National Antarctic Expedition V6, 1907-1904

NATIONAL ANTARCTIC EXPEDITION
1901-1904

NATURAL HISTORY

Vol. VI.

ZOOLOGY AND BOTANY

LONDON
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
1912
{All Rights Reserved)


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;


PREFACE TO VOL.
With

Volume VI

the issue of

VI.

the Eeport on the Natural History specimens collected

by the 'Discovery' National Antarctic Expedition (1901-4)
present at least, to a close

V

Volume

Volumes I-III appeared

;

1907,

brought,

Volume IV

for

in

the

1908,

in 1910.

The present volume was
specialist to

completing

in

is

whom
it

as

to

have included

an account

of

the

Polyzoa

;

the

the specimens were submitted has, however, been prevented from

soon as he had hoped, and

without the paper in

it

was decided to

issue

the volume

question rather than delay further the publication of the

rest of the contents.

The thanks of

naturalists

careful study of the material

are

due to the numerous workers who, by their

and by

their detailed descriptions,

knowledge of the Animals, Plants and Minerals found

A

word of acknowledgment

is

also

have added to our

in a part of the Antarctic region.

due to the sub-editor of the Zoological and

Botanical Reports, Mr. F. Jeffrey Bell, for the care and judgment with which he has

done the work entrusted to him more than seven years ago

;

during that space of

time he has been charged with the distribution of the zoological and botanical
specimens to the various investigators,
authors,

obtained the reports from the respective

prepared the manuscript for the printers, revised the proof sheets,

supervised the preparation of the numerous plates and other illustrations.

L.

FLETCHER,

Director.

British

Museum (Natueal
February 2dth, 1912.

History),

and



—

CONTENTS OF VOL.
A

Generai; Notice of the Biological Memoirs

VI.

.

.

.

Synopsis of the Contents of the Several Volumes of Memoirs.
Classified

Summary

of the Contents of Vols. I-VI

.

.

(p- vii.)

•

.

(p. xi.)

(p- xiii.)

ECHINODERMA.
III.

On a Collection

of

Young Holothurioids.

By

Prof. E.

W.

MaoBride, D.Sc, LL.D., F.R.S

(9 pp., 2 Pis.)

POLYCH^TA.
By

E.

Ehlers, Professor in Gottingen

FEESHWATEE
By

F.

E. Fritsch,

.

.

(32 pp., 3 Pis.)

.

.

(GO pp., 3 Pis.)

ALGiE.

D.Sc, Ph.D., F.L.S.



—

A GENERAL NOTICE OF THE BIOLOGICAL
MEMOIES.
The completion

of these

memoirs, descriptive of the fauna and

fifty

part of the Antarctic area which was visited by the

command

Discovery,' under the

'

of that

flora

of Captain Scott, E.N., C.V.O., oS"ers a suitable opportunity for saying a few words

on

the

and

that were obtained,

collections

some general

for suggesting

reflections

on the inhabitants of the area that was studied.

we have

First of all

and the South Polar regions
solid

animals

defenceless

which they
to assume

hungry

other,

splendid and

such

in

absence

the

polar

sea,

places

are

aggressors

which

allows
are

other

compelled

and the

as themselves

of

the

in

dominated by

have

climes,

colourations

beautiful

the

is

of its solid

same colour

assume the

In the

live.

the

to

surfaces

ever

Carnivores which,

rapacious

one there

the

in

the one case, the

In

land.

;

between the North

extraordinary difi'erences

note the

to

so

all

snow on
Penguins

the

shown

well

in

Dr. Wilson's admirable drawings.

The

region,

Antarctic

inhospitable climate, almost

devoid of

forms are described in these volumes.

"It was

of

the collection
represented

on the

usual

the

quite

the

'

to

thing

Discovery

being,

of

instead

'

life,

will

might

with

imagine,

teems with species, of which 227

its

new

Of some Amphipoda Mr. Hodgson writes
take

10,000

Schizopoda

by nearly 10,000 specimens.

Polychseta

we

as

to

30,000

contained

Students

of

at

one

a

species

Professor

And

haul."

which was

Ehlers'

report

have only a faint idea of the number of specimens

of

Harmothoe spinosa brought back by the Expedition.

The Alga Lessonia grandiflora was found
24 feet long

;

Mosses showed
existence.

to

have

a

lamina

as

much

as

on the other hand, M. Cardot noted that most of the species of
signs,

With

the

which he has
exception

of

detailed, of

some

obtained by the 'Discovery' Expedition,

the severity

Algae,

no

of their struggle

freshwater

organisms

for

were


A GENEKAL NOTICE OF THE BIOLOGICAL MEMOIRS.

viu

On

parting with these reports,

on what has

most

of all there

First

by Mr. Hodgson and Dr.

duties

appropriate

say

to

a few words

and studious attention that

me, in the prolonged

struck

have given to them.

may be

it

stands out the observant care

of their

always of

Koettlitz as collectors, under conditions

discomfort, and often of danger.

I

In the second place, one cannot but be impressed

with the varied attainments that Dr. Wilson brought to the tasks that he had to

perform

memoirs

of his

;

intelligent

lover

pencil,

Antarctic regions a necessity for

to the

visit

their

solution.

drawings added considerably to the attractiveness of Volume

his

not enter at any greater length into Dr.

