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OLIGOCHAETA


JJonbon

HENRY FROWDE
OXFORD UNIVERSITY PRESS WAREHOUSE

AMEN CORNER,

MACMILLAN &

CO.,

E.C.

66 FIFTH AVENUE


/:,

A

MONOGRAPH
OF THE ORDER OF


OLIGOCHAETA
BY

FRANK EVERS BEDDARD
M.A. (OxoN.), F.R.S.
PROSECTOR TO THE ZOOLOGICAL SOCIETY OF LONDON

AND LECTURER ON BIOLOGY AT

GUY'S HOSPITAL

OXFORD
AT THE CLARENDON PRESS
M DCCC XCV

*$
i


PRINTED AT THE CLARENDON PRESS
BY

HORACE HART

PRINTER TO THE UNIVERSITY


PREFACE
are already two works dealing with the Order Oligochaeta.

first

of these, in time of publication,

is

The

Professor Franz Vejdovsky's

'System und Morphologie der Oligochaeten,' which appeared in 1884. Six
years later Professor L^on Vaillant contributed to the volumes on the
'

'

a volume and a half dealing with the
same group. It might appear, therefore, at first sight, that the ground has
already been taken up, and that there is no occasion for the present work.
It must be recollected, however, that it is nearly ten years since the
Annelids of the

Suites k Buffon

publication of Professor Vejdovsky's treatise,

and that during

this


period

more particularly of the earthworms,
Professor Vejdovsky's object, moreover, was more

our knowledge of the Oligochaeta,

has increased enormously.
especially to give an account of the structure of the group from his own
observations, and to spend less time in dealing with the results of other
not that the previous work upon the group was by any means
neglected by him, but no great amount of detail was given as to the
genera and species of Oligochaeta not occurring in his native country.
observers

work

rather more comprehensive in scope, but there
so that the large
is no account of any researches made after the year 1 886
amount of literature which has appeared since that date has not yet been

M.

Vaillant's

is

;


M. Vaillant's contribution also is rather
incorporated into any general work.
more devoted to the systematic side of the subject than to the description
of structure.

appeared to me that there was room for
a treatise of rather wider scope than those of either Professor Vejdovskv or

Under these circumstances

it


PREFACE

vi

of M. Vaillant, and one which should deal with the entire subject up to the

date of publication.
As will be seen from the bibliography which I have
given in the Appendix, the literature of this subject is large and scattered

;

so that to bring together under one cover all of importance that

known about the group

is


as yet

of the Oligochaeta will possibly be considered not to

have been a useless performance. In attempting this labour I have received
much kindly encouragement from Professor Ray Lankester, which I gratefully
I have been so fortunate, through the kind influence of
acknowledge.
Dr. Burdon Sanderson, as to secure the assistance of the Clarendon Press,
to the Delegates of which institution I wish to express
ness.

The

my

great indebted-

cost of the necessary illustrations, which has been considerable,

has been largely defrayed by the generosity of Mr. J. P. Gassiot, F. Z. S.,
100 at my
who, at the suggestion of Mr. Sclater, has placed a sum of

Without

disposal for this purpose.

this very liberal act


of assistance on

Mr. Gassiot's part this work could hardly have been undertaken.
The
I may now say a few words about the scope of this monograph.
study of the group with which it deals is one which has occupied my leisureI have had the opportunity of dissecting
time for the past fifteen years
and examining most, if not all, of the more important types so that while
;

;

a good part of the volume
siderable proportion of

is

necessarily derived from compilation, a con-

the result of first-hand knowledge. Whilst
recent papers into the present work, I have also

it

is

have amalgamated my
incorporated with it a certain amount of new matter which
I


I

have not

published elsewhere, and I have given, in the systematic part, descriptions of

have not, however, made any lengthy investigations
for the express purpose of this monograph, but I have in several cases verified
the statements of others, and have corrected, in a few minor particulars,

a few new

species.

I

errors of observation on the part of myself, as well as of other zoologists.

