Catalogue of the Chaetopoda in the British Museum (Natural History)

B.
,

M.

(N.H.)

A


(gxiiiB^

T/i7s

Catalogtie

(Utueeum ((Uaturaf J^^^torj).

is

of

No. oCt) of
the

25

Chaetopoda,

copies

Part

of
/.,

Arenicolidie, printed on Special paper.

PRESENTED
BY

^bc

n:ru0tec0
OF

THE BRITISH MUSEUM.



c*

O


:

311

6?6«

CATALOGUE

111?,

OF THE

'-^'^''t

H x\ E T O P O D A
IN

BEITISH

THE

MUSEUM

(NATUIIAL HISTORY).

A.

POTjYCHAETA:

PART L— AREXICOLIDAE.

BY
J.

H.

ASHWOETH,

D.Sc,

Lecturer on Incertebrtttc Zoology in the Unieersity of Ediiihimjh.

LONDON
PRINTED BY ORDER OF THE TRUSTEES
OF THE BRITISH MUSEUM.
SOLD BY
LoKGMANS, Grekn & Co., 39, Paternostku Row, E.G.
B. QuARiTCH, 11, Grafton Street, New Bond Street, W.
DoLAU & Co., Ltd., 37, Soho Square, W.
AND AT the
British Museum (Natural History), Cromwell Road, S.\V.

1912.
{All rights reserved.)

^vr'rs:

!i


:

LONDON
PRINTEP BV WILLIAM CLOWES AND SONS, LIMITED,

DUKE STREET, STAMFORD STREET,

S.E.,

AND GREAT WINDMILL STREET,

W.


PEEFACE
The

present work, which was undertiikeu at the suggestion of the

Kay Lankester, K.C.B.,

former Director, Sir E.
a small

proportion

Catalogue of

tlie

ground

of the

to

Chaetopoda or even of the Polychaeta.
" l*art

Although the volume appears as
catalogue,

its

F.R.S., covers l)ut

he gone over in making a

selection

for

first

tlie

circumstance, depending on the fact

I " of

tliat

a

was

place

more general
a

fortuitous

Dr. J. H. Ashworth, the

author, had already devoted special attention to the Arenicolidae

when
not

the idea was

chosen

beginning.

first

because

But

projected.

as circumstances

thought advisa])le

to

The Family

was supposed

it

to

in question w^as thus

come

have plnced

it

naturally

the

at

there, it has

been

devote some pages to an introductory survey of

the history and classification of the Chaetopoda, and in particular
of the

Polychaeta.

by the author

Morphological characters are largely employed

in characterising the species.

Mr. r. Jeffrey

Bell,

whose

official

duties in the

Museum

include

the care of the specimens of Chaetopoda, has devoted mucli time to
the editing of this A'^olume.

The continuation
at present

provided

of the series of
for,

but

it

is

volumes on Chaetopoda

is

not

hoped that opportunities will

present themselves from time to time of continuing the series which

has been begun by Dr. Ashworth.

SIDNEY

V.

HABMEK,

Keeper of Zoology.

British

Museum (Natural

History),

London, S.W.,
November, 1912.



T.IBLE OF

CONTENTS

.....
.......

1>A«E

List of Text-Figures

Introduction

vii

xi

Historical Accou.nt of the Chaetopod.v, with Speci.\l
to the Polychaeta and their Classification

Reference

.......

Arenicolidae

1

25

Arenicola Lamarck, emend.
A General Account of the Genus Arenicola

29
32
33
38
39
55

....
......

External Characters
External Apertures

Chaetae
Gills

Coelom and Coelomic Septa
Alimentary Canal Burrowing
Nervous System and Sense-Organs
Nephridia
Reproductive Organs
:

Development

61

.

.....
.....

Post-Larval Stages, with a Discussion of the Genus Chj)ne?iidcs
Post-Larval Stages of A. marina

A. cristata
A. assimilis var. ajfi
A. ccaudata
.,
,,
,,
A. hranchialis
,,

,,
,,
Separation of the Genus Arenicola into Sections and Species
The Caudate Section of the Genus Arenicola (with a Key to th
,,

,,

,,

„

,,

,,

.

