M 11

FACTS AND ARGUMENTS

DARWIN.

BY FRITZ MÜLLER,

WITH ADDITIONS BY THE AUTHOR.

TRANSLATED FROM THE GERMAN

By W. S. DALLAS, F.L.S.,

ASSISTANT SECKETARY TO THE GEOLOGICAL SOCIETY OF LONDON.

WITH ILLUSTRATIONS.

LONDON:
JOHN MURRAY, ALBEMARLE STREET

1869.

MR. DARWIN'S WORKS.

A NATURALIST'S VOYAGE ROUND THE WORLD ;

being a Jouknal of Rkseakchks into the Natural History and Geology of
CouKTßiES Visited. Post 8vo. 9s.

THE ORIGIN OF SPECIES, by MEANS of NATURAL

SELECTION ; or, The Preservation of Favoured Eaces in the Struggle
FOR Life. Woodcuts. Post 8vo. 15s.

THE VARIOUS CONTRIVANCES by which BRITISH

and FOREIGN ORCHIDS are FERTILIZED by INSECTS, and on th? GOOD
EFFECTS of INTERCROSSING. Woodcuts. Post 8vo. 9s.

THE VARIATION OF ANIMALS AND PLANTS

UNDER DOMESTICATION. Illustrations. 2 vols., 8vo. 28s.

LONDOX: printed by W. CLOWES AND SONS, DUKE STREET, STAMFORD STRKET,
AND CHARING CROSS

TRANSLATORS PREFA

My principal reason for undertaking tlie translation
of Dr. Fritz Miiller's admirable work on the Crustacea,
entitled 'Für Darwin,' was that it was still, although
published as long ago as 1864, and highly esteemed
by the author's scientific countrymen, absolutely
unknown to a great number of English naturalists,
including some who have occupied themselves more
or less specially with the subjects of which it treats.
It possesses a value quite independent of its reference
to Darwinism, due to the number of highly interesting
and important facts in the natural history and par-
ticularly the developmental history of the Crustacea,
which its distinguished author, himself an unwearied
and original investigator of these matters, has brought
together in it. To a considerable section of English
naturalists the tone adopted by the author in speaking
of one of the greatest of their number will be a source
of much gratification.

In granting his permission for the translation of his
little book. Dr. Fritz Müller kindly offered to send
some emendations and additions to certain parts of it.
His notes included many corrections of printers' errors,
some of which would have proved unintelligible with-
out his aid, some small additions and notes which

a 2

( iv )

have been inserted in their proper places, and two
longer pieces, one forming a foot-note near the close of
Cliap. XI. (p. 119), the other at the end of Chap. XII.
(pp. 135-140), describing the probable mode of evolu-
tion of the Khizocephala from the Cirripedia.

Of the execution of the translation I will say but
little. My chief object in this, as in other cases, has
been to furnish, as nearly as possible, a literal version
of the original, regardiog mere elegance of expression
as of secondary importance in a scientific work. As
much of Dr. Müller's German does not submit itself to
such treatment very readily, I must beg his and the
reader's indulgence for any imperfections arising from
this cause.

W. S. D.

London, 15^/i Feb., 18G9.

AUTHOR'S PREFACE.

It is not the purpose of the following pages to discuss
once more the arguments adduced for and against
Darwin's theory of the origin of species, or to weigh
them one against the other. Their object is simply to
indicate a few facts favourable to this theory, collected
upon the same South American ground, on which, as
Darwin tells us, the idea first occurred to him of de-
voting his attention to "the origin of species, — that
mystery of mysteries."

It is only by the accumulation of new and valuable
material that the controversy will gradually be brought
into a state fit for final decision, and this appears to be
for the present of more importance than a repeated
analysis of what is ah'eady before us. Moreover, it is
but fair to leave it to Darwin himself at first to beat oft*
the attacks of his opponents from the splendid structure
which he has raised with such a master-hand.

F. M.

Desterro, 1th Sej^t., 1863.

CONTENTS.

Chap. tagk

I. Introdixtory 1

II. The Species of Meltta 7

III. Morphology OF Crustacea 13

lY. Sexual Peculiarities AND Dimorphism 19

V. Eespiratiox in Land Crabs 30

VI. Structure of the Heart is Edriophthalma 39

VII. Developmental History of Podophthalma 47

VIII. Developmental Hi:sTORY OF Edriophthalma 69

IX. Developmental History of Entomostraca, Ciruipedes,

AND Ehizocephala 83

X. On the Principles of Classification 97

XL On THE Progre.'>s OF Evolution 110

XII. PRoGRE.'is of Evolution IN Cru.'jtacea 122

29120

HISTORY OF CRUSTACEA

CHAPTER I.

INTRODUCTORY.

When I had read Charles Darwin's book *0n tiiet-^-
Origin of Species,' it seemed to me that there was one
mode, and that perhaps the most certain, of testing
the correctness of the views developed in it, namely,
to attempt to apply them as specially as possible
to some particular group of animals. Such an
attempt to establish a genealogical tree, whether for
the families of a class, the genera of a large family,
or for the species of an extensive genus, and to pro-
duce pictures as complete and intelligible as possible
of the common ancestors of the various smaller and
larger circles, might furnish a result in three different
ways.

1. In the first place, Darwin's suppositions when thus
applied might lead to irreconcilable ^nd contradictory
conclusions, from which the erroneousness of the sup-
positions might be inferred. If Darwin's opinions are
false, it was to be expected that contradictions would
accompany their detailed application at every step, and

B

^ HISTOKY OF CRUSTACEA. Chap. I.

that these, by their cumulative force, would entirely
destroy the suppositions from which they proceeded,
even though the deductions derived from each particular
case might possess little of the unconditional nature of
mathematical proof.

2. Secondly, the attempt might be successful to a
greater or less extent. If it was possible upon the
foundation and with the aid of the Darwinian theory,
to show in what sequence the various smaller and larger
circles had separated from the common fundamental
form and. from each other, in what sequence they had
acquired the peculiarities which now characterise them,
and what transformations they had undergone in the
lapse of ages, — if the establishment of such a genea-
logical tree, of a primitive history of the group under
consideration, free from internal contradictions, was
possible, — then this conception, the more completely
it took up all the species within itself, and the more
deeply it enabled us to descend into the details of their
structure, must in the same proportion bear in itself
the warrant of its truth, and the more convincingly
prove that the foundation upon which it is built is no
loose sand, and that it is more than merely "■ an intel-
lectual dream."

3. In the third place, however, it was possible, and
this could not but appear, jprimd facie, the most pro-
bable case, that the attempt might be frustrated by
the difficulties standing in its way, without settling the
question, either way, in a perfectly satisfactory manner.
But if it were only possible in this way to arrive for

Chap. I. INTRODUCTORY. 3'

oneself at a moderately certain independent judg-
ment upon a matter affecting the highest questions
so deej)ly, even this alone could not but be esteemed
a great gain.

Having determined to make the attempt, I had in
the first place to decide upon some particular class.
The choice was necessarily limited to those the chief
forms of which were easily to be obtained alive in some
abundance. The Crabs and Macrurous Crustacea, the
Stomapoda, the Diastylidse, the Amphipoda and Iso-
poda, the Ostracoda and Daphnidae, the Copepoda and
Parasita, the Cirripedes and Khizocephala of our coast,
representing the class of Crustacea with the deficiency
only of the Phyllopoda and Xiphosura, furnished a long
and varied, and at the same time intimately connected
series, such as was at my command in no other class.
But even independently of this circumstance the selec-
tion of the Crustacea could hardly have been doubtful.
Nowhere else, as has already been indicated by various
writers, is the temptation stronger to give to the expres-
sions " relationship, production from a common funda-
mental form," and the like, more than a mere figurative
signification, than in the case of the lower Crustacea.
Among the parasitic Crustacea, especially, everybody
has long been accustomed to speak, in a manner scarcely
admitting of a figurative meaning, of their arrest of
development by parasitism, as if the transformation of
species were a matter of course. It would certainly
never appear to any one to be a pastime worthy of the
Deity, to amuse himself with the contrivance of these

B 2

4 HISTOEY OF CKUSTACEA. Chap. I.

marvellous cripplings, and so they were supposed to
have fallen by their own fault, like Adam, from their
previous state of perfection.

That a great part of the larger and smaller groups
into which this class is divided, might be regarded as
satisfactorily established, was a further advantage not
to be undervalued; whilst in two other classes with
which I was familiar, namely, the Annelida and Aca-
lephse, all the attempted arrangements could only be
considered preliminary revisions. These undisplace-
able groups, like the sharply marked forms of the hard,
many-jointed dermal framework, were not only import-
ant as safe starting points and supports, but were also
of the highest value as inflexible barriers in a problem
in which, from its very nature, fancy must freely unfold
her wings.

When I thus began to study our Crustacea more
closely from this new stand-point of the Darwinian
theory, — when I attempted to bring their arrangements
into the form of a genealogical tree, and to form some
idea of the probable structure of their ancestors, — I
speedily saw (as indeed I expected) that it would require
years of preliminary work before the essential problem
could be seriously handled. The extant systematic
works generally laid more weight upon the characters
separating the genera, families and orders, than upon
those which unite the members of each group, and con-
sequently often furnished but httle employable material.
But above all things a thorough knowledge of develop-
ment was indispensable, and every one knows how im-

Chap. I. INTRODUCTORY. 5

perfect is our present knowledge of this subject. The
existing deficiencies were the more difficult to supply,
because, as Van Beneden remarks with regard to the
Decapoda, from the often incredible difference in the
development of the most nearly allied forms, these
must be separately studied — usually family by family,
and frequently genus by genus — ^nay, sometimes, as in the
case oiFeneus, even species by species ; and because these
investigations, in themselves troublesome and tedious,
often depend for their success upon a lucky chance.

But although the satisfactory completion of the
" Genealogical tree of the Crustacea " appeared to be
an undertaking for which the strength and life of an in-
dividual would hardly suffice, even under more favour-
able circumstances than could be presented by a distant
island, far removed from the great market of scientific
life, far from libraries and museums — nevertheless its
practicability became daily less doubtful in my eyes, and
fresh observations daily made me more favourably in-
clined towards the Darwinian theory.

In determining to state the arguments which I de-
rived from the consideration of our Crustacea in favour
of Darwin's views, and which (together with more general
considerations and observations in other departments),
essentially aided in making the correctness of those views
seem more and more palpable to me, I am chiefly influ-
enced by an expression of Darwin's : " Whoever," says
he (' Origin of Species,' p. 482), " is led to believe
that species are mutable, will do a good service
by conscientiously expressing his conviction." To the

0 HISTOEY OF CEUSTACEA. Chap. I.

desire expressed in these words I respond, for my own
part, with the more pleasure, as this furnishes me with
an opportunity of publicly giving expression in words to
the thanks which I feel most deeply to be due from me
to Darwin for the instructions and suggestions for which

1 am so deeply indebted to his book. Accordingly I
throw this sand-grain with confidence into the scale
against " the load of prejudice by which this subject is
overwhelmed," without troubling myself as to whether
the priests of orthodox science will reckon me amongst
dreamers and children in knowledge of the laws of
nature.

Chap. II. THE SPECIES OF MELITA.

CHAPTER IL

THE SPECIES OF MELITA.

A FALSE supposition, when tlie consequences proceeding
from it are followed further and further, will sooner or
later lead to absurdities and palpable contradictions.
During the period of tormenting doubt — and this was
by no means a short one — when the pointer of the
scales oscillated before me in perfect uncertainty be-
tween the p'o and the con, and when any fact leading
to a quick decision would have been most welcome
to me, I took no small pains to detect some such con-
tradictions among the inferences as to the class of
Crustacea furnished by the Darwinian theory. But I
found none, either then, or subsequently. Those which
I thought I had found were dispelled on closer con-
sideration, or actually became converted into supports
for Darwin's theory.

Nor, so far as I am aware, have any of the necessary
consequences of Darwin's hypotheses been proved by any
one else, to stand in clear and irreconcilable contradic-
tion. And yet, as the most profound students of the
animal kingdom are amongst Darwin's opponents, it
would seem that it ought to have been an easy matter
for them to crush him long since beneath a mass of ab-

8 HISTORY OF CRUSTACEA. Chap. 11.

surd and contradictory inferences, if any such were to be
drawn from his theory. To this want of demonstrated
contradictions I think we may ascribe just the same im-
portance in Darwin's favour, that his opponents have
attributed to the absence of demonstrated intermediate
forms between the species of the various strata of the
earth. Independently of the reasons which Darwin
gives for the preservation of such intermediate forms
being only exceptional, this last mentioned circumstance
will not be regarded as of very great significance by any
one who has traced the development of an animal upon
larvae fished from the sea, and had to seek in vain for
months, and even years, for those transitional forms,
which he nevertheless knew to be swarming around him
in thousands.

A few examples may show how contradictions might
come forth as necessary results of the Darwinian hypo-
theses.

It seems to be a necessity for all crabs which remain
for a long time out of the water (but why is of no con-
sequence to us here), that air shall penetrate from behind
into the branchial cavity. Now these crabs, which have
become more or less estranged from the water, belong
to the most different families — the Kaninidae (Banina),
Eriphinse (Eri^Ma gonagra), GrapsoidsB (Aratus, Ses-
arma, &c.), Ocypodidae (Gelasimus, Ocypoda), &c., and the
separation of these families must doubtless be referred
to a much earlier period than the habit of leaving the
water displayed by some of their members. The arrange-
ments connected with aerial respiration, therefore, could

Chap. II.

THE SPECIES OF MELITA.

9

not be inherited from a common ancestor, and could
scarcely be accordant in their construction. If there were
any such accordance not referable to accidental resem-
blance among them, it would have to be laid in the scale as
evidence against the correctness of Darwin's views. I
shall show hereafter how in this case the result, far from
presenting such contradictions, was rather in the most
complete harmony with what might be predicted from
Darwin^s theory.

A second example. — We are already acquainted with
four species of Melita {M. valida, settles, anisocliir, and

Fig. 1

Fresnelii\ and I can add a fifth (fig. 1), in which the
second pair of feet bears upon one side a small hand of
the usual structure, and on the other an enormous
clasp-forceps. This want of symmetry is something so

^ Fig. 1. Melita exilli n. sp., male, enlarged five times. The large
branchial lamellae are seen projecting between the legs.

10 HISTOKT OF CRUSTACEA. Chap. II.

unusual among tlie Ampliipoda, and the structure of
the clasp-forceps differs so much from what is seen else-
where in this order, and agrees so closely in the five
species, that one must unhesitatingly regard them as
having sprung from common ancestors belonging to
them alone among known species. But one of these
species, M. Fresnelii, discovered by Savigny, in Egypt,
is said to want the secondary flagellum of the anterior
antennse, which occurs in the others. From the trust-
worthiness of all Savigny's works there can scarcely be
a doubt as to the correctness of this statement. Now,
if the presence or absence of the secondary flagellum
possessed the significance of a distinctive generic cha-
racter, which is usually ascribed to it, or if there were
other important differences between Melita Fresnelii
and the other species above-mentioned, which would
make it seem natural to separate M. Fresnelii as a dis-
tinct genus, and to leave the others united with the
rest of the species of Melita — that is to say, in the
sense of the Darwinian theory, if we assume that all
the other Melitm possessed common ancestors, which
were not at the same time the ancestors of M. Fres-
nelii— this would stand in contradiction to the conclu-
sion, derived from the structure of the clasp-forceps,
that M. Fresnelii and the four other species above-men-
tioned possessed common ancestors, which were not also
the ancestors of the remaining species of Melita. It
would follow

Chap. II. THE SPECIES OF MELITA.

From the structure of the clasp-forcpps :

11

From the presence or absence of the
secoiida'v flncr^Uum.

M. palmata, kc. M.ezilii, &c. M. Fresndli. M. palinata, &c. M. exilii, kc. M. Fresnelii

As, in the first case, among the Crabs, a typical agree-
ment of arrangements produced independently of each
other would have been a very suspicious circumstance
for Darwin's theory, so also, in the second, would any
difference more profound tlian that of very nearly
allied species. Now it seems to me that the secondary
flagellum can by no means furnish a reason for doubting
the close relationship of M. Fresnelii to M. exilii, &c.,
which is indicated by the peculiar structure of the un-
paired clasp-forceps. In the first place we must con-
sider the possibility that the secondary flagellum, which
is not always easy to detect, may only have been over-
looked by Savigny, as indeed Spence Bate supposes to
have been the case. If it is really deficient it must be
remarked that I have found it in species of the ge-
nera Leucothoe, Cyrtojplnum and Amjohilochus, in which
genera it was missed by Savigny, Dana and Spence
Bate — that a species proved by the form of the epimera
(coxse Sp..B.) of the caudal feet (urojooda Westw.), &c.,
to be a true Amjphithoe^ possesses it — that in many
species of Cerapus it is reduced to a scarcely perceptible

2 I accept this and all the other genera of Amphipoda here men
tinned, with the hmits given to them by Spence Bate (^' Catal. of
podous Crustacea '). y<\ V

z:^,

12 HISTOKY OF CEUSTACEA. Chap. II.

rudiment — nay, that it is sometimes present in youth
and disappears (although perhaps not without leaving
some trace) at maturity, as was found by Spence Bate
to be the case in Acanthonotus Owenii and Atylus eari-
natus, and I can affirm with regard to an Atylus of these
seas, remarkable for its plumose branchiae — and that
from all this, at the present day when the increasing
number of known Amphipoda and the splitting of them
into numerous genera thereby induced, compels us to
descend to very minute distinctive characters, we must
nevertheless hesitate before employing the secondary
flagellum as a generic character. The case of Melita
Fresnelii therefore cannot excite any doubts as to
Darwin's theory.

Chap. III. MORPHOLOGY OF CRUSTACEA. 13

CHAPTEK III.

MORPHOLOGY OF CRUSTACEA — NAUPLIUS-LARV^.

If the absence of contradictions among the inferences
deduced from them for a narrow and consequently
easily surveyed department must prepossess us in
favour of Darwin's views, it must be welcomed as a
positive triumph of his theory if far-reaching conclu-
sions founded upon it should subsequently be confirmed
by facts, the existence of which science, in its previous
state, by no means allowed us to suspect. From many
results of this kind upon which I could report, I select
as examples, two, which were of particular importance
to me, and relate to discoveries the great significance of
which in the morphology and classification of the Crus-
tacea will not be denied even by the opponents of
Darwin.

Considerations upon the developmental history of
the Crustacea had led me to the conclusion that, if the
higher and lower Crustacea were at ^all derivable from
common progenitors, the former also must once have
passed through Nauplius-like conditions. Soon after-
wards I discovered Naupliiform larvae of Shrimps (* Ar-
chiv für Naturg.' 1860, i. p. 8), and I must admit that

14 HISTOEY OF CRUSTACEA. Chap. III.

this discovery gave me ttie first decided turn in Dar-
win's favour.

The similar number of segments ^ occurring in the

1 Like Claus I do not regard tlie eyes of the Crustacea as limbs, and
therefore admit no ocular segment ; on the other hand I count in the
median piece of the tail, to which the character of a segment is often
denied. In opposition to its interpretation as a segment of the body,
only the want of limbs can be cited ; in its favour we have the relation
of the intestine, which usually opens in this piece, and sometimes even
traverses its whole length, as in Microdeutopus and some other Amphi-
poda. In Microdeutopus, as Spence Bate has already pointed out, one
is even led to regard small processes of this tubular caudal piece as
rudimentary members. Bell also (' Brit. Stalk-eyed Crust.' p. xx.),
states that he observed limbs of the last segment in Falxmon serratus
in the form of small movable points.

The attempt has often been made to divide the body of the higher
Crustacea into small sections composed of equal numbers of segments,
these sections consisting of 3, 5 or 7 segments. None of these attempts
has ever met with general acceptance ; my own investigations lead me to
a conception which nearly approaches Van Beneden's. I assume four
sections of 5 segments each — the primitive body, the fore-body, the hind-
body, and the middle-body. The primitive body includes the segments
which the naupliiform larva brings with it out of the egg ; it is after-
wards divided, by the younger sections which become developed in its
middle, into the head and tail. To this primitive body belong the two
pairs of antennae, the mandibles and the caudal feet (" posterior pair of
pleopoda," Sp. B.). Even in the mature animal the fact that these
terminal sections belong to one another is sometimes betrayed by the
resemblance of their appendages, especially that of the outer branch of
the caudal feet, with the outer branch (the so-called scale) of the second
pair of antennaB. Like the antenna?, the caudal feet may also become
the bearers of high sensorial apparatus, as is shown by the ear of
My sis.

The sequence of the sections of the body in order of time seems
originally to have been, that first the fore-body, then the hind-body, and
finally the middle-body was formed. The fore-body appears, in the
adult animal, to be entirely or partially amalgamated with the head ;
its appendages (siagonopoda Westw.) are all or in part serviceable for
the reception of food, and generally sharply distinguished from those of
the following group. The segments of the middle-body seem always

Chap. III. MOEPHOLOGY OF CRUSTACEA. 15

Crabs and Macrura, Ampliipoda and Isopoda, in which
the last seven segments are always different from the
preceding ones in the appendages with which they are
furnished, could only be regarded as an inheritance from
the same ancestors. And if at the present day the
majority of the Crabs and Macrura, and indeed the
Stalk-eyed Crustacea in general, pass through Zoea-like
developmental states, and the same mode of transforma-
tion was to be ascribed to their ancestors, the same
thing must also apply, if not to the immediate ancestors
of the Amphipoda and Isopoda, at least to the common
progenitors of these and the Stalk-eyed Crustacea. Any
such assumption as this was, however, very hazardous,
so long as not a single fact properly relating to the

to put forth limbs immediately after their own appearance, whilst the
segments of the hind-body often remain destitute of feet through long
portions of the larval life or even throughout life (as in many female
Diastylidse), a reason, among many others, for not, as is usual, regard-
ing the middle-body of the Crustacea as equivalent to the constantly
footless abdomen of Insects. The appendages of the middle-body
(pereiopoda) seem never, even in their youngest form, to possess two
equal branches, a peculiarity which usually characterises the appendages
of the hind-body. This is a circumstance which renders very doubtful
the equivalence of the middle-body of the Malacostraca with the section
of the body which in the Copepoda bears the swimming feet and in the
Cirripedia the cirri.

The comprehension of the feet of the hind-body and tail in a single
group (as "fausses pattes abdominales," or as " pleopoda ") seems not
to be justifiable. When there is a metamorphosis, they are probably
always produced at diiferent periods, and they are almost always quite
different in structure and function. Even in the Amphipoda, in which
the caudal feet usually resemble in appearance the last two pairs
of abdominal feet, they are in general distinguished by some sort of
peculiarity, and whilst the abdominal feet are reproduced in wearisome
uniformity throughout the entire order, the caudal feet are, as is well
known, amongst the most variable parts of the Amphipoda.

16 HISTORY OF CRUSTACEA. Chap. III.

Edrioplitlialma could be adduced in its support, as the
structure of this very coherent group seemed to be
almost irreconcilable with many peculiarities of the
ZoUa. Thus, in my eyes, this point long constituted one
of the chief difficulties in the application of the Dar-
winian views to the Crustacea, and I could scarcely
venture to hope that I might yet find traces of this
passage through the Zoea-form among the Amphipoda
or Isopoda, and thus obtain a positive proof of the cor-
rectness of this conclusion. At this point Van Bene-
den's statement that a cheliferous Isopod {Tanais
Dulongii), belonging, according to Milne-Edwards,
to the same family as the common Äsellus aqua-
tieus, possesses a carapace like the Decapoda, directed
my attention to these animals, and a careful exa-

Fig. 2.2

mination proved that these Isopods have preserved,
more truly than any other adult Crustacea, many of the
most essential peculiarities of the Zoece, especially their

- Tanais duhius (?) Kr, $ , magnified 25 times, showing the orifice
of entrance (x) into the cavity overarched by the carapace, in which
an appendage of the second pair of maxillae (/) plays. On four feet
(i, k, I, m) are the rudiments of the lamellae which subsequently form
the brood-cavity.

Chap. III. NAUPLIUS-FORMS. 17

mode of respiration. Whilst in all other Oniscoida the
abdominal feet serve for respiration, these in our cheli-
ferous Isopod (fig. 2) are solely motory organs, into
which no blood-corpuscle ever enters, and the chief
seat of respiration is, as in the Zoese, in the lateral
parts of the carapace, which are abundantly traversed
by currents of blood, and beneath which a constant
stream of water passes, maintained, as in ZoiSm and
the adult Decapoda, by an appendage of the second
pair of maxilloB, which is wanting in all other Edrioph-
thalma.

