop) . 4 334| (COHEN (..B-.).- 5 - 7) Drake (B.) 530 
Brown (A.C.) . 2 . 26 | CoLtENso(J.W.)  . . 32 | DrumMMoND (Prof. Js nist 
Brown (J.A.) . ey ers | | (COLERIDGE) (8: .)) 4. ska SORVDEN  {. 5) 120 
Brown (Dr. James) . . 4 | CoLuiER (Hon. John) - 2| Du Cane (EK. F. ity 29 
Brown (T. E.) . 14 | CoLuirins (J. Churton) . 19 | DurF(SirM.E.Grant) 20 29537 
BRowneE (J. 74 B. yi . rz | CotquHouNn(F.S.) . . 14 | DuNnsmutrR (A.). : sty, 
Brownz (Sir x rape . 20 | Cotvin (Sidney) : 4, 20 | DUNTZER (H.). i ovina iG 
Browne (W.R.) ComBE (G.) - 8 | Dupré (A.) : 3 awa, 
Brunton(Dr.T. Lauder) 22, a CONGREVE (Rev. Me ye - 33 | Dyer (L.). . 4 AE 
Bryce (James) . - 9, 28, 37 | Conway(Hugh) . . 17 | Eapie (J.). 4, 30) 31 
BucHHEIM (C.A.) . 2.201) CooK(B.E) ie. . 2| EASTLAKE (Lady) $ sage 
BuckKLAND oe ): : en i5)| Coox: (C- Kinloch) . . 24 | Expers(G.) 17 
iBockiny (A.B). a rofl Coors: Gib). 7, 34 | EDGEWORTH (Prof. ie Y.). 28 
BucKNILL (Dr. Cs) . 22 | CORBETT (J.) : 4 17, 33 | EpmunpDs (Dr. W.) . 22 
BuckrTon (G. B.) . 40 | CoRFIELD (W. H.) . 11 | Epwarps- Moss (Sir J. E. ) 30 
Bunyan . - 4,19, 20 | Corry (T.H.) . £168) HimEer (G.I) A ye 6 
Burcon (J. W. ) s . 14 | COTTERILL (J. H. ie é . 8 | Experton (W.A.) . a 719 
Burke (E.) 4 . 28 | CoTToNn Gepece) : . 34 | ELLERTON (Rev. J.)- s4 
Burn (R.). 1 | Cotton (C.) BS ee i2e| erion (elon. A.) ee. 529 
BuRNETT (F. Hodgson) « 17 | CoTTon(J.S.) . 2 wezou errs (E2) é Z1RI2 
BuRNS : 14, 20 | CouEs (E.) . 40 | EMERSON one wW. )) xe 4, 20 
Bury (J. B.) Cour! .iOPE (Ww. Hie ). . 4] Evans (S.) , Aeayt 
BuTCHER (Prof. S$. ) £5,19,30 CowELL(G.) . . 23 | Everett (J.D. ) 3 2) 120 
But er (A. J.). 37 | CowPER . 3 : . 20 | FaLtconeR (Lanoe) . WELZ 
BuTLer (Rev.G.) . 2633)| Cox (G. V.) , g | FARRAR (Archdeacon) ch 3° 34 
BuTLer (Samuel) . . 4 | Crark(Mrs.)14, "a 19, 20, 37, 38 | Farrer (SirT.H.) . 29 
BuT Ler (W. Archer) 133)) Grark (EL) 8, 29 | FAULKNER (F.). 7, 
But er (Sir W. F. - . 4 | CRANE(Lucy) . 5 2, 39 | Fawcett (Prof. H. ). 28, 29 
Byron. . 20 | CRANE (Walter) . 39 | FawcetT(M.G.) . 5, 28 
CarrNEs (J. E. ) . 29 | CRAVEN (Mrs. D.) . §8| Fay (Amy) ‘ : ey 
Ca.LpeEcoTt (R.) 12, 38, 39 | CRAwForD(F.M.) . . 17 | FEARNLEY (W.) 3 Up n27, 
CaLpERwoobD (Prof. H.) CREIGHTON (Bishop M.) 4, 10 | FEaRon (D. R.) F gourd 
8, 25, 26, 33 Bs A ae ae ai 8 | FERREL(W.) . 2 De 
CaLveErT (Rev. A.) . . 31 | Cross (J. A.) . 30 | Ferrers(N.M.) . R27 