I shall

spirit of the

not only a naturalist, but he also possesses great power over brush and

is

and

be said that they breathe the true

and they raised a number of interesting questions

animals,

of

which made another
Dr. Wilson

may

it

Wilson's

observations

11.

on Birds,

any zoologist who has not yet read them should make himself acquainted with

for

them

The same

at once.

which were brought home
Morphological

observations

slits

expected at

of

seals,

and the account of two new
gill

Seals,

the skins

of

in excellent condition.

memoirs on the embryos of

chord and

Whales and

report on

true of his

is

considerable

are

interest

be found

to

the

in

the development of the feathers of the penguins,

species of

possible

indicates

a

should

also

whose possession of a noto-

Ceplialodiscus,
affinity

to

Man

that

would

not be

first sight.

attention

Special

be called

to

the

two reports

Echinoderm

on

Larvae which open up several points of general interest.

Turning to the more

new

23

species,

among
so

species

much

are

I

have to note that two

and 26 of

alone

judge,

I

may

as one containing
it

can

plants.

and

fell

into

I

my

even

As

to

these,

new

by some

;

there

appear

the validity
as

the

of

we know,

to

differ

forms, but as one which, from the very conditions
offer

congratulate myself, at any

hands,

for

I

an admirable opportunity for
rate, that the

tremble to think

how many

other workers would have made, as a result of the bewildering

exhibited

In the zoological

suggest that another collection should be treated, not

would be brought together, must

study of variation.

Echinoderms

but none among plants

proposed,

of animals,

specialists

themselves.

under which
the

new genera

new

201

new

systematic memoirs

sub-families have been created, both for Calcareous Sponges.

reports

lie

strictly

of the forms.

amount

collection

species

of

some

of variation


";

A GENEKAL NOTICE OP THE BIOLOGICAL MEMOIES.
On

whole

the

the

naturalists,

K.C.M.G., who have

Eliot,

been
Poles,

and

with the possible exception of

contributed

these

to

by any resemblance between

struck
it

ix

do not appear to have

reports,

faunas

the

and South

the North

of

was therefore with some astonishment that

Charles

Sir

I received

late

last

year

of " bipolarity

an important memoir, by Professor Theel, urging the

evidences

manifested by the Priapulids and

by the Swedish Antarctic

To

Expedition.

may

It

be remembered that one of the

was a curious

some time

had
the

before

been discovered

was found

person

many

years ago

name

quite unintelligible

Land has ten

Victoria

of the lacunar

illustration

a

legs,

common genus Nymphon,

of

may

draw

first

it

while

sea

attention.

objects of interest in the whole

found by Mr. Hodgson

spider

knowledge of zoologists that

who knew

that

ten-legged sea

a

was

it

spider

by Mr. Eights, and described by him under

Decolopoda.
differs

Eights's

was found by the Charcot Expedition
It

collected

of course, I can here only

this,

was the ten-legged Pycnogonid or

collection
it

Sipunculids

in

although

But,

no other

genus,

which

the

important

from

form
point

had marked

off

from the

peculiarities,

in another part of the Antarctic.

be observed that there are striking differences between the fauna of

the area explored by English and that examined by French, German, and Swedish
navigators, but the time for an explanation of these has hardly yet come.

Most
to

had

of the

zoologists

the

or

contributors
botanists

Common

is

it

the biological volumes bear names

has been

introducing

opportunity of

Mr. C. F. Jenkin, who

;

to

now

to

to

me an

zoological

especial

well

known

privilege that I

work and

zoological

workers

Professor of Engineering in the University of Oxford.

treatment has been followed in the nomenclature of both plants and

animals.
F.

British

have

Museum (Natural

History),

Department of Zoology.

JEFFKEY BELL.



NATURAL HISTORY
OF THE

NATIONAL ANTARCTIC EXPEDITION,
190M904.

SYNOPSIS OF THE CONTENTS OF THE
SEVEEAL VOLUMES OF MEMOIES.
Volume L
Field Geology.

.

Rock Specimens

.

Volume

.....

Mammalia (Whales and
AVES

Seals)

.

By H.
By G.

T.

Ferrar, M.A., F.G.S.

T. Prior,

M.A., D.Sc, F.G.S.

II.

By Edward
By Edward

A. Wilson, M.B.
A. Wilson, M.B.

On some

Points in the Anatomy of
the Emperor and Adelie Penguins

Pisces

Cephalodiscus

.

Cephalopoda
Gastropoda
Nudibranchiata
Amphineura
Lamellibranchiata
Brachiopoda
Decapoda
.

Cumacea

.

Volume

On Collecting
Pteropoda
Pycnogonida
Acari
Amphipoda
Leptostraca
Ostracoda
CiRRIPEDIA

CHiETOGNATHA

Nbmatoda

in

Antarctic Seas

By
By
By
By
By
By
By
By
By
By
By

W.

Pycraft.
Boulenger, F.R.S.
W. G. Ride wood, D.Su.
W. E. HoYLE, D.Sc, M.A.
P.

G. A.

Edgar A. Smith,

I.S.O.

Sir Charles Eliot, K.C.M.G. LL.D.
Edgar A. Smith, I.S.O.

ISO.

Edgar
Edgar

A. Smith,

W.

T.

W.

T.

Calman, D.Sc.
Calman, D.Sc.

A. Smith, I.S.O.

III.