For both these discoveries, it may be remarked in
passing, science is indebted less to a happy chance than
immediately to Darwin's theory.

Species of Feneus live in the European seas, as well as
here, and their NaupUus-hrood has no doubt repeatedly
passed unnoticed through the hands of the numerous
naturalists who have investigated those seas, as well as
through my o\^n,^ for it has" nothing which could attract
particular attention amongst the multifarious and often
wonderful NaupUus-iorms. When I, fancying from the
similarity of its movements that it was a young Peiieus-
Zoea, had for the first time captured such a larva, and
on bringing it under the microscope found a Nauplius
differing toto coelo from this Zoea, I might have thrown
it aside as being completely foreign to the develop-
mental series which I was tracing, if the idea of early
Naupliiform stages of the higher Crustacea, which in-

3 Mecznikow has recently found Naupliiform shrimp-larvss in the sea
near Naples.

C

18 HISTORY OF CRUSTACEA. Chap. HI.

deed I did not believe to be still extant, had not at the
moment vividly occupied my attention.

And if I had not long been seeking among the
Edriophthalma for traces of the supposititious Zoea-
state, and seized with avidity upon everything that
promised to make this refractory Order serviceable to
me, Yan Beneden's short statement could hardly have
affected me so much in the manner of an electric
shock, and impelled me to a renewed study of the
Tanaides, especially as I had once before plagued
myself with them in the Baltic, without getting any
further than my predecessors, and I have not much
taste for going twice over the same ground.

Chap. IV. TANAIS DüBIUS. 19

CHAPTEK IV.

SEXUAL PECULIARITIES AND DIMORPHISM.

Our Tanais, which in nearly all the particulars of its
structure is an extremely remarkable animal, furnished
me with a second fact worthy of notice in connection
with the theory of the origin of species by natural
selection.

When handlike or cheliform structures occur in the
Crustacea, these are usually more strongly developed in
the males than in the females, often becoming enlarged
in the former to quite a disproportionate size, as we
have already seen to be the case in Melita. A better
known example of such gigantic chelae is presented
by the males of the Calling Crabs (Gelasimus), which
are said in running to carry these claws "elevated,
as if beckoning with them " — a statement which, how-
ever, is not true of all the species, as a small and
particularly large-clawed one, which I have seen run-
ning about by thousands in the cassava-fields at the
mouth of the Cambriu, always holds them closely
pressed against its body.

A second peculiarity of the male Crustacea consists
not unfrequently in a more abundant development on

c 2

20 HISTOEY OF CRUSTACEA. Chap. IY.

the flagellum of the anterior antennse of delicate fila-
ments which Spence Bate calls "auditory cilia," and
which I have considered to be olfactory organs, as
did Leydig before me, although I was not aware of
it. Thus they form long dense tufts in the males of
many Diastylidse, as Van Beneden also states with regard
to Bodotria, whilst the females only possess them more
sparingly. In the Copepoda, Claus called attention to
the difference of the sexes in this respect. It seems to
me, as I may remark in passing, that this stronger deve-
lopment in the males is greatly in favour of the opinion
maintained by Leydig and myself, as in other cases male
animals are not unfrequently guided by the scent in
their pursuit of the ardent females.

Now, in our Tanais, the young males up to tlie last
change of skin preceding sexual maturity resemble the
females, but then they undergo an important metamor-
phosis. Amongst other things they lose the moveable
appendages of the mouth even to those which serve for
the maintenance of the respiratory current ; their in-
testine is always found empty, and they appear only to live
for love. But what is most remarkable is, that they now
appear under two different forms. Some (fig. 3) acquire
powerful, long-fingered, and very mobile chelae, and,
instead of the single olfactory filament of the female,
have from 12 to 17 of these organs, which stand two or
three together on each joint of the flagellum. The others
(fig. 5) retain the short thick form of the chelae of the
females ; but, on the other hand, their antennse (fig. 6)
are equipped with a far greater number of olfactory

Chap. IV.

TANAIS DUBIUS.

21

filaments, wliich stand in groups of from five to seven

together.

Figs. 3-6. 1

* Fig. 3. Head of the ordinary form of the male of Tanais duhius (?)
Kr. magn. 90 times. The terminal setse of the second pair of antennas
project between the cheliferous feet. Fig. 4. Buccal region of the same
from below ; A, labrum. Fig. 5. Head of the rarer form of the male,
magn. 25 times. Fig. 6. Flagellum of the same, with olfactory fila-
ments, magn. 90 times.

22 HISTORY OF CRUSTACEA. Chap. IV.

In the first place, and before inquiring into its signi-
ficance, I will say a word upon this fact itself. It was
natural to consider whether two different species with
very similar females and very different males might not
perhaps live together, or whether the males, instead of
occurring in two sharply defined forms, might not be
only variable within very wide limits. I can admit
neither of these suppositions. Our Tanais lives among
densely interwoven Confervse, which form a coat of
about an inch in thickness upon stones in the neighbour-
hood of the shore. If a handful of this green felt is put
into a large glass with clear sea-water, the walls of the
glass are soon seen covered with hundreds, nay with
thousands, of these little, plump, whitish Isopods. In
this way I have examined thousands of them with the
simple lens, and I have also examined many hundreds
with the microscope, without finding any differences
among the females, or any intermediate forms between
the two kinds of males.

To the old school this occurrence of two kinds of
males will appear to be merely a matter of curiosity.
To those who regard the "plan of creation" as the
"free conception of an Almighty intellect, matured
in the thoughts of the latter before it is manifested in
palpable, external forms," it will appear to be a mere
caprice of the Creator, as it is inexplicable either
from the point of view of practical adaptation, or
from the "typical plan of structure." From the side
of Darwin's theory, on the contrary, this fact acquires
meaning and significance, and it appears in return

Chap. IV. TANAIS DUBIUS. 23

to be fitted to throw light upon a question in which
Bronn saw " the first and most material objection
against the new theory," namely, how it is possible
that from the accumulation in various directions of the
smallest variations running out of one another, varieties
and species are produced, which stand out from the pri-
mary form clearly and sharply like the petiolated leaf
of a Dicotyledon, and are not amalgamated with the
primary form and with each other like the irregular
curled lobes of a foliaceous Lichen.

Let us suppose that the males of our Tanais, hitherto
identical in structure, begin to vary, in all directions as
Bronn thinks, for auglit I care. If the species was
adapted to its conditions of existence, if the best in this
respect had been attained and secured by natural selec-
tion, fresh variations affecting the species as a species
would be retrogressions, and thus could have no prospect
of prevailing. They must rather have disappeared
again as they arose, and the lists would remain open to
the males under variation, only in respect of their sexual
relations. In these they might acquire advantages over
their rivals by their being enabled either to seek or to
seize the females better. The best smellers would over-
come all that were inferior to them in this respect, unless
the latter had other advantages, such as more powerful
chelae, to oppose to them. The best ^claspers would over-
come all less strongly armed champions, unless these
opposed to them some other advantage, such as sharper
senses. It will be easily understood how in this manner all
the intermediate steps less favoured in the development

24 HISTORY OF CRUSTACEA. Chap. IV.

of tlie olfactory filaments or of the chelae would disappear
from the lists, and two sharply defined forms, the best
smellers and the best claspers, would remain as the sole
adversaries. At the present day the contest seems to
have been decided in favour of the latter, as they occur
in greatly preponderating numbers, perhaps a hundred
of them to one smeller.

To return to Bronn's objection. When he says that
" for the support of the Darwinian theory, and in order
to explain why many species do not coalesce by means
of intermediate forms, he would gladly discover some
external or internal principle which should compel the
variations of each species to advance in one direction,
instead of merely permitting them in all directions," we
may, in this as in many other cases, find such a principle
in the fact that actually only a few directions stand open
in which the variations are at the same time improve-
ments, and in which therefore they can accumulate and
become fixed ; whilst in all others, being either indifferent
or injurious, they will go as lightly as they come.

The occurrence of two kinds of males in the same
species may perhaps not be a very rare phenomenon
in animals in which the males differ wddely from
the females in structure. But only in those which
can be procured in sufficient abundance, will it be
possible to arrive at a conviction that we have not
before us either two different species, or animals of
different ages. From my own observation, although
not very extensive, I can give a second example. It
relates to a shore-hopper (Orchestia). The animal

Chap. 1Y.

OECHESTIA DARWmil.

25

(fig. 7) lives in marshy places in the vicinity of the sea,
under decaying leaves, in the loose earth which the
Marsh Crabs (Gelasimus, Sesarma, Cyelograpsus, &c.)

throw up around the entrance to their burrows, and
even under dry cow-dung and horse-dung. If this
species removes to a greater distance from the shore
than the majority of its congeners (although some of
them advance very far into the land and even upon
mountains of a thousand feet in height, such as 0.
tahitensis, telluris, and syhieola), its male differs still
more from all known species by the powerful chelae of
the second pair of feet. Orchestia grtjiohus, from the
sandy coast of Mönchgut, alone presents a somewhat
similar structure, but in a far less degree ; elsewhere
the form of the hand usual in the Amphipoda occurs.
Now there is a considerable difference between the
males of this species, especially in the structure of these
chelge — a difference so great that we can scarcely find a
parallel to it elsewhere between two species of the
genus— and yet, as in Tanais, we do not meet with a

- Fig. 7. Orchestia Darwinii, n. sp. male.

26

HISTORY OF CRUSTACEA.

Chap. IY.

long series of structures running into one another, but
only two forms united by no intermediate terms (figs. 8
and 9). The males would be unhesitatingly regarded

Figs. 8 and

as belonging to two well-marked species if they did not
live on the same spot, wdth undistinguishable females.
That the two forms of the chelae of the males occur
in this species is so far worthy of notice, because the
formation of the chelae^ which differs widely from the
ordinary structure in the other species, indicates that
it has quite recently undergone considerable changes,
and therefore such a phenomenon w^as to be expected
in it rather than in other species.

•^ Figs. 8 and 9. The two forms of the chelee of the male of Orchestia
Darwinii, magn. 45 times.

Chap. IY.

MELITA MESSALINA.

27

I cannot refrain from taking this opportunity of re-
marking that (so far as appears from Spence Bate's
catalogue), for two different kinds of males {Orchestia
telluris and sijhicola) which live together in the forests
of New Zealand,, only one form of female is known, and
hazardirg the supposition that we have here a similar
case. It does not seem to me to be probable that two
nearly allied species of these social Amphipoda should
occur mixed together under the same conditions of life.

As the males of several species of Melita are distin-
guished by the powerful unpaired clasp-forceps, the
females of some
other species of the
same genus are
equally distinguish-
ed from all other
Amphipoda by the
circumstance that
in them a peculiar
apparatus is de-
veloped wdiich fa-
cilitates their being
held by the male.
The coxal lamellse
of the penultimate pair of feet are produced into hook-
like processes, of which the male lays hold with the

^ Fig. 10. Coxal lamella of the penultimate pair of feet of the male (a),
and coxal lamella, .with the three following joints of the same pair of
feet of the female (&) of Melita Messalina, magn. 45 diam.

Fig. 11. Coxal lamella of the same pair of feet of the female of
M. insatiabilis.

. // /

Figs. 10 and 11.*

l^

28 HISTORY OF CRUSTACEA. Chap. IY.

bands of the first pair of feet. The two species in
which I am acquainted with this structure are amongst
the most salacious animals of their order, even females
which are laden with eggs in all stages of development,
not unfrequently have their males upon their backs.
The two species are nearly allied to Melita pahnata
Leach (Gammarus Bugesii, Edw.), which is widely
distributed on the European coasts, and has been fre-
quently investigated ; unfortunately, however, I can
find no information as to whether the females of this or
any other European species possess a similar contriv-
ance. In M. exilii all the coxal lamellae are of the
ordinary formation. Nevertheless, be this as it will,
whether they exist in two or in twenty species, the
occurrence of these peculiar hook-like processes is
certainly very limited.

Now our two species live sheltered beneath slightly
tilted stones in the neighbourhood of the shore : one of
them, Melita Messalina^ so high that it is but rarely
covered by the water ; the other, Melita insatiahilis, a
little lower; both species live together in numerous
swarms. We cannot therefore suppose that the loving
couples are threatened with disturbance more frequently
than those of other species, nor would it be more difficult
for the male, than for those of other species, in case of
his losing his female, to find a new one. Nor is it any
more easy to see how the contrivance on the body of
the female for insuring the act of copulation could be
injurious to other species. But so long as it is not
demonstrated that our species are particularly in want

Chap. IY. MELITA MESSALINA. 29

of this contrivance, or that the latter would rather be
injurious than beneficial to other species, its presence
only in these few Amphipoda will have to be regarded
not as the work of far-seeing wisdom, but as tliat of a
favourable chance made use of by Natural Selection.
Under the latter supposition its isolated occurrence is
intelligible, whilst we cannot perceive why the Creator
blessed just these few species with an apparatus whicli
he found to be quite compatible with the " general plan
of structure " of the Amphipoda, and yet denied it to
others which live under the same external conditions,
and equal them even in their extraordinary salacity.
Associated with, or in the immediate vicinity of the
two species of Melita, live two species of Allorchestes,
the pairs of which are met with almost more numerously
than the single animals, and yet their females show no
trace of the above-mentioned processes of the coxal
lamellae.

These cases, I think, must be brought to bear against
the conception supported with so much genius and
knowledge by Agassiz, that species are embodied
thoughts of the Creator; and, with these, all similar
instances in which arrangements which would be equally
beneficial to all the species of a group are wanting in
the majority and only conferred upon a few special
favourites, which do not seem to want them any more
than the rest.

30 HISTORY OF CRUSTACEA. Chap. V.

CHAPTER V.

RESPIRATION IN LAND CRABS,

Among tlie numerous facts in the natural history of
the Crustacea upon which a new and clear light is
thrown by Darwin's theory, besides the two forms of
the males in our Tanais and in Orchestia Darwinii,
there is one which appears to me of particular im-
portance, namely, the character of the branchial cavity
in the air-breathing Crabs, of which, unfortunately, I
have been unable to investigate some of the most
remarkable {Gecarcinus, Banina). As this character,
namely, the existence of an entrance behind the
branchiae, has hitherto been noticed, even as a fact,
only in Banina, I will go into it in some detail. I
have ah-eady mentioned that, as indeed is required by
Darwin's theory, this entrant orifice is produced in
different manners in the different families.

In the Frog-crab (Banina ) of the Indian Ocean,
which, according to Eumphius, loves to climb up on
the roofs of the houses, the ordinary anterior entrant
orifice is entirely wanting according to Milne-Edwards,
and the entrance of a canal opening into the hindmost
parts of the branchial cavity is situated beneath the
commencement of the abdomen.

Chap. V. RESPIEATION IN LAND CRABS. 31

The case is most simple in some of the Grapsoidae,
as in Aratus Fisonii, a charming, lively Crab which
ascends the mangrove bushes (Bhizophora) and gnaws
their leaves. By means of its short but remarkably
acute claws, which prick like pins when it runs over the
hand, this Crab climbs with the greatest agility upon
the thinnest twigs. Once, when I had one of these
animals sitting upon my hand, I noticed that it elevated
the hinder part of its carapace, and that by this means
a wide fissure was opened upon each side above the last
pair of feet, through which I could look far into the
branchial cavity. I have since been unable to pro-
cure this remarkable animal again, but on the other
hand, I have frequently repeated the same observation
upon another animal of the same family (apparently a
true Grajpsus), which lives abundantly upon the rocks
of our coast. Whilst the hinder part of the carapace
rises and the above-mentioned fissure is formed, the
anterior part seems to sink, and to narrow or entirely
close the anterior entrant orifice. Under water the
elevation of the carapace never takes place. The
animal therefore opens its branchial cavity in front or
behind, according as it has to breathe water or air.
How the elevation of the carapace is effected I do not
know, but I believe that a membranous sac, which
extends from the body cavity far into the branchial
cavity beneath the hinder part of the carapace, is in-
flated by the impulsion of the fluids of the body, and
the carapace is thereby raised.

32 HISTORY OF CRUSTACEA. Chap. Y.

I have also observed the same elevation of the cara-
pace in some species of the allied genera Sesarma and
Cyclograjpsus, which dig deep holes in marshy ground,
and often run about upon the wet mud, or sit, as if
keeping watch, before their burrows. One must, how-
ever, wait for a long time with these animals, when
taken out of the water, before they open their branchial
cavity to the air, for they possess a wonderful arrange-
ment, by means of which they can continue to breathe
water for some time when out of the water. The
orifices for the egress of the water which has served for
respiration, are situated in these, as in most Crabs, in
the anterior angles of the buccal frame (" cadre buccal,"
M,-Edw.), whilst the entrant fissures of the branchial
cavity extend from its hinder angles above the first
pair of feet. Now that portion of the carapace which
extends at the sides of the mouth between the two
orifices (" regions pterygostomiennes "), appears in our
animals to be divided into small square compartments.
Milne-Edwards has already pointed this out as a par-
ticularly remarkable peculiarity. This appearance is
caused partly by small wart-like elevations, and partly
and especially by curious geniculated hairs, which to a
certain extent constitute a fine net or hair-sieve extended
immediately over the surface of the carapace. Thus
when a wave of water escapes from the branchial
cavity, it immediately becomes diffused in this network
of hairs and then again conveyed back to the branchial
cavity by vigorous movements of the appendage of the

Chap. Y. EESPIRATION IN LAND CRABS. 33

outer maxilliped which works in the entrant fissure.
Whilst the water glides in this way over the carapace
in the form of a thin film, it will again saturate itself
with oxygen, and may then serve afresh for the pur-
poses of respiration. In order to complete this arrange-
ment the outer maxillipeds, as indeed has long been
known, bear a projecting ridge furnished with a dense
fringe of hairs, which commences in front near their
median line and passes backwards and outwards to the
hinder angle of the buccal frame. Thus the two ridges
of the right and left sides form together a triangle with
the apex turned forwards, — a breakwater by which the
water flowing from the branchial cavity is kept away
from the mouth and reconducted to the branchial
cavity. In very moist air the store of water contained
in the branchial cavity may hold out for hours, and it
is only when this is used up that the animal elevates its
carapace in order to allow the air to have access to its
branchiae from behind.

In Eri/phia gonagra the entrant orifices of the respi-
ratory cavity serving for aerial respiration are situated,
not, as in the Grapsoidae, above, but behind the last
pair of feet at the sides of the abdomen.

The swift-footed Sand-Crabs (Ocijpoda) are exclu-
sively terrestrial animals, and can scarcely live for a
single day in water ; in a much shorter period a
state of complete relaxation occurs and all voluntary
movements cease.^ In these a peculiar arrangement

1 As this was not observed in the sea, but iu glass vessels containing
sea-water, it might be supposed that the animals become exhausted

D

34

HISTORY OF CRUSTACEA.

Chap. V.

on the feet of the third and fourth pairs (fig. 12) has
long been known, although its connexion with the

branchial cavity has
not been suspected.
These two pairs of
feet are more closely
approximated than
the rest; the op-
posed surfaces of
their basal joints
(therefore the hind-
er surface on the
third, and the an-
terior surface on the
fourth feet) are
smooth and po-
lished, and their margins bear a dense border of long,
silky, and peculiarly formed hairs (fig. 13). Milne-
Edwards who rightly compares these surfaces, as to
their appearance, with articular surfaces, thinks that they
serve to diminish the friction between the two feet. In
considering this interpretation, the question could not

and die, not because they are under water but because tliey liave con-
sumed all the oxygen which it contained. I therefore put into the
same water from which I had just taken an unconscious Ocy2}oda, with
its legs hanging loosely down, a specimen of Lupea cUacanfha which
had been reduced to the same state by being kept in the air, and this
recovered in the water just as the Ocypoda did in the air.

- Fig. 12. Posterior entrance to the branchial cavity of Ocypoda
rhombea, Fab., nat. size. The carapace and the fourth foot of the right
side are removed.

2 Fig. 13. Points of some of the hairs of the basal joints of the foot,
magn. 45 diam.

Fig. 13.3

Fig. 12.

Chap. V. RESPIRATION IN LAND CRABS. 35

but arise why such an arrangement for the diminution
of friction should be necessary in these particular Crabs
and between these two feet, leaving out of consideration
the fact that the remarkable brushes of hair, which on
the other hand must increase friction, also remain un-
explained. But as I was bending the feet of a large
Sand-Crab to and fro in various directions, in order to
see in what movements of the animal friction occurred
at the place indicated, and whether these might, per-
haps, be movements of particular importance to it and
such as would frequently recur, I noticed, when I had
stretched the feet widely ajDart, in the hollow between
them a round orifice of considerable size, through which
air could easily be blown into the branchial cavity, and
a fine rod might even be introduced into it. The
orifice opens into the branchial cavity behind a conical
lobe, which stands above the third foot in place of a
branchia which is wanting in Ocyjpoda. It is bounded
laterally by ridges, which rise above the articulation of
the foot, and to which the lower margin of the carapace
is applied. Exteriorly, also, it is overarched by these
ridges with the exception of a narrow fissure. This
fissure is overlaid by the carapace, which exactly at this
part projects further downwards than elsewhere, -and in
this way a complete tube is formed. Whilst in Grapsus
the water is allowed to reach the branchise only from
the front, I saw it in OGyj)oda flow in also through the
orifice just described.

In the position of posterior entrant orifice and the
accompanying peculiarities of the third and

36 HISTOKY OF CKUSTACEA. Chap. V.

pairs of feet, two otlier non-aquatic species of the same
family, which I have had the opportunity of examining,
agree with Ocyjjoda. One of these, perhaps Gelasimiis
vocanSj which lives in the mangrove swamps, and likes
to furnish the mouth of its burrow with a thick, cylin-
drical chimney of several inches in height, has the
brushes on the basal joints of the feet in question com-
posed of ordinary hairs. The other, a smaller Gelasimus,
not described in Milne-Edwards' ' Natural History of
Crustacea,' which prefers drier places and is not afraid
to run about on the burning sand under the vertical
rays of the noonday sun in December, but can also
endure being in water at least for several weeks, re-
sembles Ocyjpoda in having these brushes composed of
non-setiform, delicate hairs, indeed even more deli-
cate and more regularly constructed than in Octj])oda.'^
What may be the significance of these peculiar hairs, —
whether they only keep foreign bodies from the
branchial cavity, — Avhether they furnish moisture to
the air flowing past them, — or whether, as their aspect,
especially in the small Gelasimus, reminds one of the
olfactory filaments of the Crabs, they may also perform
similar functions, — are questions the due discussion of
which -would lead us too far from our subject. Kever-
theless it may be remarked that in both species, es-
pecially in OcyjJoda, the olfactory filaments in their

"* This smaller Gelasimus is also remarkable because the chameleon-
like change of colour exhibited by many Crabs occurs very strikingly in
it. The carapace of a male which I have now before me shone with a
dazzling white in its hinder parts five minutes since when I captured it,
at present it shows a dull gray tint at the same place.

Chap. V. EESPIRATION IN LAND CRABS, 37

ordinary situation are very much reduced, and when
they are in the water their flagella never perform the
peculiar beating movements which may be observed in
other Crabs, and even in the larger Gelasimus ; more-
over, the organ of smell must probably be sought in
these air-breathing Crabs, as in the air-breathing Verte-
brata, at the entrance to the respiratory cavity.

So much for the facts with regard to the aerial respi-
ration of the Crabs. It has already been indicated
why Darwin's theory requires that when any peculiar
arrangements exist for aerial respiration, these will be
differently constructed in different families. That ex-
perience is in perfect accordance with this requirement
is the more in favour of Darwin, because the schoolmen
far from being able to foresee or explain such profound
differences, must rather regard them as extremely sur-
prising. If, in the nearly allied families of the Ocy-
podidse and Grapsoidse, the closest agreement prevails
in all the essential conditions of their structure ; if the
same plan of structure is slavishly followed in every
thing else, in the organs of sense, in the articulation of
the limbs, in every trabecula and tuft of hairs in the
complicated framework of the stomach, and in all the
arrangements subserving aquatic respiration, even to
the hairs of the flagella employed in cleaning the
branchiae, — why have we suddenly this exception, this
complete difference, in connexion with aerial respira-
tion?