Cameron (V.L.) . . 37 | CRossLey (E.) . g . 2 | FESSENDEN (C.) s + 26 
CAMPBELL (J. F.) . 37 | CrossLey (H.). : STA HUNG Ke (Eads) a2 Z SHAE 
CampeBELt (Dr. _M. ),. 33 ) Cummine (L.) .- ; . 26] FisHer(Rev.O.) . 26, 27 
CamMPBELL (Prof. Lewis) 5, 13 | CUNNINGHAM (C Mea 28 | Fiske (J.). 6, 10, 25, 291 34 
Capes (W.W.). } . 13 | CUNNINGHAM (Sir H.S. o 17 | Fison(L.). e : I 
Car_es (W. R.) : . 37 | CunnincuaM (Rev. J.) . 31] Firco(J.G.) . 8 
CarLyYLe(T.) . . 3 | CunNINGHAM(Rev. W)31,33,34 | Firz GERALD (Caroline) . 14 
CARMARTHEN (Lady) . 17 | CUNYNGHAME (SirA.T.). 24 | FiTzGERALD (Edward) 14, 20 
CARNARVON (Earl of) . 36 | CurTeis(Rev.G.H.) 32, 34 | Firzmaurice (Lord E.) . 5 
Carnot (N.L.G.) . . 27 | Daun (F.) : . 17 | Fieay(F.G.) . 2 Papo se 
ares (Bishop) . 33 | Daxyns (H. G.) : . 37 | FLEISCHER (E.). F ATLA, 
Carr (J.C.) E - 2| Dare (A. W. W.) . 31 | Freminc(G.) . SER, 
CARROLL (Lewis) . 26, 38 | DatTon (Rev. J. N.) . 37 | FLower (Prof. W. ial reso 
CarTER (R. EeeenelD . 23 | DANTE - 3) 13, 37 | FLicxicer(F.A.) . 23 
Casset (Dr. D.) . 9 | Davies er a Li). 20, 31, 34 | Forspes(A.)_ . J 4, 37 
Cautrey(G:S.)  . . 4 | Davies (W.) 5 | Forspes(Prof.G.)_ . at ek: 
CazENovVE (J.G.) . . 33 | Dawkins (W. B. ese . 1 | Forses (Rev. G. H.) peiga 
CmatMeErRS(J.B.) . 2 Dawson (G.M.)_. . 9| Foster (Prof.M.) . 6, 27 
CwHacMErRS (M.D.) . 29 | Dawson (Sir J. W.) . 9 | Foruercitt (Dr. J.M.) 8, 23 
CHAPMAN (Elizabeth R. 7 14 | Dawson (J.) . 3 . | FowLe (Rev. T.W.). 29, 34 
CHASSERESSE as Mao |) Davi. B:) t7: E . 17 | FowLerR(Rev.T.) . 4, 25 
Cuerry (R. R.) erat | Dav GR. E(t: 5 . 26} FowLeR(W.W.)  . . 24 
CHEYNE or EESEI)\ (3 20 02))|) DEFor (D:) 8 - 4, 20 | Fox (Dr. Wilson) . 123 
CHEyneE (T. K.) ; . 30 | DeicuTon (K.). . 15 | FoxweE tt (Prof. H. - . 28 
CuRISTIE (J.) . : . 23 | DELAMoTTE (P. H. ). . 2] Framyjr(D.) VEO 
CurisTiE(W.D.) . seco |) Derr (E.C.)0r 3 . 12 | FRANKLAND (P. F. ve er 
CuurcH(Prof.A.H.) . 6] DE Morcan(M.) . . 39 | FRASER(Bishop) . 34 
CuurcH (Rev.A.J.) 4, 30,37 | DE VERE(A.) . é . 20 | FRASER- ee (C. e ie 14 
Cuurcu (F. J.). 20, 37 | Dicey(A.V.) . 12, 29 | FRAZER (J.G.). aA 
CuurcH (Dean) 314943)19)34133 Dickens (C.) . 5,17 | FREDERICK (Mrs. > 4 tt A. 
Cuark (J. W.) . 5 DIGGLE (Rev. J. W. ). . 34 | FREEMAN (Prof. E. A.) 
CrarK (L.) p E . 2 | Ditxke (Ashton W.) . 19 2, 4s IO, 29, 32 
Cviark (S.) F - - 3 | Dike (Sir Charles W. i 22 29 | FRENCH (G. R.) A MT 