By T. V. Hodgson, F.L.S.
By Sir Charles Eliot, K.C.M.G., LL.D.
By T. V. Hodgson, F.L.S.
By Dr. E. L. Trouessart.
By A. 0. Walker, F.L.S.
By Dr. J. Thiele.
By Prof. G. S. Brady, F.R.S.
By Prof. A. Gruvel.
By Dr. G. Herbert Fowler.
By Dr. 0. von Linstow.
h 2


,

xu

SYNOPSIS OF THE CONTENTS OF

Volume

/

Cestoda
Alcyonaria
Hydroid Zoophytes

THE SEVERAL VOLUMES OF MEMOIES.
III [continued).

By Arthur E. Shipley, F.R.S.
By Prof. S. J. Hickson, F.R.S.
By Prof. S. J. Hickson, F.R.S.

.

.

F.

....
Alg^

AND

(Ph^ophyue.^!;

By A. and E.
By M. Foslie.

FLORIDEiE)

y Marine Algje (Corallinace^)

Volume
/ solenogastres

Aptera

H. Gravely.

By Dr. J. Rennie.
By R. KiRKPATRICK.
By Jules Cardot.

Tentacles of a Siphonophore
Hexactinellida
Musci

Marine

Dr. H.

Prof.
M.R.I.A.

ECHINODERMA
ECHINODERM LaRV^
.

and

Actiniae

\

.

.

Nemertinea
MedUS/E
.

.

Volume
/

A

.....

General Notice of the Biological
Memoirs
Synopsis of the Contents of the
several Volumes of Memoirs.
Classified Summary of the Contents
OF Vols. I-VI.
On a Collection
thurioids

of

Young Holo-I

.....
....

.

\

H.

Carpenter,

B.Sc.,

J.

C.

Simpson, B.Sc.

V.

By Dr. H. W. Marett Tims.
By Prof. W. A. Herdman, D.Sc, F.R.S.
By T. V. Hodgson, F.L.S.
By Prof. L. Joubin.

By Edward T. Browne.
By Dr. Otto Vernon Darbishire.

Seal-Embryos
tunicata

LiCHENES

Nierstrasz.

Traunfels.
J. A. Clubb, M.Sc.
R. KiRKPATRICK.
C. F. Jenkin, B.A.

.

,

G.

By
By
By

.

Tetraxonida
Calcarea

ISOPODA

F.

By W. F. Lanchester, M.4.
By Dr. Rudolf Ritter v. Stummer-

.

.

Volume
(

Gepp.

By W. M. Tattersall, M.Sc.
By R. NOKRIS WOLFENDEN, M.D.
By F. Jeffrey Bell, M.A.
By Prof. E. W. MagBride, M.A., F.R.S.,

sohizopoda
CbPEPODA

SiPUNCULOIDEA

S.

IV.

By
By

.

.

Myzostomid^

and

Polych^ta
Freshwater kLGM

.

.

.

)

VI.

By

F.

By

Prof. E.

Jeffrey Bell, M.A.

W. MacBride,

D:Sc.,

LL.D

F.RS.

By
By

E. Ehlers, Professor
F. E. Fritsch,

iii

Gottingen.

D.Sc, Ph.D., F.L.S.


SUMMARY OP THE

CLASSIFIED

CONTENTS OF VOLS.

On Collecting in Antarctic

}

>

Seas

By m
T.

I-VI.

ttito
-a
V. Hodgson, F.L.S.

ttith
Vol. HI.

^t

-d

.

.

.

ZOOLOGY.
VERTEBRATA.

^^™s)^

^^^'''^'^'''

Seal-Embryos

AvES

.

^""^
I

.

.

-By Dr.

.

.

.

^

g

...
.

„

n.

13

A. Wilson, M.B.

"j

PEROR AND AdBLIE PeN-

Pisces

^- ^^^'°^' ^•^-

H. W. Marett Tims

By Edward

.

.

On some Points in the

Anatomy of tbe EmGUINS

^y ^"^^^^°

I

p

p^.^^^^^^

-^

J
.

By

G. A. Boulenger, F.R.S.

TUNICATA.
By

Prof.

W.

A.

Hbrdman, D.Sc,

F.R.S.

.

.

„

V.

.

.

„

II.

PTEROBRANCHIA.
Cephalodiscus

.

.

.

By W.

G.

Ridewood, D.Sc.

.


XIV

CLASSIFIED

SUMMARY OF THE CONTENTS OF

VOLS. I-VI.

MOLLUSCA.
Cephalopoda

By W.

Gastropoda

By Edgar

Pteropoda

By

Sir Charles Eliot, K.C.M.G., LL.D.

nudibranchiata

By

Sir Charles Eliot, K.C.M.G., LL.D.

Amphineura

By Edgar

A. Smith, I.S.O.

By De. H.

F.

By Edgar

A. Smith, I.S.O.

solenogastres

.

Lamellibranchiata

E.

HoYLE, D.Sc, M.A.
A. Smith, I.S.O.

Nierstrasz

.

.

.

.

.

.

.

Vol. II.

„

II.

.

„

III.

.

„

II.

.

„

II.

„

IV.

„

II.

.

.

.

.

.

.

.

BKACHIOPODA.
By Edgar

A. Smith, I.S.O.

.

II.


CLASSIFIED

SUMMARY OF THE CONTENTS OF

XV

VOLS. I-VI.