The schoolmen will scarcely have an answer for this
question, except by placing themselves on the theo-

38 HISTORY OF CRUSTACEA. Chap. V.

logico-teleological stand-point whicli has justly fallen
into disfavour amongst us, and from which the mode
of production of an arrangement is supposed to be ex-
plained, if its " adaptation " to the animal can be demon-
strated. From this point of view we might certainly
say that a widely gaping fissure which had nothing pre-
judicial in it to Äratus Pisonii among the foliage of
the mangrove bushes, was not suitable to the OcyjQoda
living in sand ; that in the latter, in order to prevent
the penetration of the sand, the orifice of the branchial
cavity must be placed at its lowest part, directed down-
wards, and concealed between broad surfaces fringed
with protective brushes of hair. It is far from the
intention of these pages to enter upon a general refu-
tation of this theory of adaptation. Indeed there is
scarcely anything essential to be added to the many
admirable remarks that have been made upon this sub-
ject since the time of Spinoza. But this may be
remarked, that I regard it as one of the most import-
ant services of the Darwinian theory that it has de-
prived those considerations of usefulness which are still
undeniable in the domain of life, of their mystical
supremacy. In the case before us it is sufficient to
refer to the Gelasimus of the mangrove swamps, which
shares the same conditions of life with various Grapsoi-
dee and yet does not agree with them, but with the
arenicolous Ocy^oda.

Chap. VI. STIIUCTUEE OF THE HEAET. 39

CHAPTER YI.

STEUCTÜKE OF THE HEART IN THE EDEIOPHTHALMA.

Scarcely less striking than the example of the air-
breathing Crabs, is the behaviour of the heart in the
great section Edriophthalma, which may advantage-
ously be divided, after the example of Dana and Spence
Bate, only into two orders, the Amphipoda and the
Isopoda.

In the Amphipoda, to which the above-mentioned
naturalists correctly refer the Caprellidae and Cyamidae
(Latreille's Lwmodiiooda) , the heart has always the
same position ; it extends in the form of a long tube
through the six segments following the head, and has
three pairs of fissures, furnished with valves, for the
entrance of the blood, situated in the second, third,
and fourth of these segments. It was found to be of
this structure by La Yalette in Ni^pliargus {Gammarus
^uteanus), and by Claus in Phronima ; and I have
found it to be the same in a considerable number of
species belonging to the most different families.^

1 The yoimg animals in the egg, a little before their exclusion, are
usually particularly convenient for the observation of the fissures in the
heart ; they are generally sufficiently transparent, the movements of the
heart are less violent than at a later period, and they lie still even

4.0 HISTORY OF CRUSTACEA. Chap. VI.

The sole unimportant exception which I have hitherto
met with is presented by the genus Brachyscelus,^ in
which the heart possesses only two pairs of fissures, as it
extends forward only into the second body-segment, and
is destitute of the pair of fissures situated in this
segment in other forms.^

without the pressure of a glass cover. Considering the common opinion
as to the distribution of the Amphipoda, namely, that they increase in
multiplicity towards the poles, and diminish towards the equator, it
may seem strange that I speak of a considerable number of species on
a subtropical coast. I therefore remark that in a few months and
without examining any depths inaccessible from the shore, I obtained
38 different species, of which 34 are new, which, with the previously
known species (principally described by Dana) gives 60 Brazilian
Amphipoda, whilst Kröyer in his ' Grönlands Amfipoder ' was ac-
quainted with only 28 species, including 2 Ljemodipoda, from the
Arctic Seas, although these had been investigated by a far greater
number of Naturalists.

2 According to Milne-Edwards' arrangement the females of this genus
would belong to the " Hyperines ordinaires " and the previously un-
known males to the ' ' Hyperines anormales," the distinguishing charac-
ter of which, namely the curiously zigzagged inferior antennae, is only a
sexual peculiarity of the male animals. In systematising from single
dead specimens, as to the sex, age, &c. of which nothing is known, similar
errors are unavoidable. Thus, in order to give another example of very
recent date, a celebrated Ichthyologist, Bleeker, has lately distinguished
two groups of the Cyprinodontes as follows : some, the Cyiariuodontini,
have a " pinna analis non elongata," and the others, the Aplocheilini,
a " pinna analis elongata " : according to this the female of a little
fish which is very abundant here would belong to the first, and the
male to the second group. Such mistakes, as already stated, are
unavoidable by the " dry-skin " philosopher, and therefore excusable ;
but they nevertheless prove in how random a fashion the present
systematic zoology frequently goes on, without princij)les or sure
foundations, and how much it is in want of the infallible touchstone
for the value of the different characters, which Darwin's theory promises
to furnish.

2 I find, in Milne-Edwards' ' Le9ons sur la Physiol, et I'Anat. comp.'
iii. p. 197, the statement that, according to Frey and Leuckart, the
heart of Caprella linearis possesses five pairs of fissures. I have ex-

Chap. VI. STRUCTUEE OF THE HEART. 41

ConsideriDg this uniformity presented by the heart
in the entire order of the Ampliipoda, it cannot but
seem very remarkable, that in the very next order of
the Isopoda, we find it to be one of the most change-
able organs.

In the cheliferous Isopods (Tanais) the heart re-
sembles that of the Amphipoda in its elongated tubular
form, as well as in the number and position of the
fissures, but with this difference, that the two fissures
of each pair do not lie directly opposite each other.

In all other Isopoda the heart is removed towards
the abdomen. In the wonderfully deformed parasitic
Isopods of the Porcellanse (Entonisous
porcellanas), the spherical heart of the y/AJV

female is confined to a short space of ^""^f \f^
the elongated first abdominal segment, / ^i^^ -sf u
and seems to possess only a single pair =li' ]'

of fissures. In the male of Entoniscus P* '^\i
Cancrorum (n. sp.), the heart (fig. 16) ^^^^^^^^ X' ^
is situated in the third abdominal seg- j^jg^ j^ *
ment. In the Cassidinse, the heart
(fig. 14) is likewise short and furnished with two pairs
of fissures, situated in the last segment of the thorax
and the first segment of the abdomen. Lastly, in a
young Anilocra, I find the heart (fig. 15) extending
through the whole length of the abdomen and furnished

amined perfectly transparent young Caprellx (probably the young of
Caprella attenuata, Dana, with which they occurred), but can only find
the usual three pairs.

^ Fig. 14. Heart of a young Cassidina.

42

HISTORY OF CRUSTACEA.

Chap. VI.

with four (or five ?) fissures, which are not placed in
pairs but alternately to the right and left in successive

( / segments. In other
\\^ animals of this order,
f which I have as yet
,' ^ only cursorily examin-
( ed, further differences
I will no doubt occur.
But why, in two orders
so nearly allied to each
other, should we find
in the one sach a con-
stancy, in the other
such a variability, of
the same highly im-
portant organ ? From
the schoolmen we need
expect no explanation, they will either decline the dis-
cussion of the " wherefore " as foreign to their province,
as lying beyond the boundaries of Natural History, or
seek to put down the importunate question by means of
a sounding paraphrase of the facts, abundantly sprinkled
with Greek words. As I have unfortunately forgotten
my Greek, the second way out of the difSculty is closed
to me ; but as I luckily reckon myself not amongst the
incorporated masters, but, to use Baron von Liebig's
expression, amongst the " promenaders on the outskirts

5 Fig. 15.

6 Fig. 16.

Heart of a young Anilocra.

Abdomen of the male of Entoniscus Cancrorum.

h. Heart.

I. Liver.

Chap. 71. STEUCTURE OF THE HEART. 43

of Natural History," this affected hesitation of the
schoolmen cannot dissuade me from seeking an answer,
which indeed presents itself most naturally from Dar-
wm's point of view.

As not only the Tanaides (vv^hich reasons elsewhere
stated {vide supra) justify us in regarding as particularly
nearly related to the primitive Isopocl) and the
Amphipoda, but also the Decapod Crustacea, possess a
heart with three pairs of fissures essentially in the
same position ; and as the same position of the heart
recurs (vide infra) even in the embryos of the Mantis-
Shrimps (Squilla), in which the heart of the adult
animal, and even, as I have elsewhere shown, that of
the larvsB when still far from maturity, extends in the
form of a long tube with numerous openings far into
the abdomen, we must unhesitatingly regard the
heart of the Amphipoda as the primitive form of that
organ in the Edriophthalma. As, moreover, in these
animals the blood flows fi'om the respiratory organs to
the heart without vessels, it is very easy to see how
advantageous it must be to them to have these organs
as much approximated as possible. We have reason to
regard as the primitive mode of respiration, that occur-
ring in Tanais {vide su])ra). Now, where, as in the
majority of the Isopoda, branchiae were developed upon
the abdomen, the position and structure of the heart
underwent a change, as it approached them more nearly,
but without the reproduction of a common plan for
these earlier modes of structure, either because this
transformation of the heart took place only after the

44 HISTORY OF CRUSTACEA. Chap. VI.

division of the primary form into subordinate groups,
or because, at least at the time of this division, the
varying heart had not yet become fixed in any new
form. Where, on the contrary, respiration remained
with the anterior part of the body, — whether in the
primitive fashion of Zoea, as in the Tanaides, or by the
development of branchige on the thorax, as in the Am-
phipoda, — the primitive form of the heart was inherited
unchanged, because any variations which might make
their appearance were rather injurious than advan-
tageous, and disaj)peared again immediately.

I close this series of isolated examples with an obser-
vation which indeed only half belongs to the province
of the Crustacea to which these pages ought to be con-
fined, and which also has no further connexion with
the preceding circumstances than that of being an " in-
telligible and intelligence-bringing fact " only from the
point of view of Darwin's theory. To-day as I was
opening a specimen of Lepas anatifera in order to
compare the animal with the description in Darwin's
* Monograph on the Subclass Cirripedia,' I found in the
shell of this Cirripede, a blood-red Annelide, with a
short, flat body, about half an inch long and two lines
in breadth, with twenty-five body-segments, and without
projecting setigerous tubercles or jointed cirri. The
small cephalic lobe bore four eyes and five tentacles ; each
body-segment had on each side at the margin a tuft
of simple setae directed obliquely upwards, and at some
distance from this, upon the ventral surface, a group of
thicker setse with a strongly uncinate bidentate apex.

Chap. VI. ANNELIDE PAEASITIC IN LEPAS. 45

There was above each of the lateral tufts of bristles a
branchia, simple on a few of the foremost segments, and
then strongly arborescent to the end of the body. The
animal, a female filled with ova, evidently, from these
characters, belongs to the family of the Amphinomidse ;
the only family the members of which, being excellent
swimmers, live in the open sea.

That this animal had not strayed accidentally into
the Lepas, but appertained to it as a regular and per-
manent guest, is evidenced by its considerable size in
proportion to the narrow entrance of the test of the
Lejoas, by the complete absence of the iridescence which
usually distinguishes the skin of free Annelides and
especially of the Amphinomidse, by the formation and
position of the inferior setae, &c. But that a worm be-
longing to this particular family Amphinomidae living
in the high sea, occurs as a guest in the Lejpas, which
also floats in the sea attached to wood, &c., is at once
intelligible from the stand-point of the Darwinian theory,
whilst the relationship of this parasite to the free-living
worms of the open sea remains perfectly unintelligible
under the supposition that it was independently created
for dwelling in the Lejpas.

But however favourable the examples hitherto re-
ferred to may be for Darwin, the objection may be
raised against them, and that with perfect justice, that
they are only isolated facts, which, when the considera-
tions founded upon them are carried far beyond what
is immediately given, may only too easily lead us from
the right path, with the deceptive glimmer of an ignis

46 HISTORY OF CRUSTACEA. Chap. VI.

fatuus. The higher the structure to be raised, the
wider must be the assuring base of well -sifted facts.

Let us turn then to a wider field, that of the develop-
mental history of the Crustacea, upon which science
has already brought together a varied abundance of
remarkable facts, which, however, have remained a
barren accumulation of unmanageable raw-material,
and let us see how, under Darwin's hand, these scattered
stones unite to form a well-jointed structure, in which
everything, bearing and being borne, finds its significant
place. Under Darwin's hand ! for I shall have nothing
to do except just to place the building stones in the
position which his theory indicates for them. " When
kings build, the carters have to work."

Chap. VII. DEVELOPMENTAL HISTORY. 47

CHAPTER YII.

DEVELOPMENTAL HISTOEY OF PODOPHTHALMÄ.

Let us first glance over the extant facts.

Among tlie Stalk-eyed Crustacea (Podo^hthalma) we
know only a very few species which quit the egg in the
form of their parents," with the full number of well-
jointed appendages to the body. Tliis is the case accord-
ing to Eathke^ in the European fresh-water Crayfish,
and according to Westwood in a West Indian Land
Crab (Gecarcinus). Both exceptions therefore belong
to the small number of Stalk-eyed Crustacea which live
in fresh water or on the land, as indeed in many other
cases fresh-water and terrestrial animals undergo no
transformations, whilst their allies in the sea have a
metamorphosis to undergo. I may refer to the Earth-
worms and Leeches among the Annelida, which chiefly
belong to the land and to fresh water, — to the Flanaride
of the fresh waters and the Tetrastemma of the sparingly
saline Baltic among the Turbellaria,-^to the Pulmonate
Gasteropoda, and to the Branchiferous Gasteropoda of
the fresh waters, the young of which (according to

^ Authorities are cited only for facts which I have had no opportunity
of confirming.

48 HISTOKY OF CRUSTACEA. Chap. VII.

Troscliel's ' Handb. der Zoologie ') have no ciliated buccal
lobes, although such organs are possessed by the very
similar Periwinkles (Littorina).

All the marine forms of this section appear to be sub-
ject to a more or less considerable metamorphosis.
This appears to be only inconsiderable in the common
Lobster, the young of which, according to Van Beneden,
are distinguished from the adult animal, by having
their feet furnished, like those of Mysis, with a swim-
ming branch projecting freely outwards. From a figure
given by Couch the appendages of the abdomen and
tail also appear to be wanting.

Far more profound is the difference of the youngest
brood from the sexually mature animal in by far the
greater majority of the Podophthalma, wdiich quit the
egg in the form of Zoe'a. This young form occurs, so
far as our present observations go, in all the Crabs,
with the sole exception of the single species investigated
by Westwood. I say s'pecies, and not genius, for in the
same genus, Gecarcinus, Vaughan Thompson found
Zoea-brood,^ which is also met with in other terrestrial
crabs^; (Oc?/poc?a, Gelasimus, &c.). All the Anomura

2 Bell (' Brit. Stalk-eyed Crust.' p. xlv.) considers himself justified in
" eliminating " Thompson's observation at once, because he could only
have examined ovigerous females preserved in alcohol. But any one
who had paid so much attention as Thompson to the development of
these animals, must have been well able to decide with certainty upon
eggs, if not too far from maturity or badly preserved, whether a Zoea
would be produced from them. Moreover, the mode of life of the Land-
Crabs is in favour of Thompson. " Once in the year," says Troschel's
' Handbuch der Zoologie,' " they migrate in great crowds to the sea
in order to deposit their eggs, and afterwards return much exhausted

Chap. VII. ZOEA-BKOOD. 49

seem likewise to commence their lives as Zoeae :
witness the Porcellanse, the Tatuira (Hi^pa emerita) and
the Hermit Crabs. Among the Macrura we are ac-
quainted with the same earliest form principally in
several Shrimps and Prawns, such as Crangon (Du
Cane), Caridina (Joly), Eijppolyte, Falcemon, Alpheus,
&c. Lastly, it is not improbable, that the youngest
brood of the Mantis-Shrimps (Squilla) is also in the
same case.

The most important peculiarities which distinguish
this Zoea-brood from the adult animal, are as fol-
lows : —

The middle-body with its appendages, those five pairs
of feet to which these animals owe their name of Deca-
poda, is either entirely wanting, or scarcely indicated ;
the abdomen and tail are destitute of appendages, and
the latter consists of a single piece. The mandibles, as in
the Insecta, have no palpi. The maxillipedes, of which
the third pair is often still wanting, are not yet brought
into the service of the mouth, but appear in the form
of biramose natatory feet. Branchiae are wanting, or
where their first rudiments may be detected as small
verruciform prominences, these are dense cell-masses,
through which the blood does not yet flow, and which
therefore have nothing to do with respiration. An in-
terchange of the gases of the water apd blood may occur
all over the thin-skinned surface of the body ; but the

towards their dwelling places, which are reached only by a few." For
what purpose would be these destructive migrations in species whose
young quit the egg and the mother as terrestrial animals ?

E

50

HISTORY OF CRUSTACEA.

Chap. YII.

lateral parts of the carapace may unhesitatingly be in-
dicated as the chief seat of respiration. They consist,
exactly as described by Leydig in the Daphnie, of an
outer and inner lamina, the space between which is
traversed by numerous transverse partitions dilated at
their ends; the spaces between these partitions are
penetrated by a more abundant flow of blood than
occurs anywhere else in the body of the Zoea. To this

Fig. 17.3

Fig. 18.4

may be added that a constant current of fresh water
passes beneath the carapace in a direction from behind
forwards, maintained as in the adult animal, by a folia-
ceous or linguiform appendage of the second pair of
maxillae (fig. 18). The addition of fine coloured par-
ticles to the water allows this current of water to be
easily detected even in small Zoeae.

3 Fig, 17. Zoea of a Marsh Crab (Cydograpsus ?), magn, 45 diam.
■* Fig. 18. Maxilla of the second pair in the same species, magn.
180 diam.

Chap. VII. ZOE^ OF TEUE CRABS. 51

The Zoese of the Crabs (fig. 17) are usually distin-
guished by long, spiniform processes of the carapace.
One of these projects upwards from the middle of the
back, a second downwards from the forehead, and fre-
quently there is a shorter one on each side near the
posterior inferior angles of the carapace. All these
processes are, however, wanting in Maia according to
Couch, and in Eurynome according to Kinahan ; and
in a third species of the same group of the Oxyrhjnchi
(belonging or nearly allied to the genus Achseus) I also
find only an inconsiderable dorsal spine, whilst the fore-
head and sides are unarmed. This is another example
warning us to be cautious in deductions from analogy.
Nothing seemed more probable than to refer back the
beak-like formation of the forehead in the Oxyrhynchi
to the frontal process of the Zoea, and now it appears
that the young of the Oxyrhynchi are really quite
destitute of any such process. The following are more
important peculiarities of the Zoeae of the Crabs,
although less striking than these processes of the cara-
pace which, in combination with the large eyes, often
give them so singular an appearance : — the anterior
(inner) antennaß are simple, not jointed, and furnished
at the extremity with from two to three olfactory fila-
ments ; the posterior (outer) antennae frequently run
out into a remarkably long spine-like process (" styli-
form process," Spence Bate), and bear, on the outside,
an appendage, which is sometimes very minute (" squami^
form process " of Spence Bate), corresponding with the

E 2

52

HISTOKY OF CRUSTACEA.

Chap. VII.

antennal scale of the Prawns,^ and the first rudiment of
the future flagellum is often already recognisable. Of
natatory feet (afterwards maxillipeds) only two pairs
are present ; the third (not, as Spence Bate thinks, the
first) is entirely wanting, or, like the five following
pairs of feet, present only as a minute bud. The tail,
of very variable form, always bears three pairs of setae
at its hinder margin. The Zoese of the Crabs usually

Figs. 19—23.8

maintain themselves in the water in such a manner
that the dorsal spine stands upwards, the abdomen is
bent forwards, the inner branch of the natatory feet is
directed forwards, and the outer one outwards and
upwards.

^ In a memoir on the metamorphoses of the Porcellame I have errone-
ously described this appendage as the " flagelkim."

^ Tails of the Zoese of various Crabs. Fig. 19. Pinnotheres. Fig. 20.
Sesarma. Fig. 21. Xantho. Figs. 22 and 23 of unknown origin.

Chap. VII.

ZOE^ OF PORCELLAN^.

53

It is further to be remarked that the Zoeae of the
crabs, as also of the Forcellanss, of the
Tatuira and of the Shrimps and Prawns,
are enveloped, on escaping from the
egg, by a membrane veiling the spinous
processes of the carapace, the setae of
the feet, and the antennae, and that
they cast this in a few hours. In
Achseus I have observed that the tail
of this earliest larval skin resembles
that of the larvae of Shrimps and Prawns,
and the same appears to be the case in
Maia (see Bell, * Brit. Stalk-eyed Crust.,'
p. 44).

Widely as they seem to differ from
them at the first glance, the Zoeae of
the Forcellanse (fig. 24) approach those
of the true Crabs very closely. The
antennae, organs of the mouth, and nata-
tory feet, exhibit the same structure.
But the tail bears five pairs of setae, and
the dorsal spine is wanting, whilst, on
the contrary, the frontal process and
the lateral spines are of extraordinary
length, and directed straight forward
and backward.

The Zoea of the Tatuira (fig. 25)
also appears to differ but little from

" Fig. 24. Zoea of Porcellana stelUcola, F. Müll.
Magn. 15 diam.

54

HISTORY OF CRUSTACEA.

Chap. VII.

^^^^^^^

Fig. 25.

those of the true Crabs, which it likewise resembles

in its mode of locomo-
tion. The carapace
possesses only a short,
broad frontal process ;
the posterior margin
of the tail is edged
with numerous short
setse.

The Zoea of the
Hermit Crabs (fig. 26)
possesses the simple
inner antennae of the
Zoea of the true Crabs ;
the outer antennae bear
upon the outside on a
short stalk a lamella of
considerable size ana-
logous to the scale of
the antennae of the
Prawns ; on the inside,
a short, spine-like pro-
cess ; and between the
two the flagellum, still
short, but already fur-
nished with two apical

8 Fig. 25. Zoea of the Ta-
tuira {Hippa emeritd), magn.
45 diam.

9 Fig. 26. Zoea of a small
Fig. 26.' . Hermit Crab, magn. 45 diam.

Chap. YII. ZOE^ OF MACRURA. 55

setae. As in tlie Crabs, there are only two pairs of
well-developed natatory feet (maxillipedes), but tbe
third pair is also present in the form of a two-jointed
stump of considerable size, although still destitute of
setae. The tail bears five pairs of setae. The little
animal usually holds itself extended straight in the
water, with the head directed downwards.

This is also the position in which we usually see the
Zoeae of the Shrimps and Prawns (fig. 27), which agree
in their general appearance with those of the Hermit
Crabs. Between the large compound eyes there is in
them a small median eye. The inner antennae bear, at
the end of a basal joint sometimes of considerable length,
on the inside a plumose seta, which also occurs in the
Hermit Crabs, and on the outside a short terminal joint
with one or more olfactory filaments. The outer
antennae exhibit a well-developed and sometimes dis-
tinctly articulated scale, and within this usually a spini-
form process ; the flagellum appears generally to be still
wanting. The tliird pair of maxillipedes seems to be
always present, at least in the form of considerable
rudiments. The spatuliform caudal lamina bears from
five to six pairs of setae on its hinder margin.

The development of the Zoea-brood to the sexually
mature animal was traced by Spence Bate in Carcinus
masnas. He proved that the metamorphosis is a per-
fectly gradual one, and that no sharply separated stages
of development, like the caterpillar and pupa of the
Lepidoptera, could be defined in it. Unfortunate]
possess only this single complete series of obs^rvatkks * L

5ß

HISTOKY OF CRUSTACEA.

Chap. VIT.

and its results cannot be regarded at once as uni-
versally applicable ; tbus tlie young Hermit Crabs
retain the general aspect and mode of locomotion of
Zoeae, whilst the rudiments of the thoracic and abdo-
minal feet are growing, and then, when these come into

action, appear at once in a
perfectly new form, which
differs from that of the
adult animal chiefly by the
complete symmetry of the
body and by the presence of
four pairs of well-developed
natatory feet on the abdo-
men.^°

The development of the
Palinuridse seems to be very
peculiar. Claus found in the
ova of the Spiny Lobster
{Falinurus), embryos with a
completely segmented body,
but wanting the appendages
of the tail, abdomen, and last
two segments of the middle-
body; they j)ossess a single
median and considerably
compound eye ; the anterior
antennae are simple, the posterior furnished with a small

^^ Glaucothoe Peronii, M.-Edw., may be a young and still symmetrical
Pagurus of this kind.

1^ Fig. 27. Zoea of a Pdlsemon residing upon Bhizostoma cruciatum^
Less., magn. 45 diam.

Fig. 27

Chap. VII. NAUPLIUS-BROOD OF PENEUS. 57

secondary branch ; the mandibles have no palpi ; the
maxillipedes of the third pair, like the two following pairs
of feet, are divided, into two branches of nearly eqnal
length ; whilst the last of the existing pairs of feet and
the second pair of maxillipedes bear only an inconsider-
able secondary branch. Coste, as is well known, asserts
that he has bred young Phyllosomata from the ova of
this lobster — a statement that requires further proof,
especially as the more recent investigations of Claus upon
Fhyllosoma by no means appear to be in its favour.