42 

PAGE 
FRIEDMANN (P.)  . et 
Frost (A.B.) . é Aleets) 
FROUDE (J. A.). F Jetted. 
FuLLerton (W. M.) af 37 
Furniss(Harry) . o0N38 
FuRNIVALL (F. J.) . VEY 
FyFFE(C.A.) . IS Av IO 
Fyre (H.H.) . E ALO 
GAIRDNER (J.). : sang: 
GALTON (F.) . ‘ 1,127 
GamMGEE (Arthur) . «27 
GARDNER (Percy) . Bye ie: 
GARNETT (R.). § Hina 
GARNETT OW. i : oH Os 
GASKELL (Mrs.) ql ane 
Gasxoin (Mrs. H.) . 4130 
GEDDEs (W. D.) 133137. 
Grr (W.H.) |. 25, 27 
GEIKIE (Sir A.). 4; 9, 27 
GENNADIUS (J.) . 17 
Gippins (H. de B. yin iro 
GipBon (Charles) . AWweS 
GitcuRisT (A.). : e's 
GILEs (P.). F (ace 
Gitman (N. P. ) é . 28 
GiLmor: (Rev. J.) . 13 
GuapsTonE (Dr. J.H.) 7,8 
GLADSTONE (W. E. 3 of has 
GLAISTER (E.) . 2,8 
Goprray (H.) . é was 
Gopkin (G.S.). 2 PRS 
GOETHE . : 4, 14 
GOLDSMITH 4, 12, 14, 20, 21 
GoopDALeE (Prof. G. [:) KP 26 
GOopFELLOw (J.)_. II 


Gorvon (General C. G. irs 4 
Gorpon (Lady Pa 3 Yar 


GoscHEN (Rt. Hon.G. J.). 28 
GossE eeeune) i 4, 13 
Gow (J.) . i eA ito 
GRAHAM (D. ne , Utara 
GRAHAM (J. W.) Aig] 
GRAND’ HOMME (E. x: 8538 
Gray (Prof. re) . 26 
Gray (Asa) : ed16 
GRAY . = wl dy ha er 
GREEN (J. R. - 9, 10, 12, 20 
GREEN (Mrs. J. Rn i A245 i05 ZO 
GREEN (W.S.). : 2 37 
GREENHILL (W.A.). 20 
GREENWOOD (J.E.). a3 

GRIFFITHS (W. eu) 4 ate3 
GRIMM. F ag 
Grove (Sir G. ). F 9, 24 
Guest (E.) : ; 2x0 
Guest (M. J.) . é Siiixo 
GUILLEMIN (A.) . +26, 27 
GuizoT (PA PEG») vas IGS 
Gunton ((G.) . : . 28 
HALEs (J. W.). 4 16;"20 
HaAttwarD (R.F.) . Wr 
HAMERTON (P.G.) . 2, 21 
HamIittTon (Prof. D. J. aravwes 
HAmILTON (J.). + 894 
Hansury (D.). : 6, 23 
Hannay (David) . 4 
Harpwick Sie a 31) 34 
Harpy (A.8.). 17 
Harpy(T.)  . 4 ADT 
Hare (A.W.) . : - 20 
Hare (J.C.) ©, 34 
HARPER (Father Thos. ) ma 34 
Harris (Rev. G. C.). melt 
Harrison (F.). 4, 5) 21 


INDEX. 


PAGE 
HARRISON (Miss J.). eAnT 
Harve (Bret) . J Ae tf 


HartTic (Dr. R.) 6 
Hart Ley (Prof. W. Nara: eae, 


Harwoop (G.). 21, 20, 32 
Haves (A.) 4 d IUtTH 
HEADLAM (W.). é 86g6 
He tps (Sir A.). : pakoT 
HEMPEL (Dr. W..) . AI 
HERODOTUS. : o.36 
HERRICK . : A ..| 20 
HERTEL (Dr.) . Amis! 
HILy a Davenport). 7.29 
Hity.(O:)). - 29 
Hiorns (A. H. \ 3 ei i23 
Hosart (Lord) ‘ Oar 
Hosppay(E.) . 3 HG. 
Hopecson (Rev. J.T.) . 4 
Hor Fpinc (Prof. H.) su(26 
Hormann (A.W.) . Sy. 
Hote (Rev. C.). . 7, 10 
Houipay (Henry) . regs 
HOLLAND (LT. EB.) %.. 1) iesieg 


Ho.titway-CaLTHRoP(H.) 38 
Hormes (O. W. eee ) rite 


HomMeER . 13, 36 
Hooker (Sir J. ‘D. Dine 6, 37 
Hoots (C. H.). 4 soareo) 
Hooper (G.)_ . EN EH Oth. 
Hooper (W. H.) 5 a vic2 
Hore(F.J.)  . : Se) 
Hopkins (E.) . . Ly Fra. 
Hoppus (M. A.M.) . 1/628 
HORACE . { - 13, 20 
Hort (Prof. F. J./A.). |) 30, 32 
Horton (Hon. S.D.) . 28 
HovENDEN (R.M.). VAY. 
HowE Lt (George) . AB 
Howes (G. B.) . 3 1840 
Howitt (A. W. ) 3 I 


Howson (Very Rev. y; S.) 32 
HozteEr (Col. i. M 

HwtpnerR (Baron) . 37 
HuGuEs (T.) 4, 15, 18, 20, 37 


Hut (E.). 2,9 
HUuLtAH (J.) H@yi2o; 4 
Hume (D.) 