Caudate Species)

The Ecaudate Section

.......

of the

Genus Arenicola (with a Key

83
to

the Ecaudate Species)

Arenicola marina (Linnaeus)
Arenicola loveni Kinberg, emend.
Arenicola cristata Stimpson
Arenicola glacialis Murdoch
Arenicola j^unill a Quatrefages
Arenicola assiynilis Ehlers, and var. affinis Ashworth.
Arenicola ecaudata Johnston
Arenicola hranchialis Audouin and Edwards
.

.

.

...

Branchiomaldane Langerhans
Branchiomahlanr vinccnti Langerhans
The Inter-relationships of the Members of the Family Arenicolidae
The Affinities of the Arenicolidae

.......
....

63
66
71

72
73
75
77
79
80
80
81
82

84
86
103
105
111
114
123
132
138

147
147

156
159

Systematic Index to the Arenicolidae

163

Gener.\l Index

165

Description of Plates

I-XV

171



—
—

.

LIST OF TEXT-FIGURES

...

.....
....

PAGE

Fig.

1.

Arcnicola loveni.

Fig.

2.

A.

Fig.

3.

A. hrancliialis.

Anterior end, dorsal aspect

Fig.

4.

A. assimilis var.

affinis.

Fig.

5.

A. ccaudata.
specimen, 8

Fig.

a.

—A

Fig.

7.

—

Fig.

8.

A. yusilla.

Fig.

9.

A. marina.
Notopodial chaetae of a post-larval specimen,
4"3 mm. long

42

Fig. 10.

A. ccaudata. Distal halves of notopodial chaetae, from postlarval specimens about 7 mm. long

42

Fig. 11.

A. loveni. Distal third of a notopodial chaeta, from a specimen
335 mm. long

44

Fig. 12.

A. cristata.

Fig. 13.

A. marina.

Fig. 14.

A. assiniilis var. afinin.

inisilla.

Anterior end, dorsal aspect

Anterior end, dorsal aspect

Anterior end, dorsal aspect

.

....

Outline of the posterior end of a post-larval
mm. long, to show the chaetae

Outline of a larva, about '7

cristata.

the chaetae

mm.

Fig. 16.

—

.4.

Fig. 17.

—

^1.

2)usilla.

Ti^js

of

.

.

.

unworn and worn notopodial chaetae
45

Distal fourth of a notopodial chaeta (adult)

.

.

.'

End

46

Distal third of a notopodial chaeta

.

chaetae (adult)

of a fractured chaeta (adult)

...

.........

46
47

Distal portions of notopodial

Crotchet from a post-larval specimen, 5

Fig. 18.

,,

Crotchet from a young adult, 17

Fig. 19.

,,

Crotchets from a specimen, 125

Fig. 20.

40

41

Notopodial chaetae of larvae

ccaudata and A. hrancliialis.

marina.

40

show

long, to

(adult)

¥'1^.15.— A.

34

........
..........41
.........
.....
.........

.

(adult)

33

39

.

ecaudata. Crotchet from one of the posterior notopodia of
the same specimen

.4.

33

mm.

mm. long
mm. long

47

48

long

.

.

49

.

.

49

Ventral end of the neuropodial chaetal sac of a
specimen 250 mm. long, showing the formation of crotchets
,,

.

.........
........

Fig. 21.

A. assimilis v&r.
7*6 mm. long

Fig. 22.

A. assimilis var. affinis.
and 208 umi. long

affinis.

50

Crotchets from a post-larval specimen,

50

Crotchets from specimens 136 nun.
51


List of Text-figures

Vlll

Fig. 23.Fig. 24.-

Fig. 25.-

Fig. 26.-

Fig.

Fig.

27.—


—

List of Text-figures
A. assimilis.

Fig. 55.

56.

Fig.

57.-

Fig.

58.—

Fig. 59.

Anterior end, dorsal aspect

A. assimilis var.

l''ig.

ix

affinis.

B ranchioDt aldanc

Anterior end, dorsal aspect

vineenti.

Anterior end, dorsal aspect

Fig. 60.

,,

,,

Notopodial chaetae

Fig.

,,

,,

juv.

{^^ Clymniides
Notopodial chaeta

.

incertus^')
.

,,

,,

Distal portions of three notopodial

,,

.,

juv.

chaetae
("

Clymmides

incertus ")

Xeuropodial crotchet
Neuropodial crotchet

,,

Adult.

,,

„

juv.