ECHINODEKMA.
ECHINODERMA

By

F.

Jeffrey Bell, M.A.

.

I

of

HOLOTHURIOIDS

.

.

(By
By Prof.E.
Prof. E. W. MacBride, M.A., F.R.S.
F.R.S.,(
J
and J. C. Simpson, B.Sc.
-j

ECHINODERM LarvJE

On a Collection

.

Young (By Prof.
.

.

E.

F.R.S.

\

.

W. MacBride, D.Sc,

.

.

.

.

Vol. IV.

"

LL.D.,)
-j

.

jy

"

VI.

„

VI.

„

IV.

„

IV.

„

III.

„

III.

„

V.

„

III.

POLYCH^TA.
By

Prof. E. Ehlers

SIPUNCULOIDEA.
By W.

Lanchester, M.A.

F.

MYZOSTOMID^.
By Dr. Rudolf Ritter

v.

Stummer-Traunfels

CHiETOGNATHA.
By Dr.

G.

Herbert Fowler

NEMATODA.
By

Dr. 0. von Linstow

....

NEMERTINEA.
By

Prof. L. Joubin

.....

CESTODA.
By Ajrthur

E. Shipley, F.R.S.

.

COILENTERA.
By

Alcyonaria

Actinia

.

.

.

Hickson, F.R.S.

III.

....

Tentacles of a Siphonophore
.

S. J.

(By Prof. S. J. Hickson,
Gravely

Hydroid Zoophytes

Medusa

Prof.

.

.

By

Dr.

J.

Rennie

.By Edward T.
.

By

J.

F.R.S.,

A. Clubb, M.Sc.

F.

H

III.

III.

.

Browne

and

.

.

V.
IV.


XVI

CLASSIFIED

SUMMAET OF THE CONTENTS OF

VOLS. I-VI.

POEIFERA.
By

R.

Tetraxonida

By

R. Kirkpatrick

Calcarea

By

C. F.

HEXACriNELLIDA

.

.

Kirkpatrick

Jenkin, B.A.

Vol. HI.

„

IV.

„

rv.

„

HI.

„

HI.

„

HI.

„

VI.

BOTANY.
MUSCI.

By Jules

C'ardot

MARINE ALGM.
By

Pn^oPHYCEyE AND Floride^
CORALLINACEiE

.

.

A.

-By M.

and

E. S.

FoSLIE

FRESHWATER
By

Gepp

F. E. Fritsch,

ALGiE.

D.Sc, Ph.D., F.L.S.

LICHENES.
By

Dr. Otto Vernon Darbishire

,.

V.

„

I.

„

I.

GEOLOGY AND MINERALOGY.
Field-Geology
Rock-Specimens

.

.

.

.

.

By H.
By

T.

Ferrar, M.A., F.G.S.

G. T. Prior, M. A., D.Sc, F.G.S.


ECHINODEEMA.
A COLLECTION OF YOUNG
HOLOTHUBIOIDS.

III.-ON

By Professor

W. MacBride, D.Sc, LL.D.,

E.

F.R.S.,

Imperial College of Science.
(2 Plates.)

me

In 1908 Professor Jeffrey Bell asked

to report on a collection of

young Bchino-

derms, which had been collected by the National Antarctic Expedition.

came

to hand, they proved to be the post-larval stages of a Holothurioid.

was added a specimen of the Auricularia

collection

first

When

to

larva, of

be reported from Arctic or Antarctic waters.

these

To

this

some Holothurioid, the

In an earlier report (5) by

Mr. Simpson and myself on the Echinoderm larvse of the Antarctic Expedition
described

the

for

time

first

the

occurrence

of

free-swimming

the

We

Echinoidea and Ophiuroidea in Antarctic waters.

now

can

larvae

we
of

assert the existence

of three out of the four types of free-swimming Echinoderm larvse in these waters.

This

is

important in

Echinoderms

in

view of

Arctic

embryonic type without

the

opinion

free larvae.

I

shall,

specimen, and then detail the results of

were

which has

been

expressed

that

all

and Antarctic waters had developments of the shortened

my

first

of

consider

all,

work on the

this interesting

post-larval stages which

contained in the collection.

Auricularia Antarctica.

I.

(Plate

unique

This

specimen

is

fig.

distinguished

wheel-shaped calcareous bodies which
the body,

I.,

it

1.)

above

contains.

but are perhaps most numerous

all

number

by the large

These are distributed

in the anal "

field."

Each

all

of

over

consists of

a concave bowl, the sides of which are composed of 11 to 13 "spokes" connected

by a

rim.

concavity

is

seen to

At
(fig.

the bottom of the bowl
2).

Where

consist of a

bodies, but with a larger
digitata,

but

this larva

a view can

coarse

the

hub which

projects slightly into the

be obtained from the convex side

network of calcareous substance.

number of
differs

is

the hub

Similar calcareous

spokes, are described from the larva of Synapta

from that under consideration

in

the

fact that its


"

2

E.

"wheels" are few

W. MacBRIDB.

number and confined

in

the lateral processes,

to

Numerous

whereas in our larva they are extremely numerous.

13-16 spokes are described by
fished

up

at Orotava at the

Canary

by Dr. Mortensen

nudibranchiata

A

Prof.

Chun

in a peculiar Auricularia

(2)

As

all

with

which he

and which has been named Auricularia

Islands,

(8).

few words upon the general anatomy of the Auricularia larva

out of place here.