Humpury(Prof. SirG.M. yoReae 
Hunt (W.) “| MLO 
Hunt (W.M. y) Wig 267) 
Hurroni GR: EH) 7 4 4, 21 


Hux ey (T.) 4, 21, 27, 28, 29, 40 


IppinGs (J. P.). 9 
ILLINGWORTH (Rev. in R. ) 34 
InGRAM (T. D.) : 2.50 
IRVING (J.) VAG 
IRVING (Washington) vie 
Jackson (Helen) . iw a8 
Jacos (Rev. J. A.) . Srey! 
James (Henry). a), noy 21 
James (Rev. H. A.) . Pig 
JAMES (Prof: Wi)! (3 }vi26 
JARDINE (Rev. R.) . : 26 
JEANS (Rev. G.E.) . 34, 37 
Jess (Prof. R. C.) 4, 10, 13 
JELLETT (Rev. J. H.) A134 
Jenxs (Prof. Ed.) . hey 
Jennines(A.C.)  . 10, 30 
Jevons (W.S.). 4, 26, 28, 29 
Jex-BLAKE (Sophia). Bars 
JouHNson (Amy) P - 27 
JoHNSON (Samuel) . SAMS oc) 
Jones (H.Arthur) . NET 43 


Jones (Prof. D. E.) . le7 


PAGE 
Benes CES): J f Sy 
Kant : : é s1725 
Kari 39 
KAVANAGH (Rt. ‘Hn. A.M. ) 4 
Kay (Rev. W.). : 3x 
Keary (Annie). 10, 18, 39 
Keary (Eliza) . - $99 
KerATSH et « 4, 20, 21 
KELLNER (Dr. 1 Nemes itor 
KE.LoGG (Rev. S.H.) . 34 
Kempe (A. B. 


ac 26 
KENNEDY (Prof. ALB. W.) 8 


KENNEDY (B. H.) 36 
KEYNES (J. N.). 26, 28 
KIeEPERT (H.) . i a ie, 
KILLEN (W. D. ) ietg2 


KINGSLEY (Charles) . 4, 8, Io, 
I1,12,13,15,18, 21 24, 32, 375 39 


Kinestey (Henry 20, 3 

KipuineG (J. L.). é 1538 
KrpLinG (Rudyard) . Ca ae 
KIRKPATRICK (Prof.) ange 
KLEIN (Dr. E.). / 23 
KNIGHT (W.) . Sey ea 
KUENEN (Prof. A. ae 2130 
KYNASTON (Rev. H. ) 34, 37 
LABBERTON (R. H.). yn 
LAFARGUE (P.). 2 aS 
LAMB. 4, 20, 21 
LANCIANI (Prof. R. ). Ore 
LANDAUER (J.). ‘ se 
LANDOR 4) 20 
LANE-POOLE (S. )inae 4 126) 
LANFREY (P.) . Z Wi O85, 
LANG (Andrew). 2; 12, 21,(36 


LanG (Prof. Arnold). - 39 
LANGLEY (J. N.) 27 
LANKESTER (Prof. Ray) 6, 21 


LaAstenn (RS). 6 
LeEaF (W.). 5 13, 36 | 
LEAHY (Sergeant) : 130 
Lea (M.) . Hens 
eee (S:)) se : la hoary, 
LEEPER oe Fs - Sega 
LEGGE (A. O.) . |} Mirela 
Lemon (Mark) . < 120 
LEs.iz (A.) 3 
LETHBRIDGE (Sir Roper) . 10 
Levy(Amy) . . 18 
Lewis (R.) ! ‘ “une 


LIGHTFOOT(Bp. )2%30,3%331 34 
LiGHTWwoop (J. M.). f 
Linpsay (Dr. J. A.). A Pr 


LocKYER (J. N.)- . 93) 7,27 
LopcGE (ProfiO2].), a eraem 
Loewy (B.) . . 26 
LorTiz (Mrs. W. J. ). oe 


LoNGFELLow (H.W.) . 30 
LONSDALE (J. ye ») | | 205 37 
Lowe (W. H.) . 2 Ago 
LowELt (J. R. ). I5, 21 
LuBsock (Sir J.) 6, 8, 21, 22, 40 


Lucas (F.) . < “ashe 
Lupton (S.) 3 PaaS, 
LYALL (Sir Alfred) : su 
LyTeE (H. C. M.) t ie 
LytTTon (Earl of) . Ae i: 
MacALIsTER (D.) . Neg 
MacarTHuUR(M.) . te 
Macautay(G.C.) . gg 
Maccoui (Norman). 2) ge 