,,

,,

Diagram of the nephridia

,,

,,

Transverse section

,,

^.,

,,

Chjmenides incertus ")
Crotchet from one of the pos
terior notopodia
("

Young

stages

(after INfesnil)

of

.

development

—

INTEODUCTION
The

family Arenicolidae, though small in the

and

species, holds an

number

of its genera

important place in the literature of Polychaeta.

One of the members of this family Arenicola marina, the common
lugworm is the most abundant and most readily accessible Poly-

—

chaete of northern and western Europe.

worm

Probably no other marine

has been so frequently observed, collected and dissected, and,

consequently, the records and descriptions of whicli
other species of

it

is

the subject

Twenty years ago the anatomy
Arenicola was almost unknown the accounts

form a very extensive

of the

series.

of the

;

internal organs, given in descriptions of the genus Arenicola, referred

only to A. marina, and

seemed

it

to be

taken for granted that

other species agreed in structure with this.

all the

Several of the internal

organs, notably the statocysts, oesophageal caeca, nephridia

and septal

pouches have, however, been found to exhibit well-marked differences,
in form or number, in the various species, and to afford considerable
Tn preparing the diagnoses, I have

help in systematic work.
use

full

of

the

internal

characters,

especially appreciated in those cases

mine

defectively

preserveil

or

value of

the

where

it

which

be

necessary to deter-

is

examples,

incomplete

made

will

which,

as

experience has shown, can seldom be diagnosed safely by examination
of their few and imperfect external features.
By means of the keys
and diagnoses provided, the identification of llie members of tlic

family Arenicolidae will,
.ind

<^)uestions of
full

it

is

hoped,

lie

accom])lishe(l with certainty

with conquirative ease.

treatment

synonymy have

in the text.

1

received careful consideration and

have examined

all

the extant types of

the species of Arenicolu, and, in the case of those species the types
of

which are no longer preserved, I have analysed the published
and have com))are(l them with the long series of speci-

tUiscriptions,

mens

my

at

my

disposal, with

conviction that all the

the result that

known forms

can confidently state

I

into

fall

tlie

eiglit

species

described in this Catalogue.

in

addition

to

the

Ibilisb

Museum

(

"olh^etion.

the

entire


Intj-odnction

xii

Collections of Arenicola in the

Museums

of Berlin, Paris,

Cambridge

(Mass.) and the Smithsonian Institution, together with interesting
examples from the Museums of Copenhagen, Dublin, Hamburg,

Eeykjavik,

me

St.

Petersburg, Stockholm and Vienna, have been sent to

for examination.

I

am much

indebted to the authorities of these
my disposal. During the

institutions for placing their material at

have collected, and many friends have generously
of specimens of Arenicola, in various stages
of growth, selected examples of which have recently been added to
The possession of this abundant
the British Museum Collection,

material has enabled me to revise and extend previous work upon
the characters used in diagnosis, and to ascertain their range of
variation.
In addition to the acknowledgments made in the text, I
am glad to have this opportunity of tendering my sincere thanks to
the following friends, who have aided my work by the gift or loan of
Prof. W. B. Benham, P.E.S., Prof. H. C. Bumpus,
specimens
Geheimrat Prof Dr. E. Ehlers, Prof P. Pauvel, Prof. J. D. F.
Gilchrist, Prof. W. A. Haswell, F.Pt.S., Prof. Harold Heath, Prof.
A. D. Howard, Dr. E. S. Lillie, Prof. F. Mesnil, Mr. E. Southern,
and the Directors of the Marine Laboratories at Alexandrowsk (Gouv.
Archangelsk), Cette, Millport, Plymouth, Sautander, Sevastopol and
last

twelve years

sent to me, large

I

numbers

:

Trieste.

To

and to Dr. James Eitchie, of the Eoyal

Museum, Edinburgh, who have kindly read the proofs, I am

Prof. F. Jeffrey Bell

Scottish

greatly indebted for helpful suggestions.
J.

Zoological DErARTMENT,
University of Edinburgh.
Sepfemhcr 23, 1912.

H.

ASHWOiriH.


CATALOGUE
OF THE

AEENICOLIDAE
HISTORICAL ACCOUNT OF THE CHAETOPODA, WITH
SPECIAL KEFERENCE TO THE POLYCHAETA AND
THEIR CLASSIFICATION.