1-6 in each,

" wheels "

zoologists are aware, it possesses, like other

may

not be

Echinoderm

thickened band of ciliated ectoderm as locomotor organ, and this band has

larvae, a

the form of a folded loop, the longest axis of which
the larva

;

in a word, the loop has

two long

posterior cross-pieces connecting them.

The

is

parallel to the long axis of

parallel sides

and shorter anterior and

anterior cross-piece

is

folded backwards, so

as to form a frontal loop surrounding the forehead or " frontal field," whilst the
posterior cross-piece is folded forwards so as to surround an " anal field " in which
the anus opens.
This loop may be termed the anal loop. The adjacent portions of the
frontal and anal loops are termed by Dr. Mortensen (" Quersaume "), which we may
translate as anterior and posterior " transverse bars."
The mouth is situated in a

depression between the anterior and posterior bars termed the oral field.

band, in addition to the. re-entrant frontal and anal loops,

which

"processes"
Ophiopluteus
sides

the

of

anal loop.
are

are

homologous

with

loop

frontal

produced into a number of

of

the

Echinopluteus

and

Where the

;

frontal loop passes into the sides of the ciliated band, there

developed the antero-dorsal processes.

From

same

these

lateral

portions of

back intermediate-dorsal and postero-dorsal

further

Finally, where the anal loop passes into the lateral portions of the band,

we have

processes.

Miiller,

to

the postcro-lateral
showed a resemblance

was coined to designate the

Wc

ciliated

Of these the prse-oral processes are developed from the
and the post-oral processes from the sides of the

larvae.

the band are developed
processes.

arms

the

is

The

the

These

human

in the opinion of Johannes

whence the name " Auricularia

last,

ear,

larva.

have seen that A. antarctica cannot be identified with the larva of Sj/napta

digitata, since in the latter the "

processes (1-6 in each).

wheels

But there

"

are few in

number and

are confined to the

are other differences scarcely less striking.

larva oiSynapta digitata the outline of the ciliated

band

is

In the

flowingly sinuous, none of the

processes being very strongly marked, whereas in our larva the processes are

by deep re-entrant

folds

and show some secondary

frontal loop passes into the lateral portions of the ciliated
in the larva of

dorsal surface.

our larva

it is

Synapta

The

digitata,

but in our larva

oral field is comparatively

reduced to a narrow

the posterior bar at the sides.
larva not only in the great

slit,

On

number

marked off"
Then the spot where the
band is at the anterior pole

plications.

it is

displaced far back on to the

broad in the larva of Synapta, but in

the anterior transverse bar actually overlapping

the other hand our larva resembles the Orotava
and wide distribution of the " wheels," but in the

displacement of the point of union of the lateral part of the ciliated band with the


ECHINODEEMA.
frontal loop on to the dorsal surface,

and

3

narrow

also in the

nevertheless, irreconcilable differences between the

the Orotava larva in

its lateral

so that the whole animal

name A.

the

portions

is

But there

oral field.

The

two forms.

band of

ciliated

produced into numerous secondary processes,

whence

acquires the appearance of a Nudibranch mollusc,

nudibranchiata.

Further,

are,

in

this

larva the

absent, but these are well

marked (Plate I., fig.
primary processes are deeply marked there are no

post-oral

1) in our larva,

are

processes

and though the

tag-like secondary processes such

as exist in A. nudibranchiata.

In spite of
the

for

its

coelom

considerable size (4
in the

mm.

long) our

specimen

form of a single unpaired

is

a

young

larva,

communicating with the exterior by a pore-canal.
Reviewing all the evidence, we arrive at the conclusion that our larva is a
new type of Auricularia, to which the specific name "antarctica" may be given.
It

is

still

vesicle {coe.,

fig.

1)

probably belongs to some large Holothurioid of the group Synaptidae.

II.

Post-larval Stages of Cucumaria

The specimens composing
contraction

that their shape

this collection

sp.

were mostly in a state of such intense

approximated to that of a sphere, but a few were

considerably less contracted, and of these sections and whole mounts were made.

In

PI.

I.,

fig 3,

one of the most successful of the whole mounts

be seen that at the oral end of the body there
recalling the vestibule of a

young

Crinoid.

Of

is

is

shown.

an atrium overarched by

It will

five valves,

In the intervals of the valves glimpses of

we learn from transverse

The suckers of tube-feet can be seen to be developed along three radii.
But the most striking feature in the preparation is the presence of numerous
calcareous plates embedded in the -skin, so numerous indeed as to constitute a veritable
the buccal tentacles can be seen.

these there are ten, as

sections.

cuirass, especially

Each

oral valve

is

over the dorsal surface; the plates do not touch edge to edge but overlap.

supported by a special plate.

Between the rows of tube-feet there

are also calcareous bodies to be found, but these are not fully formed plates.

the semi-transparent tissues the outline of the alimentary canal can just be

and we can observe the thick oesophagus, the long stomach, and the

Through

made

out,

slightly bent

intestine.

Now
Cucumaria

the

identification

rests

(i)

of these specimens

as

the

on the number and shape of the buccal

of the calcareous plates

embedded

young of some
tentacles, (ii)

species

of

on the nature

in the skin.

With regard to the first point, Cucumaria is characterised by the possession of ten
" dendrochirote
buccal tentacles, i.e., tentacles which exhibit scattered lateral
'"

branches.