M’Cosu (Dr. J.) 
Macpona.p (G.) : a 
MacDONELL (J.) é - 290 


PAGE 
MackalL Gia: W-) 37 
LACKENZIE (Sir Morell) . 23 
Mactacan (Dr. 7.). 23 


MACcLAaREN (Rev. Alex.) . 34 
Macvaren (Archibald) 2 
Macvtean (W.C.) 
Macear(Rev. Dr. GR: < 50138 
M’LENNAN (J.F.): . 
M’LEnnan (Malcoim) 18 
Macmi.Lian (Rev. H.)22,35,38 


MaAcMILLAN (Michael) 5; 15 
Macnamara (C.) (23 
Macguorp (K.S.) 18 
Mapoc(F.) . 18 
MAGUuIRE (J. F.) 39 


pa aEry (Prof. J. P.) 


II, 13, 22, 25, 35, 


cee (F. W.) «> 12, 29 
Ware T (1u:)t, i. 18 
NAT ORY-(Six Ey) 4: - 20 
MANSFIELD (C.. B.) . alg 
MARKHAM(C.R.) . tig 
Marriotr(J. A. R.). ME 
MAarsHAL.t (Prof. A.) 28 
MarsHartt(M.P) . 28 
Marte  (C.) 3 - 24 
MarTIN (Frances) . 3, 39 
MartTIn (Frederick). 28 
Martin (H.N.). . 40 
Martineau (H.) : 


MARTINEAU (J.). 
yeaa ) 4,5,15,16,20,22, 2b 
Masson (G 3 


Masson (R. @ ) ! 16 
MaTurRIN (Rev. W.). 35 
Maups ey (Dr. H.). 26 


Maurice (Fredk. Denison) 
8, 22, 25, 30; 31, 32, 35 


Maurice (Col.F.) . 5, 4 re 
Max Mier (F.) . 

Mayer (A. M.). : - = 
Mayor (J.B.) . ie yor 
Mayor (Prof. J. E. B.) Zu 
Mazini (L.) 39 
M’CormicKk (Ww. S ya Bate) 
ME Lpora (Prof. R.). 7, 26, 27 
MENDENHALL(T.C.) . 27 
MERrcIER (Dr. C.) 23 
Mercur (Prof. J.) 24 
MEREDITH (G.). I5 
MEREDITH ((L.A.) . I2 
MEVER (E. von) E SALE 
Miatt (A.) ‘ Pits 
MIcHELET (M. ) $ SE ba: 
Mir (HioR.). 2: 5 Ede 
MILLER(R. K.). a3 
MILLicANn (Rev. W. ): Sr35 
MILToNn . 13,1 5.)20 
MrnTo (Prof. W. ) 4, 18 
MitForpD(A.B.) . 2/438 
Mivarr (St. Grote): wyZs 
MIxTeEr (W. G.) S47 
MouHAMMAD . 20 
MoLeswortTu (Mrs. ) 39 
Mottoy (G.) . 26 
aan (Gere. ee 
Monte ius (O.)  . ee eT 
Moore (C. H.). £) 2 
MoorHousE (Bishop) e 5 
Morison (J. C.) 7 Al 
Morey (John). 3, 4, a "22 
Morris (Mowbray) . My et 
Morris (R.) EOS 
MorsHEab (E. D. A.) 36 


INDEX. 


PAGE 
MoutrTon (L. C.) Is 
MupieE(C.E.) . i525 
Muir(M.M.P.) .. TAL7 
MUivier(H.) . TALG 
MULLINGER (J. B. ed II 

_Mourpuy (J. J.). 26 
Murray (D. Christie) 18 
Murray (E.C.G.) . 38 
Myers (E.) _. I5, 36 
Mvers (F. W. H.) 4, 15, 22 
MYLne (Bishop) Bs 
Napa. (E.5S.) . 22 
NETTLESHIP(H.). . 13 
NeEwcastLE (Duke and 

Duchess) 5 - 20 
NeEwcoms (S.) . sts 
Newton (Sir C. T. yA webs 
NiIcHOL(J.) . 4543 
Noe (Lady A.) +) 18 
NORDENSKIOLD (A. E.) 38 


a 


NorGATE (Kate) 26 ar 
Norris (W. E.) 18 
Norton (Charles Eliot) 3) 37 
Norton (Hon. Mrs.) 15, 1 
OLIPHANT(Mrs. M. O. W?) 


4, II, 13, 19, 20, 39 


OxvipHaANnT (T. L 22, 25 
OLIVER (Prof.D.) . 6 
OLIVER (Capt. S. m, “38 
Oman (C. W.) . tA 
OsTWALD (Prof.) : ot 27 
Orrse (E. C.) , II 
PAGE Ge Ei) 31 


PALGRAVE (Sir F.) 
PALGRAVE (F.T.) 