Those who have not followed closely the progress of knowledge in this group
of animals may be surprised at the large number of pages which it has taken

me

to set

down the

facts


:

I

have erred, however, rather on the side of

compression and omission than on that of undue prolixity. The omissions
(with a few exceptions to be noted immediately) will not be found, I trust,
to be of facts of much importance.
They chiefly concern the progress of our


PREFACE

vii

It has not appeared to me to be necessary
acquaintance with the group.
to go at great length into the history of erroneous views or of misstatements
of fact at any rate compression and omission here seemed to be more per-

missible than in other departments of the subject.

I

have deliberately omitted

any account of the early stages of the development of the Oligochaeta
the development of organs will be found treated of to a certain extent.
to give


;

determined to do this principally on account of Prof. Vejdovsky's recently
published Entwickelungsgeschichtliche Untersuchungen,' an elaborate and
I

'

work, which goes into the matter with all details, and
treats of the rest of the literature in a most thorough fashion.
Moreover,
I have personally no first-hand acquaintance with the early development
finely-illustrated

of the Oligochaeta

another circumstance which leads

me

simply to refer to

the work already quoted those who are desirous of ascertaining what is known
Another branch of the subject
about the embryology of the Oligochaeta.
which I have thought it well to abridge is the section dealing with
unrecognizable species.

No


name,

I hope,

has been omitted, and no reference

;

but I have not, except in a few instances, gone at any length into the endless
possibilities as to the identity of species imperfectly described and now
irrecoverable.

preparation two excellent revisions of two
I refer to Dr. Rosa's detailed
families of the Oligochaeta have appeared
account of the Lumbricidae, and to Dr. Michaelsen's
Synopsis of the

While

this

work was

in

:

'


'

Enchytraeidae. The appearance of these two valuable papers caused me to
hesitate a little before dealing with the respective families in this work.
I have, however, thought it best to incorporate both of them, not, I hope,

without

critical

examination.

In the preparation of the systematic part of this monograph I received
the greatest assistance from M. Vaillant's work
I gladly acknowledge that
it formed the basis of
my preliminary (manuscript) account of many of the
;

species of Oligochaeta,

compilation
descriptions

;

I believe,

and that


it

has saved

me

a vast amount of labour

however, that I have nowhere followed M. Vaillant's

and synonymies without

careful verification

In the general sketch of the structure of the group
with generalities the details of particular genera, or it
;

in

and
I

criticism.

have only dealt

may


be of species,


PREFACE

viii

be found under their respective headings.
My own experience is that
in reading elaborate monographs the multiplicity of details tends to throw
will

one

off

much

the main course of the argument.

Details of minor importance are

better referred to their proper place instead of being included in one

I have therefore attempted
long dissertation on the structure of the group.
to give in my introductory chapter such an account of the anatomy of the

be sufficient to satisfy any person not interested in the
minute details, but desirous of having the main facts stated in as few words

Oligochaeta as

as possible.

may

It

may

be thought that

I

have erred

in the brevity of this

I claim, however, to have put before the reader a more complete
chapter
account of the structure of the group than can be found in any treatise yet
:

published,

and

I

have done


my

best to avoid details not of special significance

except as generic or specific characters.
In the systematic part I have not treated each family in a precisely
similar fashion.
In the Perichaetidae, for example, the internal structure is
dealt with after the definition of the family
details are reserved

;

in the Geoscolicidae, anatomical

In the highly peculiar group
somewhat intermediate. In each case the

as generic characters.

Eudrilidae the method adopted

is

plan followed is not, as might perhaps be suspected, the result of the discontinuous preparation of this monograph, but has been deliberately selected
as being, in

my


opinion,

Dr.

Benham,

in question.

the preparation of this work I am greatly indebted
Prof. Claus, Mr. W. T. Thiselton-Dyer, C.M.G., F.R.S.,

For material used
to

most appropriate to the family
in

Mr. Chamberlain, Mr. Gustav Eisen, Mr. Everett, Mr. F. Finn, Dr. Gregory,
Prof. Loven, Dr. Michaelsen, Prof. M'Intosh, F.R.S., Mr. Alvan Millson,
Prof. T.

J.

Parker,

F.R.S.,

Prof.