Further, these tentacles are devoid of specially developed ampullae, but on

the contrary the whole tentacle ring can be retracted as an introvert.

With regard
B 2

to the

4

W. MacBRIDE.

E.

second point, the occurrence of " Gitterplatten

common

" (lattice-plates) as calcareous

Now,

occurrence in the genus Cucumaria.

number

in the

bodies

is

of

of tentacles, the

presence of an introvert, and the shape of the calcareous bodies our specimens resemble

Cucumaria.

In the presence of tube-feet on only three

[and Colochirus, F.J.B.], but in Psolus the body

form a creeping

But

Cucumaria.
is

sole,

young Cucumaria

obtained at Plymouth in 1905, one of which
that the tube-feet are not developed on

Ludwig

—probably

shown

is

for a considerable

I

find

median ventral water-vascular canal and remain
These two terminal tube-feet are clearly

in transverse sections of our specimens (PL

11.

,

fig.

8c).

show

then, belong to the genus Cucumaria, but they

They

obvious traces of the characteristic pulmonary trees.

mode

which

Here too we

4.

fig.

laotea,

the interradii, but are confined to two.

all

time the only tube-feet.

Our specimens,
in their

in

C.

genus

(4) notes that in the development of C. planci a pair of tube-feet

are formed at the termination of the

marked

almost cylindrical, as in

is

the strongest evidence of their relationship to this

seen in their resemblance to the

Prof.

they resemble Psolus

flattened on one side so as to

whereas in our specimens the body

me

to

is

radii,

of obtaining food, for the stomach

is filled

are,

as yet

no

however, quite adult

with the half-digested remains

of Algse.
It

would seem that a

similar

stage

occurs

in

the development of Stichopus

Professor Mitsukuri (8) has found that the just metamorphosed form has
" a coat of armour," consisting of " disked tables with tall spires united by several

japonious.

cross-beams, which cover the body thickly, their bases even overlapping one another

when slightly contracted.
PL L, fig. 5 shows a

The ventral

course of the alimentary canal
involved.

No

pedicels were in three rows."

longitudinal sagittal section through a specimen in which the
is

and

quite straight,

which both mouth and anus are

endeavour has been made to represent the histology, but the general

The mouth

relationships of the organs are clearly shown.

surrounded by a very thick layer of muscles.
filled as

in

The stomach

noted above with Algae.

the thin-walled cloaca which

is

strands traversing the coelom.

might be older stages

in the

On

leads into an oesophagus

this follows a

long conical stomach

leads through a short intestine into

attached to the sides of the body-wall by muscular
I

was at

first

inclined to think that our specimens

development of Cucumaria

have been described by Mr. Simpson and myself

crocea, the

embryos of which

But this cannot be the case, for
is already folded.
The stone-canal ends blindly
in
in a thin- walled sac {ax., fig. 5) embedded in the body-wall
there is no pore-canal
leading from this sac towards the exterior such as was described by Mr. Simpson
(6).

these embryos the alimentary canal

;

and myself
nor

is

in

the case of the embryos of Cucumaria crocea in an earlier report,

there as yet

adult Holothurians.

any communication with the body-cavity such

The genital base (PL
genital organs,

may

as exists in the

This sac corresponds to the axial sinus of other Echinoderms.
I., g.

b.,

fig.

5),

from which spring the rudimentary

be seen close to the stone-canal, but there

is

as yet

no genital


ECHINODERMA.
There

duct.

is

doubt that

little

of other Echinoderms.

there

this genital base corresponds to the genital stolon

In a transverse section through the region of the tentacles

to be seen in each of the other four interradii a mass of deeply staining

is

rounded

cells

(PI.

component

their

5

lumen.

Is

II.,

some

cells in

possible

it

The masses are attached to the body -wall, and
show a tendency to group themselves round a

8a).

fig.

that

cases

these

are

antimeres of the genital organs

vestigial

destined to be absorbed?

Our material does not allow us to answer this question,
shows neither the origin nor the fate of these structures.
In the body-wall
on the right hand side of the section may be seen sections of rudimentary tube-

as

it

The

feet.

the tube-foot, that

tip of

an invaginated cup {pod.

from the

From

II.)

— quite

distinct

from the outgrowth
end., figs. 5-8).

the radial nerve-cord proceeds a pedal nerve, accompanied, like

On

it

took

origin,

its

by an epineural

each side of the mouth sections

which are sections of the epineural ring
buccal tentacle on the right side
disc (or.)
is

the disc of sensory epithelium, appears as

is,

5-8, PI.

figs.

which forms the inner part of the organ (pod.

radial-canal,

from which
.

ect.,

on the

;

space.

nerve

the

of

(ep.),

the nerve

but there

ring
is

seen, outside

are

no periheemal

The

ring.

seen to be retracted into a pocket of the

is

left side a section of

one of the

of

oral

five valves of the oral vestibule

seen.

As we examine
made out, and

be

curvature into

similar sections through older specimens the

the

three

alimentary

loops

and the main portion of

straight

In

intestine.

the

latest

found

I

outgrowth of the rectum on one side {puL,

pulmonary trees and a

of the
is

the beginning of the

fig.

fig.

in

7)

acquires

adult.

the

characteristic

The stomach remains

due to the lengthening of the
the

may

a

collections,

thin-walled

be regarded as the rudiment

duct connects the axial sinus with the ccelom.