2,15; iS; 20, 21, ae 39 
PALGRAVE (REBAD?) 29 


PALGRAVE (R. H. Inglis) . 28 
PALGRAVE (W.G.) 15, 29, 38 
PALMER (Lady S.) 5) aK) 
PARKER (T. J.). 6, 39 
PARKER (W. N.) JO. 
ParRKINSON (S.) 2) 27 
PARKMAN (F.) . Beco: 
Parsons (Alfred) aM i 
PasTEuR(L.) . F ee, 
PATER (W. H.) 2 sTO. 22 
PATERSON (J.) . Aa 2 
PATMORE (Coventry) 20, 39 
PAT TESON (J. ©.) rs 
PatTison (Mark) ass 25 
Payne (E. J.) . 10, 29 
PEasopy (C. H.) , 27 
PEEL (E.). ; ris 
PEILE (J.). 25 
PELLISSIER (E. ) 25 
PENNELL (J.) . 4 Taw 
PENNINGTON (R.) BANG 
PENROSE (F.C.) - nA fe 
Perry (Prof. J.) - a 127 
PETTIGREW (J. B.) . 6, 28, 40 
PHILLIMORE (J.G.) . MTS 
PHILLIPS (J. A.) 23 


PuILuips (W. C.) } ANT > 


PicTon (J. A.) . 22 
PIFFARD (H. G.) 23 
PLATO 20 


PLUMPTRE (Dean) 
PoLvarp (A. W.) 

Potiock(SirFk., ond Bart.) a 
Pottock (Sir F. Bart. )12,22,29 


PoLLock (Lady) 3 ee 
PoLLtock (W. Hi.) +e 
Poo.e (M.E.) . . 22 


43 

PAGE 

Poor (Ro 1.) 7; MEL 
Pore. - z 4, 20 
Poste (E. ) 27, 36 
PoTrer (L.) - 22 
PoTTeR (R.) 35 
PRESTON (T. ), : 27 
Price (L. E.F.R. vis 28 
PRICKARD (A. O. JiMe bikes 
PRINCE ALBERT VICTOR. 37 
PRINCE GEORGE 37 
PROCTER (F.) . 32 
ProperT (J. L.) . 2 
RADCLIFFE (C. B.) . 3 
Ramsay (W.) . Stes, 
RANSOME (C.) . P oe rs 
RATHBONE (W. ) 8 
RAWLINSON (W Cty setae 
RAWNSLEY (H. D. Ny 15 
Ay (PKs) es ee 26 
RAYLEIGH (Lord) 27 
REICHEL (Bishop) 35 
Rep) (as. ce a7 
REMSEN (I.)_ . : Saiz 
RENDALL (Rev. F.). 31, 35 
RenpDu(M.leC.) . eS 
Reyno tps (H.R.) . 35 
REYNOLDs (J. R.) 23 
REYNOLDs (O.). ip cr 
RsCHARDSON (B. Ww.) II, 23 


RicuHeEy (A. G.). 12 


Rosrnson (Preb. H. Fenty 35 
RoBInson (J.L.) 24 
RoBINnson (Matthew) 5 
ROCHESTER pence hehe 5 
RocxksTrRo (W.S rea! 
RoceErs (J. E. Ts sits f 20720 
RomaAneEs (G. J.) APG 
Roscoe (Sir H. E. Die AVE 
Roscoe (W. C.) ‘i of PIS 
ROSEBERY (Earl of). 4 
ROSENBUSCH (H.) 9 
Ross (P.) . Ale 
RossETTI (G Gi ) 15, 39 
ROUTLEDGE (J.) 420 
Rowe (F. J.) £26 
RUCKER (Prof. A We ) 7 
RuMFORD (Count) . {ioe 
RUSHBROOKE (W. G.) 31 
RUSSELL (Dean) 35 
RUSSELL (Sir Charles) 29 
RUSSELL (W. vain 4, 19 
RyYtanD (F.) . Sire 
Ry e (Prof. H. ye 30 


St. JOHNSTON (A.) .19, 38, 39 
SADLER (H.) . A 3.°f2 
SaInTsBury (G.) 4, 13 
SaLmon (Rev.G.) . 5 FRG 
SanpForD(M.E.) . YW Es 
Sanpys (J. E.). 38 
Sayce (A.H.) . II 
ScHaFF(P.) . “730 
SCHLIEMANN (Dr. Yea ite 
SCHORLEMMER (C.) . a] 
Scotr (D: Hi) é 3-26 
Scott (Sir W.). I5, 20 
ScRATCHLEY on Peter) 24 
ScupDpER (S. H 40 
SEATON (Dr. Ec We 23 
SEELEY (J.R.). II 
SEILER (Dr. Carl) 23, 28 


SELBORNE (Earl ee 12 720,32533 
SELLERS (E.) 
SERVICE (J.) 