Poulton,


F.R.S.,

Dr. D. Sharp, F.R.S.,

W. W.

Smith, Mr. Sowerby, Prof. Baldwin Spencer, Prof. Vejdovsky,
the Rev. H. W. Woodward, the St. Petersburg Museum, and the Zoological

Mr.

Society of London.

FRANK
LONDON, February, 1895.

E.

BEDDARD.


TABLE OF CONTENTS
PAGE

DEFINITION OF OLIGOCHAETA

PAET

I.


.

,

THE ANATOMY OF THE OLIGOCHAETA.

HISTORICAL NOTE

i

THE BODY-WALL AND EXTERNAL CHARACTERS

2

THE NERVOUS SYSTEM

18

COELOM

25

NEPHHIDIA

.

31

ALIMENTARY CANAL


53

VASCULAR SYSTEM

64

BLOOD-GLANDS

77

RESPIRATORY ORGANS

81

REPRODUCTIVE SYSTEM

84

GEOGRAPHICAL DISTRIBUTION

148

PAET

II.

SYSTEMATIC.

THE CLASSIFICATION OP THE OLIGOCHAETA


155

PHYLOGENETIC ARRANGEMENT OF THE OLIGOCHAETA

162

DESCRIPTIONS OF GENERA AND SPECIES

174

......

GROUP APHANEURA

176

FAMILY AEOLOSOMATIDAE

176

GROUP MICRODHILI

.

.

.

.


FAMILY PHREOHYCTIDAE

187
187

'

FAMILY MONILIGASTHIDAE

.

b

.

192


TABLE OF CONTENTS

x

PAGE

SUPEBFAMILY LuMBRICULlDES

2O6

LUMBBICCLIDAE


.

.

.

207

FAMILY TUBIFICIDAE

226

{FAMILY
FAMILY NAIDOMORPHA

275

FAMILY ENCHYTBAEIDAE

308

GBOUP MEOADBILI

357

SUPERFAMILY MEGASCOLICIDAE

357


PEBICHAETIDAE

.

(FAMILY
FAMILY ACANTHODBILIDAE

359

.

...

FAMILY CKYPTODBILIDAE

.

.

.

443

516

FAMILY EUDBILIDAE

573

FAMILY GEOSCOLICIDAE


622

FAMILY LUMBBICIDAE

687

BIBLIOGRAPHY

725

.

INDEX OF GENERA AND SPECIES

.

753


EXPLANATION OF PLATES
PLATE

sp

gl,

spermathecae

sp,


spermiducal gland

g, gizzard

;

spermiducal glands

;

c?

male pores.

;

sp p, pore of spermathecal

sac.

;

;

male pores;

<

;


p, genital papillae.

Libyodrilus violaceus.

male pore.

;

Fig.
<$

? oviducal pore

;

Perichaeta posthnma.

Fig. 6.

:

everetti.

;

? oviducal pore

;


sp p, spermathecal pore

$ female pore

;

peptonephridium g, gizzard nph, nephridia ; dv, dorsal
v v, ventral blood-vessel.

gl,

n, nerve-cord

Fig. 5.

worm

nph, nephridia

;

Octochaetus multiporus.

Fig. 4.

;

male pore

Dissected slightly from the side,

Ventral view of

dv, dorsal vessel

;

Polytoreulus magileiisis.

Fig. 3.

gl,

sperm-sac

spermathecal pores

sp,

Ventral view of anterior segments.

sp

s,

Perichaeta

2.

Fig.

;

sp

;

caec, caeca.

;

Ventral view of anterior segments,

vessel

Pericliaeta indica.

i.

Fig.

Dissected,

I.

Eudrilus eugeniae.

7.

male pore.

PLATES

II

AND

III.

Generative organs of various types of Oligochaeta diagrammatically represented.
The male
organs are coloured pink, the female blue. To all the figures the following lettering applies:
T, testes ; F, sperm-duct funnels ; sp s, sperm-sacs ; o, ovary od, oviduct ; e s, egg-sacs ; os, sper;

mathecal sac

;

spermathecae

sp,

;

sp

spermidncal gland.

gl,

PLATE
i.

Fig.

Dissection,

orifice

o,

vv, ventral blood-vessel

;

of

spermathecal

(?)