"secondary madrep>orite" so characteristic of

the

T/iis

adult (mad.,

6).

If

we turn

to transverse

surrounding the mouth.

through the

which

is

In

fig.

In each

five radii.

the epineural

sections

canal.

canal,

and

we

see

in

fig.

8a the ten buccal tentacles

8b, a section taken lower

we observe

Internal to

epithelium, the radial perihsemal canal.

down, we can see sections

a section of the nerve cord, external to

it

we

find a space lined

Internal to this

is

by a thin

flat

the radial water vascular

this is present in each radius, although only in the three ventral are sections

of tube-feet recognisable.

two

the

in

this curvature is

which

stage

gradually

canal

marked

so well

same features can

In the adult Psolus, according to Professor Ludwig

dorsal radial canals are absent.

When

sections to their tips they do not, as in Echinoidea, Asteroidea
in terminal

(3),

the

the radial canals are followed in successive

and Ophiuroidea, end

azygous tentacles, but terminate within the body-wall.

The most

distal tentacles are laterally placed

three radial canals

which bear tube-feet.

sents a section from the

same

series

outgrowths of the median of the

These are shown in

fig.

8c,

which repre-

taken near the posterior end of the body.

6

E.

When we
we

review

I think,

are,

tlie

account that

here given of these post-larval Holothurians,

is

confronted with a certain number of Echinoid features.

The almost complete

dorsal cuirass of overlapping plates certainly suggests the

corona of an Echinoid, especially

number

W. MacBEIDE.

if

we

recollect that, in the earliest Echinoids, the

was

of rows of plates in the corona

number and the

indefinite in

plates over-

lapped.

The

closed axial sinus

Some

investigators

is

another Echinoid feature.

have made an attempt to show that Holothurioidea are

From

widely separated from the rest of the Echinoderma.
digitata, the

out, it has

only Holothurian in which the

been argued that since in

life

the development of Synapta

form the primary evaginations of the water-

this

owe

vascular rudiment give rise to the buccal tentacles, whilst the radial canals
origin

to

evaginations

secondary

worked

history has been thoroughly

alternating

with

these,

the

radial

canals

But

Holothurioidea are not homologous with those of other Echinoderms.

their

of

in our

specimens the canals occupying the interior of the tentacles can be traced into contin-

The

that Synapta

is

about the worst form that

could have been chosen to represent the Holothurioidea.

Its radial water-vascular

uity with the radial canals.

fact

is

canals are only transitory larval structures,
directly

from the water- vascular ring, but,

and

The functional importance

the early development of Cucumaria planci

If

no longer a multiple of

five.

Professor Ludwig,

who worked out

(4), points out that in this form the first

spring from the radial canals.

we compare

form with the Auricularia which was

the youngest post-larval

described in the beginning of this paper,
is

is

which prevails in

of the buccal tentacles leads to their early appearance in

ontogeny, before the transitory radial canals appear.

tentacles

buccal tentacles not only spring

number

the other groups of Holothurioidea, their

all

its

in contravention of the rule

we

see that the alimentary canal in both

The long stomach obviously corresponds in each case, as does the
The short thick oesophagus of the post-larval form corresponds to the

very similar.

short intestine.

inner part of the oesophagus of the Auricularia, whilst the shallow outer part becomes

we know from the

(as

The

life-history of other

chief difiiculty in the

way

Auricularia) the oral vestibule.

of deriving Holothurioidea from a primitive form

of Echinoidea has lain in the difference between the Echinopluteus
larvse,

and the apparent retention by the

those shown

and

by the Echinopluteus.

I

latter of

and Auricularia
more primitive features than

have shown elsewhere (5) that the stomach

Echinopluteus are directly converted into the stomach and
young
Echinoid.
Further, in our specimens the stomach remains
intestine of the
unchanged in shape as growth proceeds, and the looping of the alimentary canal
is

the

intestine

of the

due solely to the growth
Echinopluteus.

oral disc of the

the

Auricularia

in

young Echinoid

—the

length

The " amniotic space
difference

is

of
"

the
in

intestine.

This

is

also

true

of

the EchinopAuteus inside which the

formed corresponds to the

oral

between the two cases being that

vestibule
in

the

of

latter


7

ECHINODBRMA.
form the
deserts

oral vestibule at the

original

its

whilst in

the

"cBsophagus"

median

moment

of metamorphosis, as Mr.

position

and moves to the

Echinopluteus the

formed on the

is

original

shown,

the larva,

of

side

left

(1) has

new

oesophagus disappears and a

larval

left

side of

the larva.

in

respect

of

more modified development, but

Bury

the

This

history

is

of

undoubtedly the
ccBlom

the

the

Echinopluteus has retained a far more primitive arrangement than the Auricularia.

The

fact

that the original

re-formed in Echinopluteus,

mouth
is

is

retained in Auricularia whilst

paralleled

by the

fact that the

lost

and

it

is

is

retained in

mouth

the Ophiopluteus larva, but lost and re-formed in the more primitive Bipinnaria larva,

yet no one would doubt that an Ophiuroid
in the primitive Echinoids

is

derived from an Asteroid.