Natukalists as remote as Aristotle were acquainted with Chaetopoda and other worms, the records of which thus extend backwards
The chief object of the
to the earliest works on natural history.
account given in the following pages

historical

is

to

trace

the

principal stages in the growth of knowledge regarding the Chaeto-

poda,

and their classification, and to
schemes of classification proposed the
of the worms which form the subject of the present

especially

indicate
position

in

the

Polychaeta

the

different

Catalogue.
Aristotle recorded in his " Historia

Animalium

"

the occurrence

marine scolopeudrac,^ similar to their terrestrial congeners but
somewhat smaller, redder in colour, and having a larger number of
more slender feet. He stated that these animals are to be found in
the ueighboiu'hood of rocks, and that they do not occur in very deep
water.
The animals referred to were prol)ably nereidiform worms.
Aristotle also mentioned helminthes or intestinal worms.
Pedacius
Dioscorides ^ descri])ed the use in medicine oi Scolojjendra marina,
earthworms and leeches.
of

Allusions to marine scolopendrae occur in the writings of Pliny

and

Aelianns,''

Lib.

'

De

ii,

and Iho former also

cap. xiv, 2.

^yK^l\^^^rll'?>jnu

referretl lo TTiriitlo

'^

nnd Lvmhrlcus.

SaXaTTicu,

written probably about 60 a. n.]. Recens. 0. Sprengel,
Lipsiac (1829), pp. 174, 195, 708, 709.
Nat. Hist., Lib. ix, cap. Ixvii, 3 [about 78 a.d.].
*
De Natura Animal., Lab. vii, cap. xxvi [about 220 a.d.]
-

]\rateria iVfedica

|

'•

.

B


—

Catalogue of Chaetopoda

2

To Pliny and

his contemporaries,

and

to his successors

during the

next sixteen centuries, the names Hirvdo and Litmhricus^ had a
much wider significance than they have at the present day these
writers inchided leeches of all kinds under the name Hirudo, and
;

they applied the designation Lumhricus to intestinal worms or to
earthworms,- or so used it as to include both.

From
little real

During

the time of Pliny onwards for more than a thousand years

advance was made in regard to the knowledge of worms.
wholly or in part, the

this period various authors repeated,

accounts of Aristotle and Pliny, sometimes with fanciful embellishments, but, for the most part, they added little or nothing new.

Most

of

the references

to

worms

in these old writings relate to

parasitic worms, leeches and earthworms, and especially to the
medicinal use of the two latter. The treatise " De Animalibus " in
he works of Isidorus, Bishop of Seville (560-636 a.d.), is noteworthy
"
" De Vermibus
under which heading are included
for a chapter ^
Sanguisuga [leeches], parasitic worms (" Vermes carnium "), namely,
Lumhricus, Ascaridae, etc., and also Multijjes [centipedes],* Scorpio,
Limax, Bomhyx, Teredo, etc. Isidorus placed the Vermes next the
snakes, but took care to point out the fundamental distinction
between them, that is, that the former are without a backbone
" non est illi spinae rigor."
Albcrtus Magnus' (1193-1280) "De Animalibus" contains brief
notes, based chiefly on the works of previous writers, especially
Pliny, on the marine scolopendra, Seta \i.e. Gordius\, Sanguisuga and
Lumhricus.
Edward Wotton ^ (1552) gave a clearly ^vl•itten digest of previous
works, but added little new information in his description of fishes
reference is made to leeches and, in the chapter on " Insects," to
Scolopendra marina, Intestina terrae \i.e. earthworms], Ascaridae and
other parasitic worms.
Shortly after the middle of the sixteenth century there appeared,
in close succession, two great memoirs, both of which contained new
observations on worms, evidently made on living specimens. Belon ^

—

—

;

Pliny, op. cit., Lib. xxxii, cap. xlii, 2; Lib. xi, cap. lii, 1.
E.g., L. J. Mod. Columella, De Ee Eustica, Lib. vi, cap. xxv
Lib. vii,
[Probably written early in the first century.]
cap. ix.
^ Orig. sive Etymolog, Lib. xii, cap. v,
p. 106, in Opera Omnia, Emend. J.
du Breul, Coloniae Agrippinae (1617).
* The words within square brackets are not in the
they are
origiaal
explanatory comments of the present writer.
* De Differentiis Animaliimi, Paris
(1552).
De Aquatilibus, Libri duo, Paris (1553),
'

^

;

;

''


Historical

3

(1553) recorded interesting and accurate ol)servatious on the earthworm and lugworni, and on their liabits. This is the tirst definite

mention

Ingworm, whicli Belon named Lumhricua mnrinns

of the

in

contradistinction to the earthworm (L. terrestrU).