:

g,

gizzard

spermiducal glands

The

clitellar

;

sj)s,

sperm-sacs;

sp

gl.

spermiducal glands;

n, nephridia.

SipJionogaster millsoni.

2.

processes.

The remaining
letters

Libyodrilus.
sac;

dv, dorsal blood-vessel

Fig.
Tr, penial

IV.

figures bear a legend.
;

;

In

ca, calciferous glands

vd, vas deferens

;

all

of

them the following

is

the significance of the

'

;

sp,

spermathecae

;

sp

s,

sperm-sacs

;

h,

'

heart

;

od, oviduct.

segments are numbered

;

the dorsal vessel with

its

branches are coloured red.

sp

gl,


EXPLANATION OF PLATES

xii

PLATE
Fig.

V.

Stuhlmannia.

i.

Longitudinal section through anterior segments,
ea, calciferous

glands

w,

;

ventral blood-vessel

br,

brain

;

n, ventral

dv, dorsal blood-vessel

;

;

nerve-cord
s,

;

g,

gizzard

;

masses of perivisceral

corpuscles.

Octochaetus multiporus.
Fig. 2.
Transverse section through oesophageal region of young worm,
vessel

;

septum

,

;

ca, calciferous

gland

;

TO,

Eudriloides

Fig. 3.

Transverse section,
ca, calciferous glands

sps,
i,

;

gpennathecal

dv, dorsal vessel

;

vp, ventral

nerve-cord.

sac

with

cotterilli.

appended glands

dv, dorsal vessel

(gl).

;

oesophagus.
Fig. 4.

Stuhlmannia.

Section through peninl process, gl, glandular epithelium of penis ; m, septum dividing its
cavity from general coelom ; vd, vas deferens ; n, nerve-cord ; dv, dorsal vessel ; 7, intestine ;
el.

clitellar

epithelium

;

sps,

sperm-sacs

;

p, muscular sac opening at

end of

penis.

ERRATA
p.

To synonyms

183.

of Aeolonoma Itemprichn add Aeolortais hemprichii

and Aeolonais decorum of

GERVAJS.
p. 184.

To synonyms

of Aelosoma quaternarium

add Aeolonais quiiternarium of GEHVAIS.

p.

214.

To synonyms

of Lumbriculus variegatus

add Tubifex yentilinus, DUGES

p.

229,

1.

14.

For Tflmatodnlus read Telmatodrilini.

p. 251,

1.

24.

For bogdunovii read boydsmovii.

p.

266,

1.

13.

p.

275.

To

285,

4.

p. 288,

1.

12.

313,

1.

15.

p.

]>.

VAILLANT).

For.Hemitubifex ater read Hemitubifex benedii.

list

1.

(fide

of genera of

Naidomorpha add

Ripistes.

For O.josinae read N.josinae.
For P. dinguis read N. elinguis.
For vermicvlus read vermicularis.

For M. fusca read M. semifusca.
For E. hegemon read F. hegemon.

p. 332, footnote.
p.

348,

1.

!<;.

A

p. 394.

few species (10) have been accidentally omitted from the list,
of Fletrlierodrilus unicug add Cryptodrilus pdewensis, MICHAELSEN.

For
C.
3.
gravidis read C. grandis.

To synonyms

p.

481.

p.

498,

1.

1

p.

530,

1.

1

p.

552, last line.

8.

p. 608,

1.

23.

66 1,

1.

29.

p.

For A. eommunis read D. communis.
For A. multi^orus read 0. multiporus.

For P. elongatus read P.finni.
For U. pairillata read U. papillifer.

NOTB.

Formal descriptions of Brachydrilus, Megascolex hallii, aud of Perichaeta dubia, have
But the principal facts in the anatomy of these species will be found
unfortunately slipped out.
scattered through the volume, and can be referred to from the Index.

Plate

Cue

-

Beddard's Monograph on Ohgochasta.

I.



us

Eu

'



IX

'sp

\l

-sp.s

i:

IV

V

VI

VII

via

a
X

a
XI!

\m

w
AT

Wl

try/

ix

SpSOLC.J

spgl..

',1

xai

'

1

'

Monograph on Oligochasta.

Benhenma,



ca.

,

M

I

16
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;