The large compound eyes, which usually soon become
moveable, and sometimes stand upon long stalks even
in the earliest period, as well as the carapace, which
covers the entire fore-body, indicate at once that the
position of the larvae hitherto considered, notwithstand-
ing all their differences, is under the Podophthalma.
But not a single characteristic of this section is retained
by the brood of some Prawns belonging to the genus
Peneus or in its vicinity. These quit the egg with an
unsegmented ovate body, a median frontal eye, and
three pairs of natatory feet, of which the anterior are
simple, and the other two biramose — in fact, in the
larval form, so common among the lower Crustacea, to
which 0. F. Müller gave the name of NaujpUus. No
trace of a carapace ! no trace of the paired eyes ! no
trace of masticating organs near tiie mouth which is
overarched by a helmet-like hood !

In the case of one of these species the intermediate
forms w^hich lead from the ^Nauplius to the Prawn, have
been discovered in a nearly continuous series.

58

HISTOEY OF CRUSTACEA.

Chap. YII.

The youngest Nauplius (fig. 28) is immediately fol-
lowed by forms in which a fold of skin runs across the
back behind the third pair of feet, and four pairs of
stout processes (rudiments of new limbs) sprout forth
on the ventral surface. Within the third pair of feet,
powerful mandibles are developed.

Fig. 28.12

In a subsequent moult the new limbs (maxillae, and
anterior and intermediate maxillipedes) come into ac-
tion, and in this way the Nauplius becojnes a Zoea
(fig. 29), agreeing perfectly with the Zoea of the Crabs
in the number of the appendages of the body, although
very different in form and mode of locomotion and even
in many particulars of internal structure. The chief

12 j^ig_ 28. Nauplius of a Prawn, magn. 45 diam.

Chap. VII.

DEVELOPMENT OF PENEUS.

59

organs of motion are still the two anterior pairs of feet,
which are slender and furnished with long setse ; the
third pair of feet loses its branches, and becomes con-
verted into mandibles destitute of palpi. The labrum
acquires a spine di-
rected forward and
of considerable size,
which occurs in all
the Zoese of allied
species. The bira-
mose maxillipedes
appear to assist but
slightly in locomo-
tion. The forked tail
reminds us rather
of the forms occur-
ring in the lower
Crustacea, especi-
ally the Copepoda,
than of the spatuli-
form caudal plate
which characterises
the Zoese of Alpheus,
Faldßmon, Hip]JoIyte,
and other Prawns,

of the Hermit Crabs, the Tatuira ^nd the Forcellansß.
The heart possesses only one pair of fissures, and has
no muscles traversing its interior like trabeculse, whilst
in other Zoese two pairs of fissures and an interior appa-

13 -pig. 29. Young Zoea of the same Prawn, magn. 45 diam.

Fig. 29. 13

60

HISTORY OF CRUSTACEA.

Chap. VII.

ratus of trabeculae
are always distinct-
ly recognisable.

During this Zoeal
period the paired
eyes, the segments
of the middle-body
and abdomen, the
posterior maxilli-
pedes, the lateral
caudal appendages
and the stump-like
rudiments of the
feet of the middle-
body are formed
(fig.30). The caudal
appendages sprout
forth like other
limbs freely on the
ventral surface,
whilst in other
Prawns, the Porcel-
lanw, &G., they are
produced in the in-
terior of the spatuli-
form caudal j)late.

As the feet of the
middle-body come

» Fig. 30. Older Zoea
of the same Prawn, magn.
45 diam.

Chap. YII.

DEVELOPMENT OF PENEUS.

61

into action, simultaneously with other profound changes,
the Zoea passes into the My sis- or Schizopod-form (fig. 31).
The antennae cease to serve for locomotion, their place
is taken by the thoracic feet,
furnished with long setae, and
by the long abdomen which just
before was laboriously dragged
along as a useless burden, but
now, with its powerful muscles,
jerks the animal through the
water in a series of lively jumps.
The anterior antennae have lost
their long setae, and by the side
of the last (fourth) joint, en-
dowed with olfactory filaments,
there appears a second branch,
which is at first of a single joint.
The previously multi-articulate
outer branch of the posterior
antennae has become a simple
lamella, the antennal scale of
the Prawn ; beside this appears
the stump-like rudiment of the
flagellum, probably as a new
formation, the inner branch dis-
appearing entirely. The five
new pairs of feet are biramose,
the inner branch short and simple, the outer one longer?
annulated at the end, furnished with long setae, and

Fig. 31.

Fig. 31. Mysis-iorm of the same Prawn, magn. 45 diam.

62 HISTORY OF CRUSTACEA. Chap. VII.

kept, as in Mysis, in constant whirling motion. The
heart acquires new fissures, and interior muscular
trabeculae.

During the Mysis-ipeiiod, the auditory organs in the
basal joint of the anterior antennae are formed ; the
inner branches of the first three pairs of feet are deve-
loped into chelae and the two hinder pairs into ambula-
tory feet; palpi sprout from the mandibles, branchiae
on the thorax, and natatory feet on the abdomen. The
spine on the labrum becomes reduced in size. In this
way the animal gradually approaches the Prawn-form,
in which the median eye has become indistinct, the
spine of the labrum, and the outer branches of the
cheliferous and ambulatory feet have been lost, the
mandibular palpi and the abdominal feet have acquired
distinct joints and setae, and the branchiae come into
action.

In another Prawn, the various larval states of which
may be easily recognised as belonging to the same
series by the presence of a dark-yellow, sharply-defined
spot surrounding the median eye, the youngest Zoea
(fig. 32), probably produced from the Nauplius, agrees
in all essential particulars with the species just de-
scribed ; its further development is, however, very dif-
ferent, especially in that neither the feet of the middle,
nor those of the hind-body are formed simultaneously,
and that a stage of development comparable to Mysis in
the number and structure of the limbs does not occur.

Traces of the outer maxillipedes make their appear-
ance betimes. Then feet appear upon four segments

Chap. VII.

DEVELOPMENT OF PEAWNS.

63

of the middle-body, and these are biramose on the three
anterior segments, and simple, the inner branch being
deficient, on the fourth
segment. On the inner
branches the chelse are
developed ; the outer
branches are lost before
an inner branch has
made its appearance on
the fourth segment (fig.
32). The latter again
becomes destitute of ap-
pendages, so that in this
case at an early period
four, and at a later only
three, segments of the
middle-body bear limbs.
The fifth segment is still
entirely wanting, whilst
all the abdominal seg-
ments have also acquired
limbs, and this one after the other, from before back-
wards. The adult animal, as shown by the three pairs
of chelae, will certainly be very nearly allied to the
preceding species.^^

Fig. 32.

^6 Fig. 32. Youngest (observed) Zoea of another Prawn. The
minute buds of the third pair of maxillipedes are visible. The forma-
tion of the abdominal segments has commenced. Paired eyes still
wanting. Magn. 45 diam.

^7 The oldest observed larvae (see fig. 33) are characterised by the
extraordinary length of the flagella of the outer antennae, and in this

64

HISTORY OF CRUSTACEA.

Chap. YII.

The youngest larva of the Schizopod genus Eu-
phausia observed by Claus, stands very near the
youngest Zoea of our Prawns ; but whilst its anterior
antennae are already biramose, and it therefore appears
to be more advanced, it still wants the middle maxilli-
pedes. In it also Claus found the heart furnished with
only a single pair of fissures. Do not Nauplius-like
states in this case also precede the Zoea ?

Fig. 33.18

The developmental history of Mysis, the near re-
lationship of which with the Shrimps and Prawns has
recently again been generally recognised, has been

respect resemble the larva of Sergestes found by Claus near Messina
(Zeitschr. für Wiss. Zool. Bd. xiii. Taf. 27, fig. 14). This unusual length
of the antennse leads to the supposition that they belong to our com-
monest Prawn, which is very frequently eaten, and is most nearly allied
to Peneus setiferus of Florida. Claus"s Acanthosoma (I. c. fig. 13) is like
the yoimger Mtjsis-iovm of the larva figured by me in the ' Archiv für
Naturgeschichte,' 1836, Taf. 2, fig. 18, and which I am inclined to refer
to Sicyonia carinata.

^8 Fig. 33. Older larva produced from the Zoea represented in fig. 32.
The last segment and the last two pairs of feet of the middle-body are
wanting. Magn. 20 diam.

Chap. VII. DEVELOPMENT OF MYSIS. 65

described in detail by Yan Beneden. So far as I Lave
tested tliem I can only confirm bis statements. Tbe
development of tbe embryo commences witb tbe forma-
tion of tbe tail ! This makes its appearance as a simple
lobe, tbe dorsal surface of wbicb is turned towards and
closely applied to tbat of tbe embryo. (Tbe young of
otber Stalk-eyed Crustacea are, as is well known, bent
in tbe egg in sucb a manner tbat tbe ventral surfaces
of tbe anterior and posterior balves of tbe body are
turned towards eacb otber, — in tbese, tberefore, tbe
dorsal, and in Mysis tbe ventral surface appears convex.)
Tbe tail soon acquires tbe furcate form witb wbicb we
made acquaintance in tbe last Prawn-Zoea described.
Then two pairs of tbick ensiform appendages make
tbeir appearance at tbe opposite end of tbe body, and
bebind tbese a pair of tubercles wbicb are easily over-
looked. Tbese are tbe antennae and mandibles. Tbe
egg-membrane now bursts, before any internal organ,
or even any tissue, except tbe cells of tbe cutaneous
layer, is formed. The young animal migbt be called a
Nauplius ; but essentially tbere is notbing but a roiigli
copy of a Nauplius-skin, almost like a new egg-
membrane, witbin wbicb tbe Mijsis is developed. Tbe
ten pairs of appendages of tbe fore- (maxillae, maxilli-
pedes) and middle-body make tbeir aj)pearance simul-
taneously, as do tbe five pairs of abjiominal feet at a
later period. Soon after tbe young Mysis casts tbe Nau-
plius-envelope it quits the brood-poucb of tbe mother.^^

^3 Van Beneden, who regards the eye-peduncles as limbs, cannot
however avoid remarking upon Mysis : " Ce pedicule n'apparait

J,

66 HISTOEY OF CRUSTACEA. Chap. YIT.

For some time, owing to an undue importance being
ascribed to the want of a particular branchial cavity,
Mysis, Leucifer, and Phyllosoma were referred to the
Stomapoda, which are now again limited, as originally
by Latreille, to the Mantis-shrimps (Squilla), the Glass-
shrimps (Erichthiis) and their nearest allies. Of the
developmental history of these we have hitherto been
acquainted with only isolated fragments. The tracing
of the development in the egg is rendered difficult by
the circumstance, that the Mantis-shrimps do not, like
the Decapoda, carry their spawn about with them, but
deposit it in the subterranean passages inhabited by
them in the form of th.in, round, yellow plates. The
spawn is consequently exceedingly difficult to procure,
and unfortunately it becomes spoilt in a day when it is
removed from its natural hatching place, whilst on the
contrary the progress of development may be followed
for weeks together in the eggs of a single Crab kept in
confinement. The eggs of Squilla, like those removed
from the body of the Crab, die because they are de-
prived of the rapid stream of fresh water which the
mother drives through her hole for the purpose of her
own respiration.

The accompanying representation of the embryo of
Squilla shows that it possesses a long, segmented
abdomen without appendages, a bilobate tail, six pairs
of limbs, and a short heart ; the latter only pulsates
weakly and slowly. If it acquires more limbs before

aucuppTTient comme les autres appendices, et parait avoir une autre
valeur morpliologique."

Chap. YII. DEVELOPMENT OF SQUILLA.

67

Fig. 34.

exclusion, the youngest larva must stand on the same
level as the youngest larva of
Eu^hausia observed by Claus.
Of the two larval forms at
present known which are with
certainty to be ascribed, if not
to Squilla, at least to a Stonia-
pod, I pass over the younger
one ^^ as its limbs cannot
be positively interpreted, and
will only mention that in it
the last three abdominal segments are still destitute
of appendages.
The older larva
(fig. 35), which
resembles the
mature Squilla
especially in the
structure of the
great raptorial
feet and of the
preceding pair,
still wants the six
pairs of feet fol-
lowing the rapto-
rial feet. The cor-
responding body-
segments are already well developed, an unpaired eye

2° Fig. 34. Embryo of a Squilla, magn. 45 diam. a. heart.

21 'Archiv für Naturgeschichte,' 1S63. Taf. 1.

22 Fig. 35. Older larva (Zoea) of a Stomapod, magn. 15 diam»

r 2

Fig 35.

6S HISTOEY OF CKUSTACEA. Chap. VII.

is still present, the anterior antennae are already bira-
mose, whilst the flagellum is wanting in the posterior, and
the mandibles are destitute of palpi ; the four anterior
abdominal segments bear biramose natatory feet, with-
out branchiae; the fifth abdominal segment has no
appendages, and this is also the case with the tail, which
still appears as a simple lamina, fringed on the hinder
margin with numerous short teeth. It is evident that
tlie larva stands essentially in the grade of Zoea.

Chap. VIII. DEVELOPMENT OF LIGIA. 69

CHAPTEE VIII.

DEVELOPMENTAL HISTOKY OP EDRIOPHTHALMA.

Less varied than that of the Stalk-eyed Crustacea is
the mode of development of the Isopoda and Amphi-
poda, which Leach united in the section Edriophtlialma,
or Crustacea with sessile eyes.

The Rock-Slaters {Ligia) may serve as an example
of the development of the
Isopoda. In these, as in
Mysis, the caudal portion p.
of the embryo is bent
not downwards, but up-
wards ; as in Mysis also,
a larval membrane is

Fig. 36. 1

first of all formed, within

which the Slater is developed. In Mysis this first larval
skin may be compared to a Nauplius ; in Ligia it appears
like a maggot quite destitute of appendages, but pro-
duced into a long simple tail (tig. 37). The egg-mem-
brane is retained longer than in Mysis ; it bursts only
when the limbs of the young Slater are already par-
tially developed in their full number. The dorsal sur-

^ Fig. 36. Embryo of Ligia in the egg, magn. 15 diam. D. yelk ;
L. liver.

70

HISTORY OF CRUSTACEA.

Chap. YIII.

face of the Slater is united to the larval skin a little
behind the head. At this point, when the union has
been dissolved a little before the change of skin, there
is a foliaceous appendage, which exists only for a short
time, and disappears before the young Slater quits the
brood-pouch of the mother.

Fig. 37.2

The young animal, when it begins to take care of
itself, resembles the old ones in almost all parts, except
one important difference ; it possesses only six, instead of
seven pairs of ambulatory feet ; and the last segment of
the middle-body is but slightly developed and destitute
of appendages. It need hardly be mentioned that the
sexual peculiarities are not yet developed, and that in
the males the handlike enlargements of the anterior
ambulatory feet and the copulatory appendages are
still deficient.

^ Fig. 37. Maggot-like larva of Ligia, magn. 15 diam. B. remains
of the egg-membrane. We see on the lower surface, from before back-
wards : — the anterior and posterior antennse, the mandibles, the anterior
and posterior maxillae, maxillipedes, six ambulatory feet, the last seg-
ment of the middle-body destitute of appendages, five abdominal feet,
and the caudal feet.

Chap. Vin. DEVELOPMENT OF ISOPODA. 71

To the question, liow far the development of Ligia
is repeated in the other Isopoda, I can only give an
unsatisfactory answer. The curvature of the embryo
upwards instead of downwards was met with by me as
well as by Eathke in Idothea, and likewise in Cassidina,
Philoseia, Tanais, and the Bopyridae, — indeed, I failed
to find it in none of the Isopoda examined for this
purpose. In Cassidina also the first larval skin without
appendages is easily detected ; it is destitute of the long
tail, but is strongly bent in the egg, as in Ligia, and
consequently cannot be mistaken for an "inner egg-
membrane." This, however, might
happen in Philoscia, in which the
larval skin is closely applied to the
egg-membrane (fig. 38), and is only
to be explained as the larval skin by
a reference to Ligia and Cassidina. Fig. 33.:
The foliaceous appendage on the back
has long been known in the young of the common Water
Slater (ÄseUus)^ That the last pair of feet of the thorax

^ Fig. 38. Embr5^o of a Philoscia in the egg, magn. 25 diam.

^ Leydig has compared this foh'aceous appendage of the Water Slaters
with the " green gland " or " shell-gland " of other Crustacea, assuming
that the green gland has no efferent duct and appealing to the fact
that the two organs occur " in the same place." This interpretation is
by no means a happy one. In the first place we may easily ascertain
in Leucifer, as was also found to be the case by Claus, that the " green
gland" really opens at the end of the process described by Milne-
Edwards as a " tubercule auditif" and by Spence Bate as an
" olfactory denticle." And, secondly, the position is about as dif-
ferent as it can well be. In the one case a paired gland, opening
at the base of the posterior antennae, and therefore on the lower
surface of the second segment ; in the other an unpaired structure rising

72 HISTOEY OF CRUSTACEA. Chap. VIII.

is wanting in th^ young of the Wood-lice (PorcelUonides,
M.-Edw.) and Fish-lice (Cymothoadiens, M.-Edw.) has
already been noticed by Milne-Edwards. This applies
also to the Box-Slaters (Idothea), to the viviparous Globe-
Slaters (S;ph^roma) and Shield -Slaters (Cassidina), to
the Bopyridse (Bopyrus, Entoniseus, Cryptoniscus, n. g.),
and to the Cheliferous Slaters (Tanais), and therefore
probably to the great majority of the Isopoda. All the
other limbs are usually well developed in the young
Isopoda. In Tanais alone, all the abdominal feet are
wanting (but not those of the tail) ; they are developed
simultaneously with the last pair of feet of the thorax.

The last pair of feet on the middle-body of the larva,
consequently the penultimate pair in the adult animal,
is almost always similar in structure to the preceding

pair. A remarkable excep-
tion is, however, presented
in this respect by Cryjptonis-
cus and Entoniseus, — re-
markable as a confirmation

Fig. 40. 6

of Darwin's proposition that
" parts developed in an unusual manner are very vari-
able," for in the peculiarly-formed pair of feet there
exists the greatest possible difference between the three
species hitherto observed. In Cry;ptoniscus (fig. 39)
this last foot is thin and rod-like ; in Entoniseus Can-
in the median line of the back heUnd the seventh segment, (" behind the
boundary line of the first thoracic segment," Leydig),

^ Fig. 39. Embryo of Cryptoniscus planarioides, magn. 90 diam.

^ Fig. 40. Last foot of the middle-body of the larva of Entoniseus
Porcellanse, magn. 180 diam.

Chap. VIII. RETROGRADE METAMORPHOSIS. 73

crorum remarkably long and furnished with a strongly-
thickened hand and a peculiarly constructed chela ; in
Entoniscus Porcellanse very short, imperfectly jointed,
and with a large ovate terminal joint (fig. 40).

Some Isopods undergo a considerable change imme-
diately before the attainment of sexual maturity. This
is the case with the males of Tanais which have already
been noticed, and, according to Hesse, with the Pra-
nizdd, in which both sexes are said to pass into the form
known as Anceus. But Spence Bate, a careful observer,
states that he has seen females of the form of Franiza
laden with eggs far advanced in their development.

In this order we meet for the first time with an
extensive retrograde metamorphosis as a consequence
of a parasitic mode of life. Even in some Fish-lice *
(CymotJioa) the young are lively swimmers, and the
adults stiff, stupid, heavy fellows, whose short clinging
feet are capable of but little movement. In the Bopy-
ridse (Boj)yrus, Fhryxus, Ketone, &c., which might
have been conveniently left in a single genus), which
are parasitic on Crabs, Lobsters, &c., taking up their
abode chiefly in the branchial cavity, the adult females
are usually quite destitute of eyes ; the antennae are
rudimentary ; the broad body is frequently unsymme-
trically developed in consequence of the confined
space ; its segments are more or less amalgamated with
each other ; the feet are stunted, and the appendages
of the abdomen transformed from natatory feet with
long setae into foliaceous or tongue-shaped and some-
times ramified branchiae. In the dwarfish males the

74

mSTOKY OF CRUSTACEA.

Chap. VIII.

eyes, antennae, and feet, are usually better preserved
than in the females; but on the other hand all the
appendages of the abdomen have not unfrequently dis-
appeared, and sometimes every trace of segmentation.
In the females of Entonhcus, which are found in the
body-cavity of Crabs and Porcellanw, the eyes, antennae,
and buccal organs, the segmentation of the vermiform

body, and in one spe-
cies (fig. 41) the whole
of the limbs, disappear
almost without leaving
a trace ; and Gryiotonis-
cus planario'ides would
almost be regarded as a
Flatworm rather than
an Isopod, if its eggs
and young did not betray its Crustacean nature. Among
the males of these various Bopyridae, that of Entoniscus
Forcellanm occupies tlie lowest place ; it is confined all
its life to six pairs of feet, which
are reduced to shapeless rounded
lumps.

The Amphipoda are distinguish-
able from the Isopoda at an early
period in the egg by the different
position of the embryo, the hinder
extremity of which is bent downwards. In all the ani-

Fig. 42.

Fig. 43.

7 Fig. 41. Entoniscus Cancrorum, female, magn. 3 times.
^ Fig. 42. Cryptoniscus iDlanarioides, female, magn. 3 times.
^ Fig. 43. Embryo of a Corophium, magn. 90 diam.

Chap. YIII. DEVELOPMENT OF AMPHIPODA. 75

mals of this order which have been examined for it,'^
a peculiar structure makes its appearance very early
on the anterior part of the back, by which the embryo is
attached to the " inner egg-membrane," and which has
been called the " micropylar apparatus," but improperly
as it seems to me.^^ It will remind us of the union of the
young Isopoda with the larval membrane and of the un-
paired " adherent organ " on the nape of the Cladocera,
which is remarkably developed in Evadne and persists
throughout life ; but in Dai^hnia jpulex, according to
Leydig, although present in the young animals, disap-
pears v/ithout leaving a trace in the adults.

The young animal, whilst still in the egg, acquires the
full number of its segments and limbs. In cases where
segments are amalgamated together, such as the last two
segments of the thorax in DuUchia, the last abdominal
segments and the tail in Gammarus ambulans and Coro-

^° In the genera Orchestoldea, Orchestia, Allor ehestes, Montacjua,
Batea n. g., Ainphilochus, Atijlus, Microdeutopus, Leucothoe, Melita,
Gammarus (according to Meissner and La Valette), Ampliitlioe, Cerapus,
Cyrtophium, Corophium, DuUchia, Frotella and Caprella.

^^ Little as a name may actually atfect the facts, we ouglit certainly
to confine the name "micropyle" to canals of the egg-membrane, which
serve for the entrance of the semen. But the outer egg- membrane
passes over the " micropylar apparatus " of the Amphipoda without
any perforation, according to Meissner's and La Valette's own state-
ments ; it appears never to be present before fecundation, attains its
greatest development at a subsequent period of ^the ovular life, and the
delicate canals which penetrate it do not even seem to be always pre-
sent, indeed it seems to belong to the embryo rather than to the egg-
membrane. I have never been able to convince myself that the so-
called " inner egg-membrane " is really of this nature, and not perhaps
the earliest larva skin, not formed until after impregnation, as might
be supposed with reference to Ligia, Cassidina and Philoscia.

76 HISTOKY OF CRUSTACEA. Chap. YIIT.

pliium dentatum, n. sp.. and the last abdominal segments
and tlie tail in BracJiysceliis,^^ or where one or more
segments are deficient, as in Dulichia and the Caprellse,
we find the same fusion and the same deficiencies in
young animals taken out of the brood-pouch of their
mother. Even peculiarities in the structure of the
limbs, so far as they are common to both sexes, are
usually well-marked in the newly hatched young, so
that the latter generally differ from their parents only
by their stouter form, the smaller number of the an-
tennal joints and olfactory filaments, and also of the
set86 and teeth with which the body or feet are armed,
and j)erhaps by the comparatively larger size of the
secondary flagellum. An exception to this rule is pre-
sented by the Hyperinse which usually live upon Aca-
lephse. In these the young and adults often have a
remarkably different appearance; but even in these
there is no new formation of body-segments and limbs,
but only a gradual transformation of these parts.^^

^2 According to Spence Bate, in Bracliyscelus crusculum the fifth
abdominal segment is not amalgamated with the sixth (the tail) but
with the fourth, which I should be inclined to doubt, considering the
close agreement which this species otherwise shows with the two
species that I have investigated.