Doubtless

from which the Holothurioidea diverged the larval mouth

was retained.
If,

then,

we

accept provisionally the

hypothesis

that

the Holothurioidea are

descended from primitive Echinoidea, a plausible physiological reason can be suggested
for their evolution.

roundings

and the

it will

If ordinary regular sea-urchins be studied in their natural sur-

be noticed that they frequent by preference crevices among rocks

vertical faces of rocks,

where the numerous tube feet borne by their long

Now

radial canals can take hold.

from habits of climbing the transition to habits of

wriggling through narrow crevices and of burrowing

is

easy,

and what

little

is

known

of the habits of Holothurioidea points to the conclusion that their normal habits are

of this kind.

I

have myself dug up Synapta inhaerens from

and Professor Mitsukuri

is

anatomy would appear
*

corona, and
to

in

the muscularity of the body-wall

thus the

distinctive

features

and

of Holothurioid

have been evolved.

Compare F. D. Bennett's account

" They usually

mud,

in the

made in order to collect
food and known as Trepang.*

one of the species used for

Such habits would require an increase
of the

burrows

(8) records that piles of stones are

Stichopus japonicus, which

a disappearance

its

of the habits of the Trepang,

'

Whaling Voyage

'

(1840),

i.,

p.

175

:

exposed in the shallow waters, though we have very often seen them buried in beds of
coral sand, their plumy tentacles being alone exposed, and floating in the water above, apparently as a lure
for prey. Some may also be observed lying on the rocks, their bodies completely encrusted with coral sand,
lie

which may either have been accumulated by a previous burrowing, or thus used

as a disguise."

— F.

J.

B.

8

E.

LIST OF

(1)
(2)

W. MacBEIDE.

WORKS REFERRED TO IN Prof. E. W. MacBRIDE'S REPORT
ON A COLLECTION OF YOUNG HOLOTHURIOIDS.

BuEY, H. The Metamorphoses of Echinoderms. Q. J. Micr. Sci.
Chun. Atlantis. Biologische Studien fiber pelagische Organismen.

Vol. 38 (1895).

Bibliotheca Zoologica.

Heft 19

(1896).

LuDWiG, H. Echinodermen in Bronn's Klassen und Ordnungen des Thierreichs. Bd. ii., Abt.
Buch 1. Die Seewalzen. 1889-1892.
Sitzungsber. d. Berliner Akad.
Zur Entwickelungsgeschicbte der Holotburien.

(4) LuDWiG, H.

(3)

3,

d.

Wissensch., 1891.
(5)

MacBride, B. W.

The development

of

Echinus esculentus,

together with

some points

in

the

K

Vol. 95 (1903).

milaris and E. acutus.
Phil. Trans, of the Royal Society.
development of
of
the National
larv^
Echinoderm
on
the
MacBeide,
E.
Simpson,
J.
Report
W.,
and
A.
(6)
Antarctic Expedition.

Vol. iv. (1908).

Notes on the habits and life-history of Stichopus Japonicus : Annotationes Zoologicae
Japonenses. Vol. v., Part i. (1903).
Die Echinodermenlarven der Plankton-Expedition, etc. Ergebnisse der Plankton(8) MoETENSEN, Th.
Expedition der Humboldt-Stiftung. Vol. ii. J. (1898).
(7)

MiTSUKURi, K.

DESCRIPTION OF PLATES

ILLUSTRATING

Prof.

E.

I.

and

W. MacBRIDE'S REPORT

OF YOUNG HOLOTHURIOIDS.
LIST OF ABBREVIATIONS.
ax.,

II.,

ON A COLLECTION


—

ECHIISrODEEMA.

PLATE
Fig.

1.

I.

Auricularia antarctica viewed from the ventral aspect.

Magnification 25 diameters,

coe.,

the

spherical coelomic vesicle.

Fig.

Fig.

2.

— Some of the calcareous wheels of A.

Magnification 200 diameters,

antarctica.

side, b., c,

—One

of the youngest of the Holothurioids in the collection, viewed

3.

convex

and

d.,

from the concave

a.,

seen from the

side.

from the

side.

Magnification

60 diameters.
Fig.

4.

—

A young Cucumarian, presumably G. lactea, dredged at Plymouth in 1905.
110 diameters. A key is given to this plate.
PLATE

Fig.

5.

—A

median longitudinal section through one

Magnification

II.

of

the youngest Holothurioids in the collection.

Magnification about 60 diameters. The intestine is seen to be perfectly straight and very short, and
the stone canal {st. c.) opens into a closed vesicle {ax.).

Fig.

6.

—A

somewhat

Fig. 5.

lateral

longitudinal section through an older specimen than that represented in

Magnification 75 diameters.

The

axial sinus

is

seen to communicate with the ccelom by

a canal (mad.).
Fig.

—A

median longitudinal section through a specimen about the same age as that represented in
Magnification 75 diameters, pul., a diverticulum of the thin-walled rectum, which is in all
probability the rudiment of the pulmonary trees.

7.

Fig.

Fig.

6.

—

b., and c.
Three transverse sections through one of the younger specimens. Magnification
75 diameters. 8a. is through the tentacles
8b. through the middle region of the body, and
8c. through the posterior end
ff. ? curiously deeply-staining packets of cells resembling rudimentary

8, a.,

;

;

gonads.

The

restriction of the tube-feet to the three ventral interradii is seen, as also the fact that

the most terminal tube-feet belong to the mid-ventral radius.

VOL. VI.