: 32, E : 
SEWELL (E. M.) Beye 3 


INDEX. 


a4 

PAGE 
Suairp (J.C.) . 4, 15 
SHAKESPEARE . 13, 15, 20, 21 
SHANN (G.)_. » 27 
SHarP(W.)  . dS 
SHELLEY . ee ess pens 
SHIRLEY (W. N. ) 435 
SHORTHOUSE (J. H. ) EEO 
SHORTLAND (Admiral) . 24 
SHUCHHARDT (Carl). Me 
SHUCKBURGH (E.S.) 11, 36 
SHUFELDT (R. W.) . <go 
S1eson (Dr. F.) : 5 es 
Sipewick (Prof. H. ) 26, 28, 29 
SIME (J.) . g, 10 
Simpson (Rev. W. 36 in B2 
SKEAT (W. W.) - By is} 
SKRINE (Jn ELE se 55 05 
SLapE (J.H.) . cS 
SLomaN (Rey. AD ae Sh 
Smart (W.) : 3 1238 
SMALLEY (G. W. rae . 22 
SMETHAM (J. andS. ) bol She 
Situ (A.) ‘ 20 
Smitu (C. B.) . 5 LTO 
SMITH (Goldwin) 4) 5) 29 
Situ (H.) 5 5 6 
SMITH (J.) c ie HO 
Situ (Rev. T. ) ; £2135 
SmiTuH (W.G.). 4 “4 10 
SmitTH (W.S.) . 31385 
SOMERVILLE (Prof. W. ste sto 
SoUTHEY . ‘i Ew ETS 
SPENDER (J. K. ) : Des) 
SPENSER . si) 20 
SPOTTISWOODE (Ww. ). 3 27 
STANLEY(Dean)_. 35 
STANLEY (Hon. Maude) . 29 
STATHAM (R.). A 2029 
STEBBING (W.). ; thd 
STEPHEN (C.E.)  . Litho 
STEPHEN (H.) . 13 
STEPHEN (Sir J. F. vA 18395 13, 22 
STEPHEN (J.K.) 13 
STEPHEN (L.) . 0 wt) a 
STEPHENS (J.B.)  . SENEO 
STEVENSON (J. J.) . atk Sto 
STEwartT (A.) . 39 


STEWART (Balfour) "26, 27) 35 
Stewart (S. A.) 


SToxEs (Sir G. G. ys. ~ 27 
Story (R. H.) . ; Bibi, &) 
Stone (W. H.). - 27 
STRACHEY (Sir E. nee 20 20 
STRACHEY (Gen. R.). 9 
STRANGFORD(Viscountess) 38 
STRETTELL (A.) = x6 
StTupss (Rev. C. W.). AAS 
Stupss (Bishop). aot 
SUTHERLAND (A.) . hhc 
Symonps (J. A.) ed: 
Symonps (Mrs. J. A. ) rads 
Symons (A.) 31216 
Tait (Archbishop) . ees 
Tait (C. W. A.) II 


Tait (Prof. P. G.) 26, 27, 35 


: PAGE 
TANNER (H.) . ‘ creer 
TAVERNIER (J. B.) . 38 
TAYLOR (Franklin) . eed: 
TAavyLor (Isaac). 25, 35 
TAYLOR (Sedley) 24, 27 
TEGETMEIER (W. B.) a 
TEMPLE (Bishop) . ess 
TEMPLE (Sir R.) SVAG4 
TENNANT (Dorothy). “8138 
TENNIEL . 38 
TENNYSON ; “34, 16, 21 
TrENNysSON (Frederick) . 16 
TENNYSON (Hallam). —_12, 39 
THompson (D’A.W.) . 6 
THompsON (E.). ‘ 5 ite) 
THompson (S.P.) . Aa ry 
THomson (A. W.)_ . 208 
THomson (Sir C. W.) 140 
TuHomson (Hugh) . I2 
Tuomson (Sir Wm.) 24, 26, 27 
THORNE (Dr. Thorne) . 23 
THORNTON (J.). 6 
THORNTON (W. zy "26, 20, 37 
THORPE (T. E.). 7 
THRING(E.) . : 8, 22 
TuRupp (J. F.). , 4 Se) 
THupicHumM (J.L.W.)-. 7 
THURSFIELD (J. R.). sie 
ToODHUNTER (1.) . 5500. 
Torrens (W.M.) . aC As 
TourRGENIEF (1. S.) . PELO 
Touma GhLsbe) yw . DUEL 
Tozer (H.F.) . : 1 XO 
Trattt (H. D.). : 4, 29 
TRENCH (Capt. F.) . »* 29 
TRENCH (Archbishop) . 35 
TREVELYAN (Sir G.O.) . 11 
TRIBE (A.). A ea O7 
TRISTRAM (W. O. Date Bie 
TROLLOPE (A.). : 3ke 
TRUMAN (J.) 5 eh LO 
Tucker (T. G. ) 36 
TULLOCH (Principal). SA5 
TuRNER (C. Tennyson) . 16 
TURNER (G.) 5 oa EE: 
TuRNER (H. H. ) k - 27 
TuRNER (J. M. W.) . op yt 
TyLor (E. B.) . nn SEI 
TyRwuiTtT (R. St. Je ) 2, 16 