^3 In the young of Hyper i a galba Spence Bate did not find any of the
abdominal feet, or the last two pairs of thoracic feet, but this very
remarkable statement required confirmation the more because he
examined these minute animals only in the dried state. Subsequently
I had the wished-for opportunity of tracing the development of a
Ilyperia which is not uncommon upon Ctenophora, especially Beroe
gilva, Eschsch. The youngest larvse from the brood-pouch of the
mother already possess the whole of the thoracic feet ; on the other
hand, like Spence Bate, I cannot find those of the abdomen. At first
simple enough, all these feet soon become converted, like the anterior

Chap. VIII. DEVELOPMENT OF AMPHIPODA. 77

Thus, in order to give a few examples, the powerful
clielse of the antepenultimate pair of feet, of Phromina

feet, into richly denticulated prehensile feet, and indeed of three
different forms, the anterior feet (fig. 44) the two following pairs
(fig. 45) and finally the three last pairs (fig. 46) being similarly con-
structed and different from the rest. ' In this form the feet remain for a
very long time, whilst the abdominal appendages grow into powerful
natatory organs, and the eyes, wliich at first seemed to me to be want-
ing, into large hemispheres. In the transition to the form of the adult
animal the last three pairs of feet (fig. 49) especially undergo a con-

* Figs. 44-46. Feet of a half-grown Hyperia Martinezii, n. sp.'' Figs.
47-49. Feet of a nearly adult male of the same species ; 44 and 47 from the
first pair of anterior feet (gnathopoda) ; 44 and 48 from the first, and 46 and
40 from the last pair of thoracic feet. Magn. 90 diam.

b N'amed after my valued friend the amiable Spanish zoologist, M. Francisco de Paula
Martinez y Saes, at present on a voyage round the world.

78 HISTORY OF CRUSTACEA. Chap. YIII.

sedenfaria, are produced, according to Pagensteclier,
from simple feet of ordinary structure ; and vice versa,
the chelae on the penultimate pair of feet of the young
Brachyscelus, become converted into simple feet. In
the youug of the last-mentioned genus the long head is
drawn out into a conical point and bears remarkably
small eyes ; in course of growth, the latter, as in most
of the Hyperinse, attain an enormous size, and almost
entirely occupy the head, which then appears sphe-
rical, &c.

The difference of the sexes which, in the Gammarinse
is usually expressed chiefly in the structure of the

siderable change. The difference between the two sexes is considerable ;
the females are distinguished by a very broad thorax, and the males
(^Lestrigo)ms) by very long antennae, of which the anterior bear an
unusual abundance of olfactory filaments.

Their youngest larvae of course cannot swim ; they are helpless little
animals which firmly cling especially to tl}e swimming laminae of their
host ; the adult Hijperise, which are not unfrequently met with free in the
sea, are, as is well known, the most admirable swimmers in their order.
(" II nage avec une rapidite' extreme," says Van Beneden of H. LatreilUi
M.-Edw.)

The transformation of the Hyperise is evidently to be regarded as
acquired and not inherited, that is to say the late appearance of the
abdominal appendages and the peculiar structure of the feet in the
yoimg are not to be brought into unison with the historical development
of the Amphipoda, but to be placed to the account of the parasitic
mode of life of the young.

As in Brachyscelus, free locomotion has been continued to the adult
and not to the young, contrary to the usual method among parasites.
Still more remarkable is a similar circumstance in Caligus, among the
parasitic Copepoda. The young animal, described by Burmeister as
a peculiar genus, Chalimus, lies at anchor upon a fish by means of
a cable springing from its forehead, and having its extremity firmly
seated in the skin of the fish. "When sexual maturity is attained, the
cable is cut, and the adult Caligi, which are admirable swimmers, are
not unfrequently captured swimming freely in the sea. (See ' Archiv,
für Naturg.' 1852, I. p. 91>

Chap. VIII. SEXUAL DIFFEEENCES IN AMPHIPODA. 79

anterior feet (gnathopoda, Sp. Bate) and in the Hype-
rinae in the structure of the antennae, is often so great
that males and females have been described as distinct
species, and even repeatedly placed in different genera
(Orchestia and Talitrus, Cerapus and Bercotlioe, Lestri-
ff onus and Hyperia) or even families {Hy]perines anor^
males and Hyi^erines orclinaires). Nevertheless it is
only developed when the animals are nearly full-grown.
Up to this period the young resemble the females in a
general way, even in some cases in which these differ
more widely than the males from the " Type " of the
order. Thus in the male Shore-hoppers (Orcliestia) the
second pair of the anterior feet is provided with a
powerful hand, as in the majority of the Amphipoda,
but very differently constructed in the females. The
young, nevertheless, resemble the female. Thus also, —
and this is an extremely rare case," — the females of
Brachyscelus are destitute of the posterior (or inferior)
antennae; the male possesses them like other Amphi-
podcc; in the young I, like Spence Bate, can find no
trace of them.

It is, however, to be particularly remarked, that the
development of the sexual peculiarities does not stand
still on the attainment of sexual maturity.

For example, the younger sexually mature males of
Orcliestia Tueurauna, n. sp., have slender inferior an-
tennae, with the joints of the flagellum not fused toge-
ther, the clasping margin (" palipa^,'" Sp. Bate) of the

^4 " I know of no case in which the inferior (antennae) are obsolete,
when the superior are developed," Dana. (Darwin, ' Monograph on the
Subclass Cirripedia, Lepadidse,' p. 15.)

80 HISTORY OF CRUSTACEA. Chap. VIII.

hand in the second pair of feet is uniformly con-
vex, the last pair of feet is slender and similar
to the preceding. Subsequently the anteimse become
thickened, two, three, or four of the first joints of the
fiagellum are fused together, the palm of the hand
acquires a 'deep emargination near its inferior angle,
and the intermediate joints of the last pair of feet
become swelled into a considerable incrassation. No
museum-zoologist would hesitate about fabricating two
distinct species, if the oldest and youngest sexually
mature males were sent to him without the uniting in-
termediate forms. In the younger males of Orcliestia
Tucuratinga, although the microscopic examination of

Fig. 50. 15 Fig. 51. 15

their testes showed that they were already sexually
mature, the emargination of the clasping margin of the
hand (represented in fig. 50) and the corresponding pro-
cess of the finger, are still entirely wanting. The same
may be observed in Cerajpus and Caprella, and probably
in all cases where hereditary sexual differences occur.

15 Fig. 50. Foot of the second pair (" second pair of gnathopoda ") of
the male and fig. 51 of the female, of Orcliestia TucuraUnga, magn. 15
diam.

Chap. VIII. DIASTYLID^. 81

Next to the extensive sections of tlie Stalk-eyed and
Sessile-eyed Crustacea, but more nearly allied to the
former than to the latter, comes the remarkable family
of the DiasUjlidse or Gumacea. The young, which Kröyer
took out of the
brood-pouch of n^
the female, and ^
which attained
one -fourth of Fig.52.i6
the length of

their mother, resembled the adult animals almost in all
parts. Whether, as in Mysis and Ligia, a transformation
occurs within the brood -pouch, which is constructed in
the same way as in Mijsis, is not known.^^ The caudal

^^ Fig. 52. Male of a Bodotria, magn. 10 diam. Note the long
inferior antennse, which are closely applied to the body, and of which
the apex is visible beneath the candal appendages.

J" A trustworthy English Naturalist, Goodsir, described the brood-
pouch and eggs of Cuma as early as 1813. Kröyer, whose painstaking
care and conscientiousness is recognised with wonder by every one who
has met him on a common field of work, confirmed Goodsir's state-
ments in 184G, and, as above mentioned, took out of the brood-poucli
embryos advanced in development and resembling their parents. By
this the question whether the Diastylidse are full-grown animals or
larvEB, is completely and for ever set at rest, and only the famous names
of Agassiz, Dana and Milne-Edwards, who would recently reduce them
again to larvaj (see Van Beneden, ' Eech. sur la Faune littor. deBelgique,'
Crustace'es, pp. 73, 74), induce me, on the basis of numerous investiga-
tions of my own, to declare in Van Beneden's words ; " Parmi toutes les
formes embryonnaires de podophthalmes ou d'e'driophthalmes que nous
avons observe'es sur nos cotes, nous n'en avons pas vu ime seule qui eut
meme la moindre ressemblance avec un Cuma quelconque." The only
thing that suits the larvse of Hippolyte, Palxmon and Alplieus, in the
family character of the Cumacea as given by Kröyer which occupies
three pages (Kröyer, ' Naturh. Tidsskrift, Ny Ksekke,' Bd. ii. pp. 203—
206j is : " Duo autenuarum paria." And this, as is well known

G

82 HISTOEY OF CRUSTACEA. Chap. VIII.

portion of the embryo in the Biastylidse, as I have
recently observed, is curved upwards as in the Isopoda,
and the last pair of feet of the thorax is wanting.

Equally scanty is our knowledge of the developmental
history of the Ostracoda. We know scarcely anything
except that their anterior limbs are developed before
the posterior one (Zenker). The development of Cyj)ris
has recently been observed by Claus : — " The youngest
stages are shell-bearing Nauplius-forms."

applies to nearly all Crustacea. How well warranted are we therefore
in identifying the latter with the former. However, it is sufficient for
any one to glance at the larva of Palxmon (fig. 27) and the Cumaceau
(fig. 52) in order to be convinced of their extraordinary similarity !

Chap. IX. DEVELOPMENT OF BKANCHIOPODA. 83

CHAPTEK IX.

DEVELOPMENTAL HISTORY OF ENTOMOSTEACA, CIERI-
PEDES, AND EHIZOCEPHALA.

The section of the Brancliiopoda includes two groups
differing even in their development, — the Phyllopoda
and the Cladocera. The latter minute animals, pro-
vided with six pairs of foliaceous feet, which chiefly
belong to the fresh waters, and are diffused under,
similar forms over the whole world, quit the egg with
their full number of limbs. The Phyllopoda, on the
contrary, in which the number of feet varies between
10 and 60 pairs, and some of which certainly live in
the saturated lie of salterns and natron-lakes, but of
which only one rather divergent genus (Nehalia) is
found in the sea,^ have to undergo a metamorphosis.
Mecznikow has recently observed the development of
Nehalia, and concludes from his observations "that
Nehalia, during its embryonal life, passes through the

^ If the Phyllopoda may be regarded as tlie nearest allies of the
Trilobites, they would furnish, with Lepidosteus and Folypterus,
Lepidosiren and Frotopterus, a further example of the preservation
in fresh waters of forms long since extinguished in the sea. The
occurrence of the Artemise in supersaline water would at the same time
show that they do not escape destruction by means of the fresh water,
but in consequence of the less amount of competition in it.

G 2

84 HISTOKY OF CRUSTACEA. Chap. IX.

Nauplius- and Zoea-stages, which in the Decapoda occur
]3artly (in Feneus) in the free state." "Therefore,"
says he, " I regard Nebalia as a Phyllopodiform Deca-
pod." The youngest larvae [of the Phyllopoda] are
Nauplii, which we have ah-eady met with exceptionally
in some Prawns, and which we shall now find repro-
duced almost without exception. The body-segments
and feet, which are sometimes so numerous, are formed
gradually from before backwards, without the indication
of any sharply-discriminated regions of the body either
by the time of their appearance or by their form. All
the feet are essentially constructed in the same manner
and resemble the maxillae of the higher Crustacea.^
We might regard the Phyllopoda as Zoeae which have
not arrived at the formation of a peculiarly endowed
abdomen or thorax, and instead of these have repeatedly
reproduced the appendages which first follow the
Nauplius-limbs.

Of the Copepoda — some of which, living in a free
state, people the fresh waters, and in far more multi-
farious forms the sea, whilst others, as parasites, infest
animals of the most various classes and often become
wonderfully deformed — the developmental history, like
their entire natural history, was, until lately, in a very
unsatisfactory state. It is true, that we long ago knew
that the Cyclopes of our fresh waters were excluded in
the Nauplius-form, and that we were acquainted with
some others of their young states; we had learnt,

2 "The maxilla of the Decapod-larva (Krebslarve) is a sort of
Phyllopodal foot" (Claus).

Chap. IX. DEVELOPMENT OF COPEPODA. 85

througli Norclmann, that the same earliest form be-
longed to several parasitic Crustacea, which had pre-
viously passed, almost universally, as worms ; but the
connecting intermediate forms which would have per-
mitted us to refer the regions of the body and the limbs
of the larvae to those of the adult animal, were wanting.
The comprehensive and careful investigations of Claus
have filled up this deficiency in our knowledge, and
rendered the section of the Copepoda one of the best
known in the whole class. The following statements
are derived from the works of this able naturalist.
From the abundance of valuable materials which they
contain I select only those which are indispensable for
the comprehension of the development of the Crustacea
in general, because, in what relates to the Copepoda in
particular, the facts have already been placed in the
proper light by the representation of their most recent
investigator, and must appear to any one whose eyes
are open, as important evidence in favour of the Dar-
winian theory.^

All the larvöß of the free Copepoda investigated by
Claus, have, at the earliest period, three pairs of limbs
(the future antennae and mandibles), the anterior with a
single, and the two following ones with a double series
of joints, or branchiae. The unpaired eye, labrum, and
mouth, already occupy their permanent positions. The
posterior portion, which is usually short and destitute of
limbs, bears two terminal setae, between which the juaus

3 I am still unacquainted with Claus' latest and largöl^Wo^i:/^^!!! fio< ^
doubt the same may be said of it. /-'-■'' '^^""^ " *"^ '^'^ * -^

86

HISTOEY OF CKUSTACEA.

Chap. IX.

is situated. The form in tliis Kaupli us -brood is ex-
tremely various, — it is sometimes compressed laterally,
sometimes flat, — sometimes elongated, sometimes oval,
sometimes round or even broader tlian long, and so
forth. The changes which the first larval stages under-
go during the progress of growth, consist essentially in
an extension of the body and the sprouting forth of
new limbs. " The following stage already displays a

fourth pair of ex-
tremities, the future
maxillse." Then fol-
io w at once three new
pairs of limbs (the
maxillipedes and the
two anterior pairs of
natatory feet). The
larva still continues
like a Nauplius, as
the three anterior
pairs of limbs repre-
sent rowing feet ; at
the next moult it is
converted into the
youngest Cyclops-
like state, when it
resembles the adult animal in the structure of the
antennae and buccal organs, although the number of
limbs and body-segments is still much less, for only the

^ Figs. 53 and 54. Nauplii of Copepoda, the former magn. 90, the
hitter 180 diam.

Fig. 54. 4

Chap. IX. DEVELOPMENT OF COPEPODA. 87

rudiments of the tliird and fourth pairs of natatory feet
have made their appearance in the form of cushions
fringed with setae, and the body consists of the oval
cephalothorax, the second, third, and fourth thoracic
segments, and an elongated terminal joint. In the
Cyclopidaß the posterior antennae have lost their
secondary branch, and the mandibles have completely
thrown off the previously existing natatory feet, whilst
in the other families these appendages persist, more or
less altered. " Beyond this stage of free development,
many forms of the parasitic Copepoda, such as Lernan-
thro])iis and Chondracantlius, do not pass, as they
do not acquire the third and fourth pairs of limbs, nor
does a separation of the fifth thoracic segment from the
abdomen take place ; others (AcJitheres) even fall to a
lower grade by the subsequent loss of the two pairs of
natatory feet. But all free Copepoda, and most of the
parasitic Crustacea, pass through a longer or shorter
series of stages of development, in which the limbs
acquire a higher degree of division into joints in con-
tinuous sequence, the posterior pairs of feet are de-
veloped, and the last thoracic segment and the different
abdominal segments are successively separated from
the common terminal portion."

There is only one thing more to be indicated in the
developmental history of the parasitic Crustacea, namely
that some of them, such as AcJitheres jperearum, certainly
quit the eg^ like the rest in a Nauplius-like form, inas-
much as the plump, oval, astomatous body bears two
pairs of simple rowing feet, and behind these, as traces

88

HISTORY OF CRUSTACEA.

Chap. IX.

of the third pair, two inflations furnished each with a
long seta, but that beneath this Nauplius-skin a very-
different larva lies ready prepared, which in a few hours
bursts its clumsy envelope and then makes its appearance

in a form " which
agrees in the seg-
mentation of the
body and in the de-
velopment of the ex-
tremities with the
first Cyclops-stage "
(Claus). The en-
tire series of Nau-
plius-stages which
are passed through
by the free Copepo-
da, are in this case
completely over-
leapt.

A final and very peculiar section of the Crustacea is
formed by the two orders of the Cirripedia and Ehizo-
cephala.^

^ Fig. 55. Nauplius of Tetradita porosa after the first moult, magn.
90 diam. The brain is seen surrounding the eye, and from it the
olfactory filaments issue ; behind it are some delicate muscles passing
to the buccal hood.

^ The most various opinions prevail as to the position of the Cirripedia.
Some ascribe to them a very subordinate position among the Copepoda;
as Milne-Edwards (1852). In direct opposition to this notion of his
father's, Alph. Milne-Edwards places them (as JBasinotes) opposite to
all the other Crustacea (Eleutheronotes). Darwin regards them as
forming a peculiar sub-class equivalent to the Podophthalma, Edri-
ophthalma, &c. This appears to me to be most convenient. I would not

Fig. 5

Chap. IX. CIREIPEDIA AND EHIZOCEPHALA. 89

In these also the brood bursts out in the Nauplius-
form, and speedily strips off its earliest larva- skin which
is distinguished by no peculiarities worth noticing. Here
also we find again the same pyriform shape of the un-
segmented body, the same number and structure of the
feet, the same position of the median eye (which, how-
ever, is wanting in Sacculina purpurea, and according
to Darwin in some species of Lepas), and the same
position of the "buccal hood," as in the Nauplii of the
Prawns and Copepoda. From the latter the Nauplii of
the Cirripedia and Khizocephala are distinguished by
the possession of a dorsal shield or carapace, which
sometimes {Sacculina purpurea) projects far beyond the
body all round ; and they are distinguished not only
from other Nauplii, but as far as I know from all other
Crustacea, by the circumstance that structures which
are elsewhere combined with the two anterior limbs
(antennae), here occur separated from them.

The anterior antennae of the Copepoda, Cladocera,
Phyllopoda (Leydig, Claus), Ostracoda (at least the
Cypridinae), Diastylidae, Edriophthalma, and Podoph-
thalma, with few exceptions relating to terrestrial ani-
mals or parasites, bear peculiar filaments which I have
already repeatedly mentioned as " olfactory filaments."

combine the Ehizocephala with the Cirripedia, as Liljeborg has done,
but place them in opposition as equivalent, like the Amphipoda and
Isopoda. The near relationship of the Cirripedia to the Ostracoda is
also spoken of, but the similarity of the so-called " Cypris-like larvse,"
or Cirriped-pupsB as Darwin calls them, to Cypris is so purely external,
even as regards the shell, that the relationship appears to me to be
scarcely greater than that of Peltogaster socialis (fig. 59) with the
family of the sausages.

90

HISTOEY OF CEUSTACEA.

Chap. IX.

A pair of similar filaments spring, in the larvae of the
Cirripedia and Ehizocephala, directly from the brain.

Fig. 56.7

At the base of the inferior antennae in the Decapoda
the so-called '* green-gland" has its opening; in the
Macrura at the end of a conical process. A similar
conical process with an efferent duct traversing it is very-
striking in most of the Amphipoda. In the Ostracoda,
Zenker describes a gland situated in the base of the
inferior antennae, and opening at the extremity of an ex-
traordinarily long " spine." In the Nauplii of Cy clops

7 Fig. 56, Nauplius of Sacculina purpurea, shortly before the second
moult, magn. 180 diam. We may recognise in the first pair of feet the
future adherent feet, and in the abdomen six pairs of natatory feet with
long setae.

Chap. IX. CIRRIPEDIA AND RHIZOCEPHALA. 91

and Ci/cIoj)s{ne, Claus finds pale "shell-glands," which
commence in the intermediate pair of limbs (the poste-
rior antennae). On the other hand in the Nauplii of the
Cirripedia and Rhizocephala the *' shell-glands " open
at the ends of conical processes, sometimes of most re-
markable length, which spring from the angles of the
broad frontal margin, and have been interpreted some-
times as antennae (Burmeister, Darwin) and sometimes
as mere " horns of the carapace " (Krohn). The con-
nexion of the "shell-glands" with the frontal horns
has been recognised unmistakably in the larvae of
Lepas, and indeed the resemblance of the frontal horns
with the conical processes on the inferior antennae of
the Amphipoda, is complete throughout.^

Notwithstanding their agreement in this important
peculiarity, the Nauplii of these two orders present
material differences in many other particulars. The
abdomen of the young Cirripede is produced beneath
the anus into a long tail-like appendage which is fur-
cate at the extremity, and over the anus there is
a second long, spine-like process ; the abdomen in
the Rhizocephala terminates in two short points, — in a
" moveable caudal fork, as in the Rotatoria," (0.
Schmidt). The young Cirripedes have a mouth,
stomach, intestine, and anus, and their two posterior
pairs of limbs are beset w^ith multifarious teeth, setae,
and hooks, which certainly assist in the inception of
nourishment. All this is wanting in the young Rhizo-

8 In connexion with this it may be mentioned that, in the females of
Brachijscelus, in which the posterior antennae are deficient, the conical
processes with the canal permeating them are nevertheless retained.

92

HISTORY OF CRUSTACEA.

Chap. IX.

cephala. The Nauplii of the Cirripedia have to undergo
several moults whilst in that form ; the Nauplii of the
Kliizocephala, being astomatous, cannot of course live
long as Nauplii, and in the course of only a few days
they become transformed into equally astomatous
" pupge," as Darwin calls them.

The carapace folds itself together, so that the little
animal acquires the aspect of a bivalve shell, the fore-
most limbs become transformed into very peculiar
adherent feet (" prehensile antennae," Darwin), and the
two following pairs are cast off, like the frontal horns.
On the abdomen six pairs of powerful biramose nata-
tory feet with long setae have been formed beneath the

Nauplius-skin, and
behind these are
two short, setige-
rous caudal appen-
dages (fig. 58).

The pupae of the
Cirripedia (fig. 57),
which are likewise
astomatous, agree
completely in all
these parts with
those of the Khizo-
cephala, even to the minutest details of the segmenta-

^ Fig. 57. Pupa of a Balanide {Chthamalus 9), magn. 50 diam. The
adherent feet are retracted -within the rather opaque anterior part of
the shell.

^0 Fig. 58. Pupa of Sacculina purpurea, magn. 180 diam. The fila-
ments on the adherent feet may be the commencements of the future
roots.

Fig. 57.9

Fig. 58. 10

Chap. IX. CmRIPEDIA AND KHIZOCEPHALA. 93

tion and bristling of the natatory feet;^^ they are
especially distinguished from them by the possession of
a pair of composite eyes. Sometimes also traces of the
frontal horns seem to persist. ^^

As the Cirripedia and Khizoeephala now in general
resemble each other far more than in their Nauplius-
state, this is also the case with the individual members
of each of the two orders.

The pupae in both orders attach themselves by means
of the adherent feet ; those of the Cirripedes to rocks,
shells, turtles, drift-wood, ships, &c., — those of the
Ehizocephala to the abdomen of Crabs, Porcellance, and
Hermit Crabs. The carapace of the Cirripedes be-
comes converted, as is well-known, into a peculiar test,
on account of which they were formerly placed among
the Mollusca, and the natatory feet grow into long cirri,
which whirl nourishment towards the mouth, which is
now open. The Khizoeephala remain astomatous ; they
lose all their limbs completely, and appear as sausage-
like, sack-shaped or discoidal excrescences of their
host, filled with ova (figs. 59, 60) ; from the point of

^^ Compare the figure given by Darwin (Balanidse, PI. xxx. fig, 5) of
the first natatory foot of the pupa of LejMS australis, with that of
Lernseodiscus Porcellame published in the 'Archiv für Naturgeschichte'
(1863, Taf, iii. fig. 5). The sole distinction, that in the latter there are
only 3 setae at the end of the outer branch, whilst in the Cirripedia
there are 4 on the first and 5 on the following natatory feet, may be
due to an error on my part.

1- Darwin describes as *' acoustic orifices " small apertures in the
shell of the pupae of the Cirripedia, which, frequently surrounded by a
border, are situated, in Lepas j^ectinata, upon short, hornlike processes.
I feel scarcely any hesitation in regarding the apertures as those of the
" shell-glands," and the hornlike processes as remains of the frontal
horns.

94

HISTORY OF CRUSTACEA.

Chap. IX.