The work

liondeletius (1554, 1555) surpasses that of any
Evidently a keen observer, especially nf niainne

of

previous writer.

animals, IJondeletius described, and gave good woodcuts

Hirudo marina, Vermis

of Scolojx'ndrae marinac,

of,

two kinds
and

mieroryncliotcros

Vermis macrorynchoteros [two Sipunculids], Vermes in tnhulis delitesand Pcnicilkts marimis [a Sabellid].^ In a further
paragraph on marine worms - " I)e vermibus stagni marini " he
gave a short account of " Lumbrici stagni," and in another place ^
referred to Lnmhricns marinus as being similar to Lnmhricus terrenus.
His figures are, in most cases, faithful representations and were
freely copied by (lesner, Aldrovaudus and others.
Rondeletius
added notes on the use in medicine of some of the worms. Gesner's
ccntcs [Serpulids]

—

—

(1558)* is, at any rate as regards the articles on worms, a
compilation from the works of previous writers, especially Belon and
treatise

liondeletius.

Some
letius,

of the later authors above mentioned, for example, Eondewithout proposing any systematic classification of animals, so

arranged those which they described as to indicate the possession
of some idea of their affinities.
One of the earliest classifications
whicli worms are included is tlie tabular statement of the
subdivisions of " Insects " given by Aldrovandus at the beginning
in

"De Animalibus

of his treatise
" Insects "

each of

divided

are

these

absence of

is

feet.

into

again

Insectis "(1602).

" Terrestria "

divided,

and

according

to

"

In this Table
Aquatica,"

the

and

presence

or
the Terrestria without feet are Lumhricus

Among

in man and animals

among the
Scolopendra marina and " Vermes in tubulis
in the subdivision " Aquatica apoda " are
delitescentes " [Serpulids]

terrestris
"

and worms

Aquatica I'edata

found

;

" are

;

Hirudo paludosa and marina, Lvmbricus
and Vermis microrinclioteros
"
[Sipunculids described by Rondeletius].
Liber \\
" De Vermibus
of the treatise of Aldrovandus deals with the Terrestria Apoda,
including Lumhricus terrestris and parasitic worms, but does not
Seta

aquatit'ct

marinus.

[Gordius],

Vermis

macrorinchoteros

—

—

Univ. Aquatil. Hist., pars altera, Lugduni (1555), Liber de Insect, et
Zooph., pp. 108-111.
'

2
*

'

Ojy. cU., p.

145.

Libri de Piscibue Marinis, Lugduni (1554), p. 399.
Hist. Aniro., Liber iiii, Tiguri (1558), pp. 503, 513, 597, 818, 1226, 1227.

B 2


Catalogue of Chaetopoda

4

any of the marine worms above mentioned the description
is given in the following book, which is headed " De
A similar practice was followed by a number of subAquaticis."
sequent writers, for instance, Eay and Linnaeus, who placed marine
worms among " Insects," and earthworms and parasitic worms in the
refer to

;

of these

class "

Vermes."

During the next century and a half there appeared several
memoirs which included studies and figures of worms. Among these,
works by Columna and Bonannus are noteworthy for good figures
Molyneux ^ gave figures
of a Serpulid and of a Nereid and its jaws
of the external characters and " an account of a not yet described
Scolopendra marina" evidently an Ajjhrodite ; Ellis * descril)ed and
figured tubicolous worms, and Peysonnel ^ published " Observations

upon the Sea Scolopendra," in which he described the external
features and the action of the pharynx of a nereidiform worm.
During this period there were also four works of more outstanding
importance by Willis, Eedi, Ray and Bonnet. In Willis' classical
account ^ of the earthworm the external features, gut, circulatory and
reproductive systems are described, and minutiae, such as chaetae,
Eedi ^ described and figured
dorsal pores, etc., are discussed.
portions of the anatomy of Hirudo and Lwnibriais terrestris, he noted
"^

^

;

the occurrence of different species of earthworms, figured Scolopendra
marina [a nereid] and its alimentary canal, gave descriptions and
figures of Hystrix

marina [Aphrodite] and

its alimentary canal, and
in his " Historia Insectorum " (1710)

Eay
figures of a Serpulid.
divided " Insects " into those which

two

undergo metamorphosis and
subdivided
into " Apoda " and
were
The latter
those which do not.
"
"
" Pedata."
Terrestria
were
placed Lumbrici
Apoda
Among the
kinds),
four
and
distinguished
some parasitic
terrestres (of which Eay
"
the
"Apoda
among
Aquatica"
intestinorum
worms "Lumbrici
" were classified according
"
The

Pedata
Hirudines.
were ranged the
six, eight, fourteen or many.
Those
to the number of feet present

—

;

—

1

Aquat. et Terr, aliquot Anim., pp. xxi,

xxii, in

Minus Cogn.