VauGHAN (C. J.) 31, 32, 35, 36 
VAUGHAN (Rev. D. 20, 36 
VAUGHAN (Rev. E.T.) . 36 
VAUGHAN (ey. Rie 536 
VELEy (M.) A uO 
VENN (Rev. J. ‘ 26, 36 
VERNON (Hon. W. W.) bP hate 


VERRALL (A. W.) 13, 36 
VERRALL (Mrs.) 5 war 
Wain (Louis) . 5 Bo 4 2I0) 
WALDSTEIN (C.) sarye 


WALKER (Prof. F. A.) . 28 
Wa.tace (A. R.) 6, 24, 28 
Wa tvace (Sir D. M.) . 29 
WaLpoLe(S.) . ; p20 
Watton (I.) . ° <n re 


PAGE 
Warp (A. W.) . J (AS E3,)'0 
Warp (H. M. is : 
Warp (5.). 5 LEG 
Warp (T. H. Dd: by iG 
Warp (Mrs. T JL ¥:6 IQ, 39 
Warp (W.) 3 5, 32 
W ARINGTON (G. vias Lago 
Waters (C. A.) 28 
WATERTON (Charles) 245 38 
Watson (E.) . 5 5 
Watson (R. S.) 4 2.138 
Wess (W.T.) . 26 


WEBSTER (Mrs. A.) . 39 
WELBY-GREGORY (Lady) . 32 
WELLDON (Rev. J. E.C.). 36 
Westcott (Bp.) 30, 31, 3% 36 
WESTERMARCK (E.). 


WETHERELL(J.) . ; Be 
WHEELER(J.T.)  . Be ie 
WHEWELL (W.). ¢. es 
Wuite (Gilbert) 3 . 24 
WHITE (Dr. W. Hale) . 23 
WHITE (W.) . : A Boe, 
WuitHaM(J.M.) . Re 3G) 
WHITNEY (W.D.) . 4 tae 


WHITTIER(J.G.) . 
Wickuam (Rev. E.€.) . 36 
WICKSTEED (P. H.). 
WIEDERSHEIM (R.) . ; pe 
WILBRAHAM (F. M. 5 

WILKINS (Prof. A.S.) 2, 13, 36 


WILKINSON (S.) : A. G24 
Wituiams (G. H.) . BA 0) 
Wituiams (Montagu) . 5 
Witiiams(S.E.)  . 4) hee 
WILLouGHBY(F.) . - 930 
WILLs (W.G.) . : Ae ale, 
Witson (A. J.). ; . 29 
WItson (Sir C.) : Aiea 


Witson (Sir D.) / ii feyee 


WItson (Dr. G.) ay See 
Witson (Archdeacon) . 36 
WILson (Mary). F 13 
WINGATE (Major F. R. ia 24 
WINKWORTH (C.)_ . 5 
WoOLSELEY (Gen. Viscount) 24 
Woop (A.G.) . “i ibe 
Woop (Rev. E.G.) . hao 
Woops (Rev. F. H.). Ripe ilee 
Woops (MissM.A.). 17, 33 
Woopwarp (C. M.). was 
WOooLNER (7T.) . : . 16 
WORDSWORTH ._ 5, 14, 16, 21 
WorTHEY(Mrs.) . Ape fe) 
WriGHT (Rey. A.) . Men 
WriGcHT(C. E.G.) . ot his 
WRIGHT (J.) . 0 2 
Wriacut (L.) 27 
WRIGHT (W. Aldis) 8, 15, 20) 31 
Wortz (Ad. 7 
Wyatt (Sir M. 'D. Wi. 2 


YoncE (C. M.) 5, 6, 8, 10, 11, 

IQ, 21, 25; 39, 39 
Younce (E. W.) : Pp 2) 
ZIEGLER) (Orbs) eens ee 


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