Fig. 59,

Fig. 60.

attachment closed tubes, ramified like roots, sink
into the interior of the host, twisting round its
intestine, or becoming diffused among the sac-like tubes
of its liver. The only manifestations of life which

persist in these non
^lus ultras in the
series of retrogres-
sively metamor-
phosed Crustacea,
are powerful con-
tractions of the
roots, and an alter-
nate expansion and
contraction of the
body, in consequence of which water flows into the
brood-cavity and is again expelled, through a wide
orifice.^^

Out of several Cirripedes, which are anomalous both in
structure and development, Gryptojphialus minutus must
be mentioned here ; Darwin found it in great quantities
together in the shell of ConchoIej)as peruviana on the

^^ Fig. 59. Young of Peltogaster socialis on the abdomen of a small
Hermit Crab ; in one of them the fasciculately ramified roots in the
liver of the crab are shown. Animal and roots deep yellow.

^4 Fig. 60. Young Sacculina imrpurea with its roots ; the animal
purple-red, the roots dark grass-green. Magn. 5 diam.

15 The roots of Sacculina purpurea (fig. 60) which is parasitic upon
a fcmall Hermit Crab, are made use of by two parasitic Isopods, namely
a Bopyrus and the before mentioned Cryptoniscus planario'ides (fig. 42).
These take up their abode beneath the Sacculina and cause it to die
away by intercepting the nourishment conveyed by the roots; the
roots, however, continue to grow, even without the Sacculina, and
frequently attain an extraordinary extension, especially when a Bopyrus
obtains its nourishment from them.

Chap. IX. CIRRIPEDIA AND ßHIZOCEPHALA.

95

Clionos Islands. The egg, whicli is at first elliptical,
soon, according to Darwin, becomes broader at the
anterior extremity, and acquires three club-shaped
horns, one at each anterior angle and one behind ; no
internal parts can as yet be detected. Subsequently
the posterior horn disappears, and the adherent feet
may be recognised within the anterior ones. From this
" egg-like larva " — (Darwin says of it, " I hardly know
what to call it ") — the pupa is directly produced. Its
carapace is but slightly compressed laterally and hairy,
as- in SaccuUna purpurea ; the adherent feet are of con-
siderable size, and the natatory feet are wanting, as, in
the adult animal, are the corresponding cirri. As I
learn from Mr. Spence Bate, the Nauplius-stage appears
to be overleaped and the larvae to leave the egg in the
pupa-form, in the case of a Khizocephalon {Peltogaster ?)
found by Dr. Powell in the Mauritius.

I will conclude this general view with a iew w^ords
upon the earliest pro- ..^^^
cesses in the develop- /^U^
ment of the Crustacea.
Until recently it w^as
garded as a general rule
that, by the partial seg- üiV/k^^^^
mentation of the vitel- 1 ^J

lus a germinal disc was
formed, and in this, cor-

64

Figs. 61, 62, 63, 6i. 'G

16 Figs. 01 — 63, Eggs of Tetradlta porosa in segmentation, magn.
90 diam. The larger of the two first-formed spheres of segmentation is
always turned towards the pointed end of tlie ej
Lernsiodiscus Forcellanx, in segmentation, magn. 90 diam.

96 HISTORY OF CRUSTACEA. Chap. IX.

responding to the ventral surface of the embryo, a primi-
tive band. We now know that in the Copepoda (Clans),
in the Ehizocephala (fig. 64), and, as I can add, in the
Cirripedia (figs. 61-63) the segmentation is complete, and
the embryos are sketched out in their complete form
without any preceding primitive band. Probably tlie
latter will always be the case where the young are
hatched as true Naujßii (and not merely with a Nau-
plius-skin, as in Addheres). The two modes of deve-
lopment may occur in very closely allied animals, as
is proved by Aehtheres among the Copepoda.^"^

17 I have not mentioned the Pycnogonidse, because I do not regard
them as Crustacea ; nor the XijDhosura and Trilobites, because, having
never investigated them myself, I knew too little about them, and
especially because I am unacquainted with the details of the explana-
tions given by Barrande of the development of the latter. According
to Mr. Spence Bate " the young of Trilobites are of the Nauplius-
form."

Chap.X. classification. 97

CHAPTEK X.

ON THE PKINCIPLES OF CLASSIFICATION.

Pekhaps some one else, more fortunate than myself,
may be able, even without Darwin, to find the guiding
clue through the confusion of developmental forms,
now so totally different in the nearest allies, now so sur-
prisingly similar in members of the most distant groups,
which we have just cursorily reviewed. Perhaps a
sharper eye may be able, with Agassiz, to make out " the
plan established from the beginning by the Creator," ^
who may have written here, as a Portuguese proverb
says " straight in crooked lines." ^ I cannot but think
that we can scarcely speak of a general plan, or typical
mode of development of the Crustacea, differentiated
according to the separate Sections, Orders, and Fami-
lies, when, for example, among the Macrura, the Kiver
Crayfish leaves the egg in its permanent form ; the

1 " A plan fully matured in the beginning and undeviatingly pursued ;"
or "In the beginning His plan was formed and from it He has never
swerved in any particular" (Agassiz and Gould, 'Principles of Zoology').

2 " Deos escrive direito em linhas tortas." To read this remarkable
writing we need the spectacles of Faith, which seldom suit eyes
accustomed to the Microscope.

H

98 HISTORY OF CRUSTACEA. Chap. X.

Lobster with Schizopodal feet ; Faleemon, like the Crabs,
as a Zoea ; and Peneus, like the Cirripedes, as a Nau-
pHus, — and when, still, within this same sub-order Ma-
crura, Palinurus, Mysis and Eujphausia again present
different young forms, — when new limbs sometimes
sprout forth as free rudiments on the ventral surface,
and are sometimes formed beneath the skin which
passes smoothly over them, and both modes of deve-
lopment are found in different limbs of the same animal
and in the same pair of limbs in different animals, —
when in the Podophthalma the limbs of the thorax and
abdomen make their appearance sometimes simultane-
ously, or sometimes the former and sometimes the
latter first, and when further in each of the two groups
the pairs sometimes all appear together, and some-
times one after the other, — when, among the Hyperina,
a simple foot becomes a chela in Flironima and a chela
a simple foot in Brachtjscelus, &c.

And yet, according to the teaching of the school, it is
precisely in youth, precisely in the course of develop-
ment, that the " Type " is mostly openly displayed.
But let us hear what the Old School has to tell us as to
the significance of developmental history, and its rela-
tion to comparative anatomy and systematic zoology.

Let two of its most approved masters speak.

*' Whilst comparative anatomy," said Johannes Müller,
in 1-844, in his lectures upon this science (and the
opinions of my memorable teacher were for many
years my own), " whilst comparative anatomy shows us
the infinitely multifarious formation of the same organ

Chap. X. CLASSIFICATION. 99

in the Animal Kingdom, it furnishes us at tlie same
time with the means, by the comparison of these various
forms_, of recognising the truly essential, the type of
these organs, and separating therefrom everything un-
essential. In this, developmental history serves it as a
check or test. Thus, as the idea of development is not
that of mere increase of size, but that of progress from
what is not yet distinguished, but which potentially
contains the distinction in itself, to the actually dis-
tinct,— it is clear, that the less an organ is developed, so
much the more does it approach the type, and that,
during its development, it more and more acquires
peculiarities. The types discovered by comparative
anatomy and developmental history must therefore
agree."

Then, after Johannes Müller has combated the idea
of a graduated scale of animals, and of the passage
through several animal grades during development, he
continues : — " What is true in this idea is, that every
embryo at first bears only the type of its section, from
which the type of the Class, Order, &c., is only after-
wards developed."

In 1856, in an elementary work,^ in which it is usual
to admit only what are regarded as the assured acquisi-
tions of science, Agassiz expresses himself as follows : —

" The ovarian eggs of all animals are perfectly identi-
cal, small cells with a vitellus, germinal vesicle and
germinal spot" (§ 278). " T7ie organs of the lody are

3 ' Principles of Zoology/ Part I. Comparative Physiology. By Louis
Agassiz and A. A. Gould., Eevised Edition. Boston, 1856.

H 2

100 HISTORY OF CRUSTACEA. Chap. X.

formed in the sequence of their organic importance ; the
most essential always a])]pear first. Thus the organs of
vegetative life, the intestine, &c., appear later than
those of animal life, the nervous system, skeleton, &c.,
and these in turn are preceded by the more general
phenomena belonging to the animal as such " (§ 318).
" Thus, in Fishes, the first changes consist in the seg-
mentation of the vitellus and the formation of a germ,
processes which are common to all classes of animals.
Then the dorsal furrow, characteristic of the Vertebrate,
appears — the brain, the organs of the senses ; at a later
period are formed the intestine, the limbs, and the per-
manent form of the respiratory organs, from which the
class is recognised with certainty. It is only after ex-
clusion that the peculiarities of the structure of the
teeth and fins indicate the genus and species" (§ 319).
'^ Hence the embryos of different animals resemble each
other the more, the younger they are'' (§ 320). " Conse-
quently the high importance of developmental history
is indubitable. For, if the formation of the organs takes
^lace in the order corresponding to their imjjortance, this
sequence must of itself be a criterion of their compara-
tive value in classification. The peculiarities which
appear earlier should be considered of higher value
than those which appear subsequently" (§ 321). "Ä
system, in order to he true and natural, must agree with
the sequence of the organs in the development of the
embryo'' (§322).

I do not know whether any one at the present day
will be inclined to subscribe to this proposition in its

Chap. X. CLASSIFICATION. 101

whole extent.'' It is certain, however, that views essen-
tially similar are still to be met with everywhere in
discussions on classification, and that even within the
last few years, the very sparingly successful attempts
to employ develoj)mental history as the foundation
of classification have been repeated.

But how do these propositions agree with our obser-
vations on the developmental history of the Crustacea ?
That these observations relate for the most part to
their "free metamorphosis" after their quitting the
egg, cannot prejudice their application to the pro^Dosi-
tions enunciated especially with regard to " embryonal
development " in the egg ; for Agassiz himself points
out (§ 391) that both kinds of change are of the same
nature and of equal importance and that no " radical
distinction " is produced by the circumstance that the
former take place before and the latter after birth.

" The ovarian eggs of all animals are identical, small
cells with vitellus, germinal vesicle and germinal
spot." Yes, somewhat as all Insects are identical,
small animals with head, thorax, and abdomen ; that is
to say if, only noticing what is common to them, we
leave out of consideration the difference of their de-
velopment, the presence or absence and the multifa-

^ Agassiz' own views have lately become essentially different, so far
as can be made out from Eud. Wagner's notice of his ' Essay on Classi-
fication.' Agassiz himself does not attempt any criticism of the above
cited older views, which, however, are still widely diffused. With his
recent conception I am unfortunately acquainted only from K. Wagner's
somewhat confused report, and have therefore thought it better not to
attempt any critical remarks upon it.

102 HISTORY OF CRUSTACEA. Chap. X.

rious structure of the vitelline membrane, the varying
composition of the vitellus, the different number and
formation of the germinal spots, &c. Numerous exam-
ples, which might easily be augmented, of such pro-
found differences, are furnished by Leydig's * Lehrbuch
der Histologie.' In the Crustacea the ovarian eggs
actually sometimes furnish excellent characters for the
discrimination of species of the same genus ; thus, for
example, in one Forcellana of this country they are
blackish-green, in a second deep blood-red, and in a
third dark yellow ; and within the limits of the same
order they present considerable differences in size,
which, as Yan Beneden and Claus have already pointed
out, stands in intimate connexion with the subsequent
mode of development.

*' The organs of the tody are formed in the sequence of
their organic importance ; the most essential always
aj>pear first.'' This proposition might be characterised
ä jQriori as undemonstrable, since it is impossible either
in general, or for any particular animal, to establish a
sequence of importance amongst equally indispensable
parts. Which is the more important, the lung or the
heart ? — the liver or the kidney ? — the artery or the
vein ? Instead of giving the preference, with Agassiz,
to the organs of animal life, we might with equal
justice give it to those of vegetative life, as the latter
are conceivable without the former, but not the ibrmer
without the latter. We might urge that, according to
this proposition, provisional organs as the first pro-
duced must exceed the later-formed permanent organs
in importance.

Chap. X. CLASSIFICATION. 103

But let us stick to the Crustacea. In Poly^phemus
Leydig finds the first traces of the intestinal tube even
during segmentation. In Mysis a provisional tail is
first formed, and in Ligia a maggot-like larva-skin.
The simple median eye appears earlier, and would
therefore be more important than the compound paired
eyes ; the scale of the antennae in the Prawns would
be more important than the ilagellum; the maxilli-
pedes of the Decapoda would be more important than
the chelae and ambulatory feet, and the anterior six
pairs of feet in the Isopoda, than the precisely similarly
formed seventh pair ; in the Amphipoda the most im-
portant of all organs would be the "micropylar ap-
paratus," which disappears without leaving a trace soon
after hatching ; in CijcloiJS the setae of the tail w^ould
be more important than all the natatory feet ; in the
Cirripedia the posterior antennae, as to which we do not
know what becomes of them, would be more important
than the cirri, and so forth. The most unimportant of
all organs would be the sexual organs, and the most
essential peculiarity would consist in colour, which is to
be referred back to the ovarian egg.

" The emhryos, or young states of different animals,
resemble each other the more, the younger they are,'' or, as
Johannes Müller expresses it, '' they ajo^proach the more
closely to the common typeJ' Different as may be the
ideas connected with the word " type," no one will dis-
pute that the typical form of the penultimate pair of
feet in the Amphipoda is that of a simple ambulatory
foot, and not that of a chela, for the latter occurs in no

104 HISTOKY OF CRUSTACEA. Chap. X.

single adult Amphipod ; we know it only in tlie young
of tlie genus Brachyscelus, which therefore in this
respect undoubtedly depart more widely than the
adults from the type of their order. This applies also
to the young males of the Shore-hoppers (Orchestia)
with regard to the second pair of anterior feet (gnatJio-
jooda). In like manner no one will hesitate to accept
the possession of seven pairs of feet as a "typical"
peculiarity of the Edriophthalma, which Agassiz, on
this account, names Tetradecapoda ; the young Isopoda,
which are Dodecapoda, are also in this respect further
from the " type " than the adults.

It is certainly a rule, and this Darwin's theory would
lead us to expect, that in the progress of development
those forms which are at first similar gradually depart
further from each other ; but here, as in other classes,
the exceptions, for which the Old School has no ex-
planation, are numerous. Not unfrequently we might
indeed directly reverse the proposition and assert that
the difference becomes the greater, the further we go
back in the development, and this not only in those
cases in which one of two nearly allied species is di-
rectly developed, and the other passes through several
larval stages, such as the common Crayfish and the
Prawns which are produced from Nauplius-brood.
The same may be said, for example, of the Isopoda
and Amphipoda. In the adult animals the number of
limbs is the same ; at the first sight of a CyrtopJiium or
a Dulichia, and even after the careful examination of a
Tanais, we may be in doubt whether we have an

Chap. X. CLASSIFICATION. 105

Isopod or an Ampliipod before us ; in the newly-hatched
young the number of limbs is different, and if we go
back to their existence in the egg, the most passing
glance to see whether the curvature is upwards or
downw^ards sufSces to distinguish even the youngest
embryos of the two orders.

In other instances, the courses which lead from a
similar starting-point to a similar goal, separate widely
in the middle of the development, as in the Prawns
with Nauplius-brood already described.

Finally, so that even the last possibility may be
exhausted, it sometimes happens that the greatest
similarity occurs in the middle of the development.
The most striking example of this is furnished by the
Cirripedia and Khizocephala, whether we compare the
two orders or the members of each with one another ;
from a segmentation quite different in its course (see
figs. 61-64) proceed different forms of ISTauplius, these
become converted into exceedingly similar pupge, and
from the pupae again proceed sexually mature animals,
differing from each other toto coelo.

" If the formation of the organs occurs in the order
corresponding to their imijortanee, this sequence must of
itself he a criterion of their comparative value in classifi-
cation^' THAT IS TO SAY, SUPPOSING THE PHYSIOLOGICAL
AND CLASSIFICATIONAL VALUE OF AN ORGAN TO CO-
INCIDE ! Just as in Christian countries there is a
catechismal morality, which every one has upon his
lips, but no one considers himself bound to follow, or
expects to see followed by anybody else, so also has

106 HISTORY OF CRUSTACEA. Chap. X.

Zoology its dogmas, whicli are as universally acknow-
ledged, as they are disregarded in practice. Such a
dogma as this is the supposition tacitly made by Agassiz.
Of a hundred who feel themselves compelled to give
their systematic confession of faith as the introduction
to a Manual or Monographic Memoir, ninety-nine will
commence by saying that a natural system cannot be
founded upon a single character, but that it has to take
into account all characters, and the general structure of
the animal, but that we must not simply sum up these
characters like equivalent magnitudes, that we must not
count but weigh them, and determine the importance
to be ascribed to each of them according to its physio-
logical significance. This is probably followed by a
little jingle of words in general terms on the com-
parative importance of animal and vegetative organs,
circulation, respiration, and the like. But when we
come to the work itself, to the discrimination and ar-
rangement of the species, genera, families, &c., in all
probability not one of the ninety-nine will pay the least
attention to these fine rules, or undertake the hopeless
attempt to carry them out in detail. Agassiz, for
example, like Cuvier, and in opposition to the majority
of the German and English zoologists, regards the
Kadiata as one of the great primary divisions of the
Animal Kingdom, although no one knows anything
about the significance of the radiate arrangement in
the life of these animals, and notwithstanding that the
radiate Echinodermata are produced from bilateral larvae.
The " true Fishes " are divided by him into Ctenoids

Chap. X. CLASSIFICATION. 107

and Cycloids, according as the posterior margin of their
scales is denticulated or smooth, a circumstance the
importance of which to the animal must be infinitely
small, in comparison to the peculiarities of the dentition,
formation of the fins, number of yertebrse, &c.

And, to return to our Class of the Crustacea, has any
particular attention been paid in their classification
to the distinctions prevailing in the ^' most essential
organs " ? For instance, to the nervous system ? In the
Corycseidse, Claus found all the ventral ganglia fused
together into a single broad mass, and in the Calanidae
a long ventral chain of ganglia, — the former, therefore,
in this respect resembling the Spider Crabs and the
latter the Lobster ; but no one would dream on this
account of supposing that there was a relationship be-
tween the Corycseidse and the Crabs, or the Calanidae
and the Lobsters. — Or to the organs of circulation ?
We have among the Copepoda, the Cyclopidse and
Corycaeidae without a heart, side by side . with the
CalanidsG and Pontellidae with a heart. And in the
same way among the Ostracoda, the Cy;pridinoe, which
I find possess a heart, place themselves side by side
with Cy])ris and Cythere which have no such organ. —
Or to the respiratory apparatus? Milne-Edwards did
this when he separated Mysis and Leucifer from the
Decapoda, but he himself afterwards saw that this was
an error. In one Cypridina I find branchiae of con-
siderable size, which are entirely wanting in another
species, but this does not appear to me to be a reason
for separating these species even generically.

108 HISTOEY OF CRUSTACEA. Chap. X.

On the other hand, what do we know of the physio-
logical significance of the number of segments, and all
the other matters which we are accustomed to regard
as tyjDical peculiarities of the different organs, and
to which we usually ascribe the highest systematic
value ?

" Those jpecuUarities ivMch first a])]pear, sliould he more
higlihj estimated than those ivhich a])])ear subsequently. A
system, in order to he true and natural, must agree with
the sequence of the organs in the development of the
embryo." If the earlier manifested peculiarities are to
be estimated more highly than those which afterwards
make their appearance, then in those cases in which
the structure of the adult animal requires one position
in the system, and that of the larva another, the
latter and not the former must decide the point. As
the Lerncece and Cirripedes, on account of their ISTauplius-
brood, were separated from their previous connexions
and referred to the Crustacea, we shall, for the same
reason, have to separate Peneus from the Prawns and
unite it with the Copepoda and Cirripedia. But the
most zealous embryomaniac would probably shrink
from this course.

A " true and natural system " of the Crustacea to
be in accordance with the sequence of the phenomena
would have to take into account in the first place the
various modes of segmentation, then the position of
the embryo, next the number of limbs produced within
the egg and so forth, and might be represented some-
what as follows : —

Chap. X. CLASSIFICATION. 109

CLASSIS CRUSTACEA.

Sub-class I. HoLOSCHiSTA. — Segmentation complete. No primitive

band. Nauplius-brood.
Ord. 1. Ceratometopa. — Nauplius with frontal horns. (Cirripedia,

Ehizocephala.)
Ord. 2. Leioäietopa.— Nauplius without frontal horns. (Cope-

poda, without Ächtheus, &c., Phyllopoda, Feneus.^

Sub-class II. Hemischista. — Segmentation not complete.

A. Nototropa. — Embryo bent upwards.

Ord. 3. Protura.— The tail is first formed. {Mysis.)
Ord. 3. Saccomorpha. — A maggot-like larva-skin is first formed.
(^Isopoda.)

B. Gasterotropa. — Embryo bent ventrally.

Ord. 5. Zoeogona. — Full number of limjDS not produced in the egg.

Zoea-brood. (The majority of the Podophthalmata.)
Ord. 6. Ämetahola. — Full number of limbs produced in the egg.

(^Astacus, Gecarcinus, Amphipoda less Hyperia?)

This sample may suffice. The farther we go into
details in this direction, the more brilliantly, as may
easily be imagined, does the naturalness of such an
arrangement as this force itself upon us.

All things considered, we may apply the judgment
which Agassiz pronounced upon Darwin's theory, with
far greater justice to the propositions just examined : —
" No theory," says he, " however plausible it may be,
can be admitted in science, unless it is supported by
facts."

110 HISTORY OF CRUSTACEA. Chap. XL

CHAPTEK XI.

ON THE PEOGRESS OF EVOLUTION.

From this scarcely unavoidable but unsatisfactory side-
glance upon the old school, which looks down with so
great an air of superiority upon Darwin's " intellectual
dream " and the " giddy enthusiasm " of its friends, I
turn to the more congenial task of considering the de-
velopmental history of the Crustacea from the point of
view of the Darwinian theory.

Darwin himself, in the thirteenth chapter of his book,
has already discussed the conclusions derived from his
hypotheses in the domain of developmental history.
For a more detailed application of them, however, it is
necessary in the first place to trace these general con-
clusions a little further than he has there done.

The changes by which young animals depart from
their parents, and the gradual accumulation of which
causes the production of new species, genera, and
families, may occur at an earlier or later period of life,
— in the young state, or at the period of sexual ma-
turity. For the latter is by no means always, as in
the Insecta, a period of repose ; most other animals
even then continue to grow and to undergo changes.

Chap. XI. EVOLUTION. Ill

(See above, the remarks on the males of the Amphi-
poda.) Some variations, indeed, from their very nature,
can only occur when the young animal has attained the
adult stage of development. Thus the Sea Caterpillars
(Polynoe) at first possess only a few body-segments,
which, during development, gradually increase to a
number which is different in different species, but con-
stant in the same species ; now before a young animal
could exceed the number of segments of its parents, it
must of course have attained that number. We may
assume a similar supplementary progress wherever the
deviation of the descendants consists in an addition of
new segments and limbs.

Descendants therefore reach a new goal, either hy de-
viating sooner or later whilst still on the way toivards the
form of their parents, or hy ^passing along this course
without deviation, hut then, instead of standing still, ad-
vance still farther.

The former mode wdll have had a predominant action
where the posterity of common ancestors constitutes a
group of forms standing upon the same level in es-
sential features, as the w^hole of the Amphipoda, Crabs,
or Birds. On the other hand we are led to the as-
sumption of the second mode of progress, when we seek
to deduce from a common original form, animals some
of which agree with young states of others.

In the former case the developmental history of the
descendants can only agree with that of their ancestors
up to a certain point at Avhich their courses separate, —
as to their structure in the adult state it will teach us

112

HISTOKY OF CRUSTACEA.

Chap. XI.

nothing. In the second case the entire develojpment of the
progenitors is also passed through hy the descendants, and,
therefore, so far as the production of a species depends
upon this second mode of progress, the historical develop-
ment of the species will be mirrored in its developmental
history. In the short period of a few weeks or months,

the changing forms of
the embryo and larvae
will pass before us, a
more or less complete
and more or less true
picture of the transfor-
mations through which
the species, in the course
of untold thousands of
years, has struggled up
to its present state.

One of the simplest
examples is furnished by
the development of the
Tubicolar Annelids; but
from its very simplicity
it appears well adapted
to open the eyes of many who, perhaps, would rather

Figs. 65, 66, 67. 1

1 Figs. 65-67. Young Tubicolar worms, magnified with the simple
lens about 6 diam. : 65.* without operculum, Protula-stage ; 66. with a
barbate opercular peduncle, Filograna-stage ; 67. with a naked opercular
peduncle, Serpula-st&ge.

■ Fig. 65 is drawn from memory, as the liltle animals, which I at first took for young
Protulce, only attracted my attention when I remarked the appearance of the operculum,
which induced me to draw them.

Chap. XL EVOLUTION. 113

not see, and it may therefore find a place here. Three
years ago I found on the walls of one of my glasses some
small worm-tubes (fig. 65), the inhabitants of which bore
three pairs of barbate branchial filaments, and had no
operculum. According to this we should have been
obliged to refer them to the genus Protula. A few
days afterwards one of the branchial filaments had be-
come thickened at the extremity into a clavate oper-
culum (fig. 66), when the animals reminded me, by the
barbate opercular peduncle, of the genus Filograna,
only that the latter possesses two opercula. In three
days more, during which a new pair of branchial fila-
ments had sprouted forth, the opercular peduncle had
lost its lateral filaments (fig. 67), and the worms had
become Serj)ulce. Here the supposition at once pre-
sents itself that the primitive tubicolar worm was a
Protula, — that some of its descendants, w^hich had
already become developed into perfect Protulce, subse-
quently improved themselves by the formation of an
operculum which might protect their tubes from
inimical intruders, — and that subsequent descendants
of these latter finally lost the lateral filaments of the
opercular peduncle, which they, like their ancestors,
had developed.

What say the schools to this case ? Whence and
for what purpose, if the Serpulce were produced or
created as ready-formed species, these lateral filaments
of the opercular peduncle ? To allow them to sprout
forth merely for the sake of an invariable plan of
structure, even when they must be immediately re-

I

114 HISTOEY OF CRUSTACEA. Chap. XL

tracted again as superfluous, would certainly be an
evidence rather of childish trifling or dictatorial pe-
dantry, than of infinite wisdom. But no, I am mis-
taken; from the beginning of all things the Creator
knew, that one day the inquisitive children of men
would grope about after analogies and homologies, and
that Christian naturalists would busy themselves with
thinking out his Creative ideas ; at any rate, in order
to facilitate the discernment by the former that the
opercular peduncle of the Ser^ulce is homologous with
a branchial filament, He allowed it to make a detour in
its development, and pass through the form of a bar-
bate branchial filament.

The historical record preserved in developnental his-
tory is gradually effaced as the development strikes into
a constantly straighter course from the egg to the ^perfect
animal, and it isfreq^uently sophisticated hy the struggle
for existence which the free-living larvce have to undergo.

Thus as the law of inheritance is by no means strict,
as it gives room for individual variations with regard
to the form of the parents, this is also the case with
the succession in time of the developmental processes.
Every father of a family who has taken notice of such
matters, is well aware that even in children of the same
parents, the teeth, for example, are not cut or changed,
either at the same age, or in the same order. Now in
general it will be useful to an animal to obtain as
early as possible those advantages by which it sustains
itself in the struggle for existence. A precocious ap-
pearance of peculiarities originally acquired at a later

Chap. XT. EVOLUTION. 115

period will generally be advantageous, and their re-
tarded appearance disadvantageous ; the former, when
it appears accidentally, will be preserved by natural
selection. It is the same with every change which
gives to the larval stages, rendered multifarious by
crossed and oblique characters, a more straightforward
direction, simplifies and abridges the process of deve-
lopment, and forces it back to an earlier period of life,
and finally into the life of the egg.^

As this conversion of a development passing through
different young states into a more direct one, is not the
consequence of a mysterious inherent impulse, but de-
pendent upon advances accidentally presenting them-
selves, it may take place in the most nearly allied
animals in the most various ways, and require very
different periods of time for its completion. There is
one thing, however, that must not be overlooked here.
The historical development of a species can hardly
ever have taken place in a continuously uniform flow ;
periods of rest will have alternated with periods of
rapid progress. But forms, which in periods of rapid
progress were severed from others after a short dura-
tion, must have impressed themselves less deeply upon
the developmental history of their descendants, than
those which repeated themselves unchanged, through a
long series of successive generations -in periods of rest.
These more fixed forms, less inclined to variation, will
present a more tenacious resistance in the transition
to direct development, and will maintain themselves
in a more uniform manner and to the last, however

I 2

116 HISTORY OF CRUSTACEA. CiiAr. XL

different may be the course of this process in other
respects.

In general, as already stated, it will be advanta-
geous to the young to commence the struggle for exist-
ence in the fqrm of their parents and furnished with all
their advantages — in general, but not without excep-
tions. It is perfectly clear that a brood capable of
locomotion is almost indispensable to attached animals,
and that the larvae of sluggish Mollusca, or of worms
burrowing in the ground, &c., by swarming briskly
through the sea perform essential services by dispers-
ing the species over wider spaces. In other cases a
metamorphosis is rendered indispensable by the cir-
cumstance that a division of labour has been set up
between the various periods of life ; for example, that
the larva? have exclusively taken upon themselves the
business of nourishment. A further circumstance to
be taken into consideration is the size of the eggs, — a
simpler structure may be produced with less material
than a more compound one, — the more imperfect the
larva, the smaller may the egg be, and the larger is
the number of these that the mother can furnish with
the same expenditure of material. As a rule, I believe
indeed, this advantage of a more numerous brood will
not by any means outweigh that of a more perfect
brood, but it will do so in those cases in which the
chief difficulty of the young animals consists in finding
a suitable place for their development, and in which,
therefore, it is of importance to disperse the greatest
possible number of germs, as in many parasites.

Chap. XI. EVOLUTION. 117

As the conversion of the original development
with metamorphosis into direct development is here
under discussion, this may be the proper place to say a
word as to the already indicated absence of metamor-
phosis in fresh-water and terrestrial animals the
marine allies of which still undergo a transformation.
This circumstance seems to be explicable in two ways.
Either species without a metamorphosis migrated espe-
cially into the fresh waters, or the metamorphosis was
more rapidly got rid of in the emigrants than in their
fellows remaining in the sea.

Animals without a metamorphosis would naturally
transfer themselves more easily to a new residence, as
they had only themselves and not at the same time
multifarious young forms to adapt to the new condi-
tions. But in the case of animals with a metamor-
phosis, the mortality among the larv«, always consider-
able, must have become still greater under new than
under accustomed conditions, every step towards the
simplification of the process of development must there-
fore have given them a still greater preponderance over
their fellows, and the effacing of the metamorphosis
must have gone on more rapidly. What lias taken
place in each individual case, whether the species has
immigrated after it had lost the metamorphosis, or lost
the metamorphosis after its immigration, will not
always be easy to decide. When there are marine
allies without, or with only a slight metamorphosis, like
the Lobster as the cousin of the Cray-fish, we may
take up the former supposition; when allies with a

118 HISTORY OF CRUSTACEA. Chap. XI.

metamorpliosis still live upon the land or in fresh
water, as in the case of Gecarcinus, we may adopt the
latter.

That besides this gradual extinction of the primitive
history, a falsification of the record preserved in the
developmental history takes place by means of the
struggle for existence which the free-living young
states have to undergo, requires no further exposition.
For it is perfectly evident that the struggle for exist-
ence and natural selection combined with this, must act
in the same way, in change and development, upon
Inrvae which have to provide for themselves, as upon
adult animals. The changes of the larvae, independent
of the progress of the adult animal, will become the
more considerable, the longer the duration of the life
of the larva in comparison to that of the adult animal,
the greater the difference in their mode of life, and the
more sharply marked the division of labour between
the different stages of development. These processes
have to a certain extent an action opposed to the
gradual extinction of the primitive history ; they in-
crease the differences between the individual stages of
development, and it will be easily seen how even a
straightforward course of development may be again
converted by them into a development with metamor-
phosis. By this means many, and it seems to me valid
reasons may be brought up in favour of the ojDinion
that the most ancient Insects ap]Droached more nearly
to the existing Orthoptera, and perhaps to the wingless
Blattidse, than to any other order, and that the " com-

Chap. XL EVOLUTION. 119

plete metamorphosis " of the Beetles, Lepidoptera, &c.,
is of later origin. There were, I believe, perfect
Insects before larvae and pupae ; but, on the contrary,
Nauplii and Zoeae far earlier than perfect Prawns. In
contradistinction to the inherited metamorphosis of the
Prawns, we may call that of the Coleoptera, Lepidoptera,
&c., an acquired metamorphosis.^

" I will here briefly give my reasons for the opinion that the so-
called " complete metamorphosis " of Insects, in which these animals
quit the egg as grubs or caterpillars, and afterwards become quiescent
pupse incapable of feeding, was not inherited from the primitive ancestor
of all Insects, but acquired at a later period.

The order Orthoptera, including the Pseudoneuroptera (Ephemera,
Libellula, &c.) appears to approach nearest to the primitive form of
Insects. In favour of this view we have : —

1. The structure of their buccal organs, especially the formation of the
labium, " which retains, either perfectly or approximately, the original
form of a second pair of maxillae " (Gerstäcker).

2. The segmentation of the abdomen ; " like the labium, the abdomen
also very generally retains its original segmentation, which is shown in the
development of eleven segments" (Gerstäcker). The Orthoptera with
eleven segments in the abdomen, agree perfectly in the number of their
body-segments with the Prawn-larva represented in fig. 33, or indeed,
with the higher Crustacea (Podophthalma and Edriophthalma) in
general, in which the historically youngest last thoracic segment (see
p. 123), which is sometimes late-developed, or destitute of appendages,
or even deficient, is still wanting.

3. That, as in the Crustacea, the sexual orifice and anus are placed
upon different segments ; " whilst the former is situated in the ninth
segment, the latter occurs in the eleventh " (Gerstäcker).

4. Their palseontological occurrence ; "in a fossil state the Orthoptera
make their appearance the earliest of all Insects, namely as early as the
Carboniferous formation, in which they exceed all others in number "
(Gerstäcker).

5. The absence of uniformity of habit at the present day in an order
so small when compared with the Coleoptera, Hymenoptera, &c. For
this also is usually a phenomenon characteristic of very ancient groups
of forms which have already overstepped the climax of their develop-
ment, and is explicable by extinction in mass. A Beetle or a Butterfly
is to be recognised as such at the fii'st glance, but only a thorough

120 HISTORY OF CRUSTACEA. Chap. XI.

Which of the different modes of development at
present occiiiTing in a class of animals may claim to be

investigation can demonstrate the mutual relationships of Termes,
Blat'a, Mantis, Forficula, IHphemera, Libellula, &c. I may refer to
a corresponding remarkable example from the vegetable world : amongst
Ferns the genera Aneimia, Schizxa and Lygodium, belonging to the
group ScMzR'cicex \yhich is very poor in species, dißer much more from
each other than any two forms of the group Polypodiacex which
numbers its thousands of species.

If, from all this, it seems right to regard the Orthoptera as the order
of Insects approaching most nearly to the common primitive form, we
must also expect that their mode of development will agree better with
that of the primitive form, than, for example, that of the Lepidoptera, in
the same way that some of the Prawns (Peneus) approaching most closely
the primitive form of the Decapoda, have most truly preserved their
original mode of development. Now, the majority of the Orthoj)tera
quit the egg in a form which is distinguished from that of the adult
Insect almost solely by the want of wings ; these larvse then soon
acquire rudiments of wings, which appear more strongly developed
after every moult. Even this perfectly gradual transition from the
youngest larva to the sexually mature Insect, preserves in a far higher
degree the picture of an original mode of development, than does the
so-called complete metamorphosis of the Coleoptera, Lepidoptera, or
Diptera, with its abruptly separated larva-, pupa- and imago-states.

The most ancient Insects would probably have most resembled these
wingless larvse of the existing Orthoptera. The circumstance that there
are still numerous wingless species among the Orthoptera, and that
some of these {Blattidx) are so like certain Crustacea (Isopods) in
habit that both are indicated by the same name (" Baratta " j by
the people in this country, can scarcely be regarded as of any im-
portance.

The contrary supposition that the oldest Insects possessed a " com-
plete metamorphosis," and that the " incomplete metamorphosis " of the
Orthoptera and Hemiptera is only of later origin, is met by serious
diflßculties. If all the classes of Arthropoda (Crustacea, Inseeta,
Myriopoda and Arachnida) are indeed all branches of a common stem
(and of tliis there can scarcely be a doubt), it is evident tliat the water-
inhabiting and water-breathing Crustacea must be regarded as the
original stem from which the other terrestrial classes, with their
tracheal respiration, have branched off. But nowhere among the
Crustacea is there a mode of development comparable to the "complete
metamorphosis " of the Inseeta, nowhere among the young or adult

Chap. XL EVOLUTION. 121

that approaching most nearly to the original one, is
easy to judge from the above statements.

The jprimitive history of a species will he preserved in
its developnental history the more perfectly, the longer the
series of yowig states through which it passes hy uniform
steps ; and the more truly, the less the mode of life of the
young departs from that of the adults, and the less the
peculiarities of the individual young states can he con-
ceived as transferred hack from later ones in previous
periods of life, or as independently acquired.

Let us apply this to the Crustacea.

Crustacea are there forms "which might resemble the maggots of the
Diptera or Hymenoptera, the larvae of the Coleoptera, or the caterpillars
of the Lepidoptera, still less any bearing even a distant resemblance to
the quiescent pupse of these animals. The pupse, indeed, cannot at all
be regarded as members of an original developmental series, the
individual stages of which represent permanent ancestral states, for
an animal like the mouthless and footless pupa of the Silkworm,
enclosed by a thick cocoon, can never have formed the final, sexually
mature state of an Arthropod.

In the development of the Insecta we never see new segments added
to those already present in the youngest larvae, but we do see segments
which were distinct in the larva afterwards become fused together or dis-
appear. Considering the parallelism which prevails throughout organic
nature between j^alseontological and embryonic development, it is there-
fore improbable that the oldest Insects should have possessed fewer
segments than some of their descendatits. But the larvse of the Cole-
optera, Lepidoptera, &c., never have more than nine abdominal segments,
it is therefore not probable that they represent the original young form
of the oldest Insects, and that the Orthoptera, with an abdomen of
eleven segments, should have been subsequently developed from them.

Taking into consideration on the one hand these difficulties, and on
the other the arguments which indicate the Orthoptera as the order
most nearly approaching the primitive form, it is my opinion that the
"incomplete metamorphosis" of the Orthoptera is the primitive one,
inherited from the original parents of all Insects, and the "complete
metamorphosis" of the Coleoptera, Diptera, &c., a subsequently acquired

122 HISTORY OF CRUSTACEA. Chap. XII.

CHAPTEE XII.

PROGRESS OF EVOLUTION IN CRUSTACEA.

According to all the characters established in the last
j)aragraph, the Prawn that we traced from the Naupliiis
through states analogous to Zoea and Mysis to the form
of a Macrurous Crustacean appears at present to be the
animal, which in the section of the higher Crustacea
(Malacostraca) furnishes the truest and most complete
indications of its primitive history. That it is the most
complete is at once evident. That it is the truest must
be assumed, in the first place, because the mode of life
of the various ages is less different than in the majority
of the other Podophthalma ; for from the Nauplius to
the young Prawn they w^ere found swimming freely in
the sea, whilst Crabs, Forcellance, the Tatuira, Squilla,
and many Macrura, when adult usually reside under
stones, in the clefts of rocks, holes in the earth, subter-
ranean galleries, sand, &c., not to mention other devia-
tions in habits such as are presented by the Hermit
Crabs, Pinnotheres, &l\, — and secondly and especially
because the peculiarities which distinguish the Zoea of
this species particularly from other Zocce (the employ-
ment of the anterior limbs for swimming, the furcate
tail, the simple heart, the deficiency of the paired eyes

Chap. XIT. EVOLUTION IN CRUSTACEA. 123

and abdomen at first, &c.) are neither to be deduced
from a retro-transfer of late-acquired advantages to this
early period of life, nor to be regarded at all as ad-
vantages over other Zoese which the larva might have
acquired in the struggle for existence.

A similar development must have been once passed
through by the primitive ancestor of all Malacostraca,
probably differing from that of our Prawn, especially in
the circumstance that it would go on more uniformly
without the sudden change of form and mode of locomo-
tion produced in the latter by the simultaneous sprout-
ing forth and entering into action in the Nauplius of
four and in the Zoea of five pairs of limbs. It is to be
supposed that, not only originally but even still, in the
larvae of the first JMalacostraca, the new body-segments
and pairs of limbs are formed singly, — first of all the
segments of the fore-body, then those of the abdomen,
and finally those of the middle-body, — and, moreover,
that in each region of the body the anterior segments
were formed earlier than the posterior ones, and there-
fore last of all the hindermost segment of the middle-
body. Of this original mode more or less distinct traces
still remain, even in species in which, in other respects,
the course of development of their ancestors is already
nearly effaced. Thus the abdominal feet of the Prawn-
larva represented in fig. 33, are formed singly from
before backwards, and after these the last feet of the
middle-body ; thus, in Falinurus, the last two pairs of
feet of the middle-body are formed later than the rest ;
thus in the young larvse of the Stomapoda the last

124 HISTOEY OF CRUSTACEA. Chap. XII.

three abdominal segments are destitute of limbs, wbich
are still wanting on the last of them in older larvae ;
and thus, in the Isopoda, the historically newest pair of
feet is produced later than all the rest. In the Cope-
poda this formation of new segments and limbs, gradu-
ally advancing from before backwards, is more perfectly
preserved, than in any of the higher Crustacea.^

The original development of the Malacostraca start-
ing from the Nauplius, or the lowest free-living grade
with which we are acquainted in the class of Crustacea,
is now-a-days nearly effaced in the majority of them.
That this extinction has actually taken place in the way
already deduced as a direct consequence from Darwin's
theory, will be the more easily demonstrated, the more
this process is still included in the course of life, and
the less completely it is already worn out. We may
hope to obtain the most striking examples in the still
unknown developmental history of the various Schizo-
poda, Peneidae, and, indeed, of the Macrura in general.
At present the multifarious Zoea-forms appear to be

1 It is well known that, in many cases, even in adult animals the last
segment of the middle-body, or some of its last segments, either want
their limbs or are themselves deficient {Entoniscus Porcellanse S, Leucifer,
&c,). This might be due to the animals having separated from the
common stem before these limbs were formed at all. But in those
cases with which I am best acquainted, it seems to me more probable
that the limbs have been subsequently lost again. That these particular
limbs and segments are more easily lost than others is explained by the
circumstance that, as the youngest, they have been less firmly fixed by
long-continued inheritance. ("Mr. Dana believes, that in ordinary
Crustaceans, the abortion of the segments with tlieir appendages almost
always takes place at the posterior end of the cephalothorax." — Darwin,
BalanidsB, p. 111.)

Chap. XIL EVOLUTION IN CRUSTACEA. 125

particularly instructive. Almost all the peculiarities
by whicli they depart from the primitive form of the
Zoea of Peneus (figs. 29, 30, 32), may in fact be con-
ceived as transferred back from a later period into this
early period of life. This is the case with the large
compound eyes, — with the structure of the heart, — with
the raptorial feet in Sqitilla, — and with the powerful,
muscular, straightly-extended abdomen in Falcemon,
Aljplieus, Hipjpolijte, and the Hermit Crabs. (In the
latter, indeed, the abdomen of the adult animal is a
shapeless sac filled with the liver and generative organs,
but it is still tolerably powerful in the Gimicothoe-stsigey
and was certainly still more powerful when this stage
was still the permanent form of the animal.) It is also
the case with the abdomen of the Zoeae of the Crabs,
the Poreellance, and the Tatuira, which is still powerful,
although usually bent under the breast ; the two last
swim tolerably by means of the abdomen, even when
adult, as do the true Crabs in the young state known as
Megalops. It is the case, lastly, with the conversion of
the two anterior pairs of limbs into antennae. The
second pair of antennae, which, in the various Zoeae
always remains a step behind that of the adult animal,
is particularly remarkable. In the Crabs the " scale "
is entirely wanting ; their Zoeae have it indicated in the
form of a moveable appendage, which is often exceed-
ingly minute. In the Hermit Crabs a similar, usually
moveable, spiniform process occurs as the remains of
the scale ; their Zoeae have a well-developed but in-
articulate scale. A precisely similar scale is possessed

126 HISTORY OF CRUSTACEA. Chap. XII.

by the adult Prawns, in the Zoeae of which it exists
still in a jointed form, like the outer branch of the
second pair of feet of the ISTauplius or Peneus-TiOea,.

The long, spiniform processes on the carapace of the
Zoeae of the Crabs and PorceUanse are not to be ex-
plained in this way, but their advantage to the larvae is
evident. Thus, for example, if the body of the Zoea of
Porcellana stellicola (fig. 24), without the processes
of the carapace and without the abdomen, which how-
ever is not rigidly extensible, is scarcely half a line in
length, whilst with the processes it is four lines long, a
mouth of eight times the width is necessary in order to
swallow the little animal when thus armed.^ Conse-
quently these processes of the carapace may be regarded
as acquired by the Zoea itself in the struggle for
existence.

The formation of new limbs beneath the skin of the
larvae is also to be referred to an earlier occurrence of
processes which originally took place at a later period.
The original course must have been that they sprouted
forth in a free form upon the ventral surface of the
larva in the next stage after the change of skin ; whilst
now they are developed before the change of skin, and
thus only come into action a stage earlier. In larvae
which, for other reasons, must be regarded as more
nearly approaching the primitive form, the original

2 Perseplione, a rare Crab, belonging to the family Leucosiidse, is
served in the same manner by its long chelate feet. If we seize
the animal, it extends them most obstinately straight downwards,
so that in all probability we should more easily break than bend
them.

Chap. XII. EVOLUTION IN CRUSTACEA. 127

mode usually prevails in this particular also. Thus the
caudal feet (the " lateral caudal lamellae ") are formed
freely on the ventral surface in Eujphausia and the
Prawns with Nauplius-brood, and witliin the caudal
lamellae in the Prawns with Zoea-brood, in Fagurus and
Forcellana.

A compression of several stages into one, and there-
by an abridgement and simplification of the course of
development^ is expressed in the simultaneous appear-
ance of several new pairs of limbs.

How earlier young states may gradually be completely
lost, is shown by 3Iysis and the Isopoda. In My sis
there is still a trace of the Nauplius-stage ; being trans-
ferred back to a period when it had not to provide for
itself, the Nauplius has become degraded into a mere
skin ; in Ligia (figs. 36, 37) this larva-skin has lost the
last traces of limbs, and in Fhiloscia (fig. 38) it is
scarcely demonstrable.

Like the spinous processes of the Zoeae, the chelae on
the penultimate pair of feet of the young Braeliyscelus
are to be regarded as acquired by the larva itself. The
adult animals swim admirably and are not confined to
their host ; as soon as the specimens of Chrysaora Bios-
sevillei, Less., or Rhizostoma cruciatum, Less., on which
they are seated, become the sport of the waves in the
neighbourhood of the shore, they escape from them, and
are only to be obtained from lively Acalephs. The
young are helpless creatures and bad swimmers; a
special apparatus for adhesion must be of great service
to them.

128 HISTORY OF CRUSTACEA. Chap. XII.

To review the developmental history of the different
Malacostraca in detail would furnish no results at
all correspondent to the time occupied by it, — if our
knowledge was more complete it would be more profit-
able. I therefore abandon it, but will not omit to
mention that in it many difficulties which cannot at
present be satisfactorily solved would present them-
selves. To these isolated difficulties I ascribe the less
importance, however, because even a little while ago,
before the discovery of the Prawn-Nauplius, this entire
domain of the development of the Malacostraca was
almost inaccessible to Darwin's theory.

Nor will I dwell upon the contradictions which appear
to result from the application of the Darwinian theory
to this department. I leave it to our opponents to
find them out. Most of them may easily be proved to
be only apparent. There are two of these objections,
however, which lie so much on the surface that they
can hardly escape being brought forward, and these, I
think, I must get rid of.

" The peculiarities in which the Zoese of the Crabs,
the Forcellance, the Tatuira, the Hermit Crabs, and the
Prawns with Zoea-brood agree, and by which they are
in common distinguished from the larvae of Peiieus
produced from Nauplii, forces us (it might be said) to
the supposition that the common ancestor of these
various Decapods quitted the egg in a similar Zoea-
form. But then neither Peneus with its Nauplius-
brood, nor even apparently the Falinuri could be re-
ferred back to this ancestor. The mode of development

Chap. XII. EVOLUTION IN CKUSTACEA. 129

of Peneus and Palinurus, as also several peculiar larvae
of unknown origin, but which are in all probability to
be attributed to Macrurous Crustacea, necessitate on
the contrary the opposite supposition, namely, that the
different groups of the Macrura have passed from their
original to their present mode of development inde-
pendently of each other and also independently of the
Crabs." To this we may answer that the occurrence of
the Zoea-form in all the above-mentioned Decapoda, its
existence in PenHus during the whole of that period of
life which is richest in progress and in which the wide
gap between the Nauplius and the Decapod is filled up,
its recurrence even in the development of the Stoma-
poda, the occurrence of a larval form closely approaching
the youngest Zoea of Peneus in the Schizopod genus
Euphausia, and the reminiscence of the structure of Zoea,
which even the adult Tanais has preserved in its mode
of respiration, — all indicate Zoea as one of those steps
in development which persisted as a permanent form
throughout a long period of repose, perhaps through a
whole series of geological formations, and thus has also
made a deeper impression upon the development of its
descendants, and formed a firmer nucleus in the midst
of other and more readily effaced young states. It
cannot, therefore, surprise us that in transitions from
the original mode of metamorphosis, to direct develop-
ment, even when produced independently, the larval
life commences in the same way with this Zoea-form in
different families, in which the earlier stages of de-
velopment are effaced. But except what is common to

R

130 HISTORY OF CRUSTACEA. Chap. XII.

4

all Zoese, and what may easily be explained as being trans-
ferred back from a later into this stage, the Zoese of the
Crabs, for example, agree with those of Fagurus and
Falcemon in no single peculiarity of structure which
leads us to suppose a common inheritance. Conse-
quently we may apparently assume, without hesitation,
that when the Brachyura and Macrura separated, the
primitive ancestors of each of these groups passed
through a more complete metamorphosis, and that the
transition to the present mode of development belongs
to a later period. With regard to the Brachyura, it
may be added that in them this transition occurred
only a little later and indeed before the existing families
separated. The arrangement of the processes of the
carapace, and, still more, the similar number of the
caudal setae in the most different Zoese of Crabs (figs.
19-23) prove this. Such an accordance in the number
of organs apparently so unimportant is only ex]3licable
by common inheritance. We may predict with cer-
tainty that amongst the Brachyura no species will
occur which, like Feneus, still produces Nauplius-
brood.^

As we have already seen, Mijsis and the Isopoda
depart from all other Crustacea very remarkably by
the fact that their embryos are curved upwards, instead

3 I must not omit remarking that what has been said as to the
development of the Crabs applies essentially only to the groups
Cyclometopa, Catometopa and Oxyrhyncha, placed together by Alph.
Milne-Edwards as " Eustomes." Among the Oxystomata, as also among
the " Anomura apterura," Edw., which approach so nearly to the Crabs,
I am unacquainted with the earliest young states of any of the species.

Chap. XII. EVOLUTION IN CRUSTACEA. 131

of, as elsewhere, downwards. Does not so isolated a
phenomenon as this, it might be asked, in the sense of
Darwin's theory, indicate a common inheritance ? Does
it not necessitate that we should unite as the descend-
ants of the same primitive ancestors, Mysis with the
Isopoda on the one hand, and on the other the rest of
the Podophthalma with the Amphipoda ? I think not.
Such a necessity exists only for those who estimate a
peculiarity at a higher value because it makes its ap-
pearance at an earlier period of the egg-life. Whoever
regards species as not created independently and un-
changeably, but as having gradually become what they
are, will say to himself that, when the ancestors of our
Mijsides came (probably much later than those of the
Amphipoda and Isopoda) to develope numerous body-
segments and limbs whilst still embryos, as they could
no longer find room in the egg when extended straight
out, and were therefore compelled to bend themselves,
this could only take place either upwards or down-
wards, and whatever conditions may have decided the
direction actually adopted, any near relationship to
either of the two orders of Edriophthalma could hardly
have taken part in it.

It may, however, be remarked, that the different cur-
vature of the embryo in the Amphipoda and Isopoda is so
far instructive, as it proves that their present mode of de-
velopment was adopted only after the separation of these
orders, and that, in the primitive stock of the Edrioph-
thalma, the embryos were, if not Nauplii, at least short
enough in the body to find room in the egg in an

K 2

132 HISTOKY OF CRUSTACEA. Chap. XII.

extended position, like the larvae of Achtheres enclosed
by the Nauplius-skin. On the other hand the uni-
formity of development that prevails in each of the two
orders — which is expressed in the Amphipoda for exam-
ple in the formation of the " micropylar apparatus," in
the Isopoda in the want of the last pair of ambulatory
feet — testifies that the present mode of development has
come down from a very early period and extends back
beyoüd the separation of the present families. In these
two orders also, as well as in the Crabs, we can hardly
hope to find traces of earlier young states, unless it
be in the family of the Tanaidse.'* If any one will
furnish me with an Amphipod or an Isopod with Nau-
plius-brood, the existence of which would not be more
remarkable in independently produced species than
that of a Prawn with Nauplius-brood, I will abandon
the whole Darwinian theory.

With regard to the Crabs, and also to the Isopoda
and Amphipoda, we were led to the assumption, that,
about the period when these groups started from the

•* Whether the want of the abdominal feet in the young of Tanais be
an inheritance from the time of the primitive Isopoda, or a subsequently
acquired peculiarity, which appears to me the more admissible view at
present, may perhaps be decided with some certainty, when we become
acquainted with the development and mode of life of its family allies,
Apseudes and Bhcea. The latter, as is well known, is the only Isopod
which possesses a secondary fiagellum on the anterior antennae. I have
recently obtained a new and unexpected pioof that the Tanaidx
(" Asellotes heteropodes " M.-Edw.) of all known Crustacea approach
most closely to the primitive form of the Edriophthalma. Mr. C. Spence
Bate wTites to me : " Apseudes, as far as I know, is the only Isopod in
which the antennal scale so common in the Macrura is present on the
lower antennae."

Chap. XII. EVOLUTION IN CEUSTACEA. lo3

common stem, a simplification of their process of deve-
lopment took place. This also seems to be intelligible
from Darwin's theory. When any circumstances favour-
able to a group of animals caused its wider diffusion
and divergence into forms adapting themselves to new
and various conditions of existence, this greater varia-
bility, which betrays itself in the production of new
forms, will also favour the simplification of the deve-
lopment which is almost always advantageous, and
moreover, exactly at this j)eriod, during adaptation to
new circumstances, as has already been indicated with
regard to fresh-water animals, this simplification will
be doubly beneficial, and therefore, in connexion with
this, a doubly strict selection will take place.

80 much for the development of the higher Crustacea.

A closer examination of the developmental history
of the lower Crustacea is unnecessary after what has
been said in general upon the historical significance of
the young states, and the application of this which has
just been made to the Malacostraca. We may see,
without further discussion, how the representation given
by Claus of the development of the Copepoda may pass
almost word for word as the primitive history of those
animals ; we may find in the Nauplius-skin of the larvae
of Aehtheres and in the egg-like larva of Cri/j)toj>hialus,
precisely similar traces of a transition towards direct
development, as were presented by the Nau])Ims-en\e-
lope of the embryos of Mysis and the maggot-like larva
of Ligia, &c.

It will be sufficient to indicate an essential difference

134 HISTOEY OF CRUSTACEA. Chap. XII.

in the process of development in tlie higher and lower
Crustacea. In the latter all new body- segments and
limbs which insert themselves between the two termi-
nal regions of the Naiiplius, are formed in uninterrupted
sequence from before backwards ; in the former there is
further a new formation in the middle of the body (the
middle-body), which pushes itself in between the fore-
body and the abdomen in the same way, as these have
done on their part between the head and tail of the
Nauplius. Thus, that which appears probable even
from the comparison of the limbs of the adult animal,
finds fresh support in the developmental history, namely,
that the lower Crustacea, like the Insects, are entirely
destitute of the region of the body corresponding to
the middle-body of the Malacostraca. It seems pro-
bable that the swimming feet of the Copepoda, as also
of the pupse of Cirripedia and Khizocephala, represent
the abdominal feet of the Malacostraca, that is to say,
are derived by inheritance from the same source with
them.

It would be easy to weave together the separate
threads furnished by the young forms of the various
Crustacea, into a general picture of the primitive his-
tory of this class. Such a picture, drawn with a little
skill, and finished in lively colours, would certainly
be more attractive than the dry discussions which I
have tacked on to the developmental history of these
animals. But the mode of weaving in the loose threads
would still in many cases be arbitrary, and to be
effected with equal justice in various ways ; and many

Chap. XIL EVOLUTION OF THE RHIZOCEPHALA. 1-35

gaps would still have to be filled up by means of more
or less bold assumptions. Those who have not wan-
dered much in this reoion of research would then
readily believe that they were standing upon firm
ground, where mere fancy had thrown an airy bridge ;
those acquainted with the subject, on the other hand,
would soon find out these weak points in the structure,
but would then be easily led to regard even what was
founded upon well considered facts, as merely floating
in the air. To obviate these misconceptions of its true
contents from either side, it would be necessary to ac-
company such a picture throughout with lengthy, dry
explanations. This has deterred me from further filling
in the outline which I had already sketched.

I will only give, as an example, the probable history
of the production of a single group of Crustacea, and
indeed of the most abnormal of all, the Rhizocephala,
which in the sexually mature state differ so enormously
even from their nearest allies, the Cirripedia, and from
their peculiar mode of nourishment stand quite alone
in the entire animal kingdom.

I must preface this with a few words upon the homo-
logy of the roots of the Ehizocephala, i.e. the tubules
which penetrate from its point of adhesion into the body
of the host, ramify amongst the viscera of the latter, and
terminate in caecal branchlets. In the pupae of the
Ehizocephala (fig. 58) the foremost limbs (" prehensile
antennae ") bear, on each of the two terminal joints, a
tongue-like, thin-skiuned appendage, in which we may
generally observe a few small strongly refractive gra-

136 HISTOEY OF CRUSTACEA. Chap. XII.

miles, like those seen in the roots of the adult animal.
I have therefore supposed these appendages to be the
rudiments of the future roots. A perfectly similar
appendage, " a most delicate tube or ribbon," was
found by Darwin in free-swimming pupae of Lepas
australis on the last joints of the " prehensile antennae."
From the perfect accordance in their entire structure
shown by the pupae of the Rhizocephala and Cirripedia,
there can be no doubt that the appendages of SaceuUna
and Lejpas, which are so like each other and spring from
the same spot, are homologous structures.

Now in three species of Lejpas, in Bichelaspis War-
wichii and in Scdl^ellum Peronii, Darwin saw, on tear-
ing recently-affixed animals from their point or support,
that a long narrow band issued from the same point of
the antennae ; its end was torn away, and in Dielielaspis,
judging from its ragged appearance, it had attached
itself firmly to the support. From this it follows that
this appendage in Le]jas australis can hardly be any-
thing but a young cement-duct. If, therefore, the
supposition that the appendages on the antennae of
the pupae of Khizocephala are young roots be cor-
rect, the roots of the Khizocephala are homologous
with the cement-ducts of the Cirripedia. And this,
strange as it may appear at the first glance, seems to me
scarcely doubtful. It is true that the act of adhesion
of the Rhizocephala has never yet been observed, but it
is more than probable that they attach themselves, just
like the Cirripedia, by means of the antennae, and that
therefore the points of attachment in the two groups

Chap. XII. EVOLUTION OF THE EHIZOCEPHALA. 137

indicate homologous parts of the body. From the
point of attachment in the Khizocephala the roots pene-
trate into the body of the host, whilst in the Cirripedia,
the cement-ducts issue from the same point. The roots
are blind tubes, ramified in different ways in different
species. The cement-ducts in the basis of the Balanidse
likewise constitute a generally remarkably complicated
system of ramified tubes, with regard to the mode of
termination of which nothing certain has yet been
made out. Individual csecal branches are not unfre-
quently seen even in the vicinity of the carina; and,
at least in some species, in which the cement-ducts
divide into extremely numerous and fine branchlets,
forming a network which gradually becomes denser
towards the circumference of the basis, these seem
nowhere to possess an orifice.

JN'ow as to the question: How were Cirripedia con-
verted by natural selection into Khizocephala ?

A considerable number of existing Cirripedia settle
exclusively or chiefly upon living animals ; — on Sponges,
Corals, Mollusks, Cetaceans, Turtles, Sea-Snakes, Sharks,
Crustaceans, Sea Urchins, and even on Acalephs. Di-
clielas])is Darwinii was found by Filippi in the bran-
chial cavity of Falinuriis vulgaris, and I have met with
another species of the same genus in the branchial
cavity of Lupea diacantha.

The same thing may have taken place in primitive
times. The supposition that certain Cirripedes might
once upon a time have selected the soft ventral surface
of a Crab, Porcellana or Pagurus, for its dwelling-place.

138 HISTOEY OF CRUSTACEA. Chap. XII.

has certainly notliing improbable about it. If then the
cement-clucts of such a Cirripede instead of merely
spreading on the surface, pierced or pushed before
them the soft ventral skin and penetrated into the inte-
rior of the host, this must have been beneficial to the
animal, because it would be thereby more securely
attached and protected from being thrown off during
the moulting of its host. Variations in this direction
were preserved as advantageous.

But as soon as the cement-ducts penetrated into the
body-cavity of the host and were bathed by its fluids,
an endosmotic interchange must necessarily have been
set up between the materials dissolved in these fluids
and in the contents of the cement-ducts, and this inter-
change could not be without influence upon the nourish-
ment of the parasite. The new source of nourishment
opened up in this manner was, as constantly flowing,
more certain than that offered by the nourishment
accidentally whirled into the mouth of the sedentary
animaL The individuals favoured in the development
of the cement-ducts now converted into nutriferous
roots, had more than others the prospect of abundant
food, of vigorous growth, and of producing a numerous
progeny. With tho further development, assisted by
natural selection, of the roots embracing the intestine
of the host and spreading amongst its hepatic tubes, the
introduction of nourishment through the mouth and all
the parts implicated in it, such as the whirling cirri, the
buccal organs, and the intestine, gradually lost their
importance, became aborted by disuse, and finally dis-

Chap. XII. EVOLUTION OF THE EHIZOCEPHALA. 139

appeared without leaving a trace of tlieir existence.
Protected by the abdomen of the Crab, or by the shell
inhabited by the Pagurus, the parasite also no longer
required the calcareous test, in which, no doubt, the
first Cirripedes settling upon these Decapods rejoiced.
This protective covering, having become superfluous,
also disappeared, and there remained at last only a soft
sack filled with eggs, without limbs, without mouth or
alimentary canal, and nourished, like a plant, by means
of roots, which it pushed into the body of its host.
The Cirripede had become a Khizocephalon.

If it be desired to form a notion of what our parasite
may have looked like when half way in its progress
from the one form to the other, we may consult the
figures given by Darwin, (Lepadidae PI., iv., figs. 1-7) of
Anelasma squalicola. This Lepadide, which lives upon
Sharks in the North Sea, seems, in fact, to be in the
best way to lose its cirri and buccal organs in the same
manner. The widely-cleft, shell-less test is supported
upon a thick peduncle, which is immersed in the skin
of the Shark. The surface of the peduncle is beset
with much-ramified, hollow filaments, which " penetrate
the Shark's flesh like roots " (Darwin). Darwin looked
in vain for cement-glands and cement. It seems to me
hardly doubtful, that the ramified hollow^ filaments are
themselves nothing but the cement-ducts converted into
nutritive roots, and that it is just in consequence of the
development of this new source of nourishment, that
the cirri and buccal organs are in the highest degree
aborted. All the parts of the mouth are extremely

,; n Y

■>;;:---n^^'

140 HISTORY OF CRUSTACEA. Chap. XII.

minute; tlie palpi and exterior maxillse have almost
disappeared ; the cirri are thick, inarticulate, and desti-
tute of bristles ; and the muscles both of the mouth and
cirri are without transverse striation. Darwin found
the stomach perfectly empty in the animal examined
by him.

Having reached the Nauplius, the extreme outpost of
the class, retiring furthest into the gray mist of primi-
tive time, we naturally look round us to see whether
ways may not be descried thence towards other border-
ing regions. By the structure of the abdomen in
Nauplius we might be remiuded, like Oscar Schmidt,
of the moveable caudal fork of the Eotatoria, which
many regard as near allies of the Crustacea, or at any
rate of the Arthropoda ; in the six feet surrounding
the mouth we might imagine an originally radiate
structure, and so forth. But I can see nothing certain.
Even towards the nearer provinces of the Myriopoda
and Arachnida I can find no bridge. For the Insecta
alone, the development of the Malacostraca may per-
haps present a point of union. Like many Zoese, the
Insecta possess three pairs of limbs serving for the
reception of nourishment, and three pairs serving for
locomotion ; like the Zoeae they have an abdomen with-
out appendages ; as in all Zoece the mandibles in
Insects are destitute of palpi. Certainly but little in
common, compared with the much wliich distinguishes
these two animal-forms. Nevertheless the supposition
that the Insecta had for their common ancestor a Zoea

Chap. XII. ^ CONCLUSION. 141

which raised itself into a life on land, may be recom-
mended for further examination.

Much in what has been adduced above may be erro-
neous, many an interpretation may have failed, and
many a fact may not have been placed in its proper
light. But in one thing, I hope, I have succeeded, — in
convincing unjprejudiced readers, that Darwin's theory
furnishes the key of intelligibility for the develop-
mental history of the Crustacea, as for so many other
facts inexplicable without it. The deficiencies of this
attempt, therefore, must not be laid to the charge of
the plan drawn out by the sure hand of the master, but
solely to the clumsiness of the workman, who did not
know how to find the proper place for every portion of
his material.

INDEX.

ACANTHONOTUS.

ENTOXISCUS.

AcANTHONOTrs Owenü, 12.
AcANTHOSOMA, 64 note.
AcH^us, 51, 53.

ACHTHERES, 131.

percarum, 87, 96.

Allorghestes, 29, 75 note.
Alpheüs, 49, 59, 125.
Amphilochus, 11, 75 note.
Amphipoda, 15, 39, 69, 74, 131,
Amphitiioe, 11, 78 note.
Anceus, 73.

Anelasma squalicola, 139.
Anilocra, 41.
Aratus, 8.

Pisonii, 31.

Artemia, 83 note.
Asellvs, 71.
Atyll-s, 75 note.
carinatus, 12.

Batea, 75 note.
Bodotria, 20, 81.
BoPYRiD^, 71, 72.
BoPYRES, 72, 73.

Brachyscelüs, 40, 76, 78, 91 note,
98, 104, 127.

crusculum, 76 note.

Brachyüra, 130.
Branchiopoda, 83.

Calanid^, 107.
Caligus, 78 note.
Oapkella, 75 note, 76, 80.

attenuata, 41 note.

linearis, 40 note.

Carcinus msenas, 55.

Oaridina, 49.

Cassidina, 41, 71, 72, 75 note.

Cerapus, 11, 75 note, 79, 80.
Chaliml'S, 78 note.
Chondracanthus, 87,

CPITHAMALrS, 92.

Cirripedia, 88, 96, 105.
Cladocera, 83.

Copepoda, 20, 84, 96, 107, 123.
CoROPHiUM, 74, 75 note.

dentatum, 76,

CORYC^ID^, 107.

Crangon, 49.

Crayfish, 97.

Cryptoniscus plauarioides, 72, 74.

Cryptophialus, 133. '

minutus, 94.

CüMA, 81 note.

CüMACEA, 81.

CYCLOGRAPsrs, 25, 32, 50.
Cy'clopiu^, 107.
Cyclops, 84-86, 90, 103.
Cyclopsine, 91.
Cymothoa, 73.
Cyniothoadiens, 72.
Cytridina, 107.
Cyprts, 82, 107.
Cyrtofhium, 11, 75 note, 104.
Cythere, 107.

Daphnia piüex, 75.

Dercothoe, 79.

DlASTYLID^, 20, 81.

Dichelaspis Warwickii, 136.
DuLiCHiA, 75. 76, 104.

Edriophthalma, 15, 39, 69.
Entomostraca, 83.
Entoniscus, 72.

cancrorum, 41, 72, 74.

porcellanffi, 41, 73, 74, 124 note.

ERICHTHUS.

INDEX.

POKTELLIDiE.

143

Eeichthus, 66.
Eeiphia gonagra, 8, 83.
EuFHAisiA, 64, 66, 98, 127, 128.

EURYNOME, 51.
EVADNE, 75.

FiLOGRANA, 112.

Gammarus, 75 7iote.

ambulans, 75.

Dugesii, 28.

piiteanus, 39.

Gecarcinus, 47, 48.
Gelasimis, 8, 19, 25, 36, 49.

vocans, 36.

Glaxjcothoe Peronii, 56.
Geapst:s, 31, 35.

Hermit Crabs, 49, 54, 59, 125.
HipPA emerita, 49, 54.
HipPOLYTE, 49, 59, 125.
Hyperia galba, 76 note.

Latreillei, 78 note.

Martinezii, 77 7iote.

^^ Hypcrines anormales et ordi-
naires," 40 note, 79.

Idothea, 71, 72.
Insecta, 119 7wte.
IsopoDA, 15, 39, 43, 69, 131.

Kepone, 73.

Ljjmodipoda, 39.
Lepas, 89, 91.

anatifera, 44.

anstralis, 93 note, 136.

LERN^omscrs porcellause, 93 note.
Lernanthropis, 87.
Lestrigonus, 78 note, 79.
Leucifer, 66, 71 note, 107, 123 note.
Leucothoe, 11, 75 note.
LiGiA, 69, 75 note, 103. 127.
Lobster, 48, 98.
LuPEA diacantha, 34 note.

Macrüra, 49, 130.
Maia, 51, 53.
Megalops, 125.
Melita, 75 note.

anisochir, 9.

exilii, 9, 11, 28.

Fresnelii, 9, 10, 11.

insatiabilis, 27, 28.

Messalina, 27, 28.

palmata, 11, 28.

setipes, 9.

valida, 9.

IMiCRODEETOPUS, 14 note, 75 note.

MoNTAGUA, 75 note.

Mysis, 64, 69, 98, 103, 107, 127, 130.

" NArPLiis-LARv.5i:," 13, 14, 17, 58.

82, 84, 86, 89, 96 note, 122, 124.
Nebalia, 83.
NlPHARGUS, 39.

OcYPODA, 8, 33, 36, 49.

rhombea, 34.

Orchestia, 75 note, 79, 104.

Darwinii, 25, 26.

gryphus, 25.

sylvicola, 25, 27.

tahitensis, 25.

telhiris, 25, 27.

Tucurauna, 79. .

Tucuratinga, 80.

Orchestoiijea, 75 note.

PAGURrs, 127, 130.
PALJi:MON. 49, 56, 59, 98, 125, 130.
Palinurus, 56, 98, 123, 128.
Peltogastee, 95.

socialis, 94.

Peneus, 17, 98, 125, 128.

setiferus, 64 note.

Persephone, 126 note.
PhiloscIa, 71, 75 note,'J.27.
Phronima, 39, 98.

sedentaria, 77.

Phryxus, 73.
Phyllopoda, 83.
Phyllosoma, 57, 66.
Pinnotheres, 52.
Podofhthalma, 47.

144

PORCELLANA.

INDEX.

ZOE^.

Polyphemus, 103.
pontellid^, 107.

PoRCELLANA, 49, 59, 60, 102, 125,
127.

stellicola, 53, 126.

porcellionldes, 72.
Praniza, 73.
Prawns, 55-64, 123.
Protella, 75 note.
Protula, 112.
Pycnogonid^, 96 note.

Kanina, 8, 30.

Khizocephala, 88, 93-96, 105, 135.

Sacculina purpurea, 89, 90, 92, 94,

95.
SCALPELLUM Peioiiii, 136.
Sergestes, 64 note.

Serpul^, 112.
Sesarma, 8, 25, 32, 52.
SI ir imps, 55.
Sph^roma, 72.
Squilla, 43, 49, 66, 125.

Taliteus, 79.

Tanais, 41, 43, 71, 73, 104, 128.

dubius, 21.

Dulongii, 16, 19.

Tatuira, 49, 53, 59, 125.
Teteaclita porosa, 88, 95.
Trilohites, 96 note.

Xantho, 52.
XiPHOSURA, 96 note.

Z0EJ3, 15, 17, 44, 48, 49-68, 122.
124, 128.

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