Stirp.,

Eoma

(1616).
"

Recreatio Mentis et Oculi, in Obs. Anim. Test.,

cap. V, p. 30, tab.

Romae

(1684),

pars

i,

ii.

Phil. Trans. R. Soe. Lond., xix (1695), p. 405.
Nat. Hist, of Corallines, London (1755), p. 90, pi. 36, " Tubularia
"
arenosa Anglica" YSnhellaria] ; p. 92, pi. 34, " Corallina tnbularia nielitensis
='

*

[a Sabellid] .
Phil. Trans.
•

<'

"
"

dami

R. Soc. Lond.,

li

(1759), p. 35.

De Aninia Brutorum, Londini (1672), p. 47.
De Animalculis vivis. Ex Etruscis Latinas
(1708),

[1st Edit. 1684.]

fee.

P.

Costac,

Amstelae-


5

Historical
with nuinv

and

feel

—

" A<[natica "

tlie

"rdypoda"

— were

divided into

" Terrestiia "

the section
Scolopeuflra while the " Aqnatica " inchided "
;

,

"

"

comprised Jidus and

Lngs

"

and l^colopcndra

This chissification of the anietaholous " Insects

marina.

Teirestria

'"

followed,

Aldiovandus the extensions were due to Eay's friend
Francis Willughhy.
Bonnet* instituted an extensive series of
experiments and observations on Naids, with special reference to the
regeneration following removal of the anterior and posterior ends.
A preliminary account of these researches was given in letters to
Sir Hans Sloane,- in which also remarks were made on similar
studies on regeneration in earthworms.
The first edition of Linnaeus' " Systema Naturae" (1735) marks
no advance on previous knowledge as regards worms the only
marine worm mentioned therein is Scol(ypcndra marina, which is
in part, that of

;

;

Among

placed in the class Insecta.

Vermes Eeptilia " (the
and Zoophyta) are GorcUiis,

the "

class being divided into Iteptilia, Testacea

Taenia, Zumhricus (including Intcstinum tcrrae, L. latus, Ascaris),
Hirudo and Limax. In the sixth edition of the " Systema " (1748)
the genera of " Vermes lleptilia " are Gordlus, Ascaris, Lumhricus,
Taenia,

Fasciola,

Zoophyta,"

Amphitrite

are
is

Hiriido,

the

and,

genera

included

among

Amphitrite, Nereis

^

the "Vermes
and Aphrodita.

not mentioned in the tenth or twelfth editions

name

;

the

due
to the fact that Miiller had, in the interval between the twelfth and
thirteenth editions, founded a genus of worms with this designation.
In the tenth edition of the Systema (1758), which is now regarded
as the foundation of zoological nomenclature, Linnaeus included the
genera Lumhricus (including the species terrestris and marinus),
Hirudo, Aphrodita, Nereis and Serpula, the species of which together
number more than forty; in the twelfth edition (1767) T'erthella

and Sabella (and Sipuneulus) were added to the list. The names
Lumhricus and Hiriido had been in use since the time of Pliny,
though the former, as defined in the tenth edition, has a much
reappearance of this

less

extensive significance

in the thirteenth (Gmelin's) edition

than

(see p. 2); the other generic

it

bore

in

names appear

is

pre-Linnaean writings
have been used for

to

first time by Linnaeus.
In the tenth and twelfth editions of the "Systema," Linnaeus
divided animals into six classes Mammalia, Aves, Amphibia, Pisces,

the

—

'

*
'

Traite d'Insectologie, 2 Partie, Paris (1745).
Phil. Trans. R. Soc. Lond., xlii (1743), pp. 468-487.
Under which ScolopeiuJra marina is given as a sjTionyiu.


—

Catalogue of Chaetopoda

6

— distinguished

by the number of chambers in
first two classes with
" sanguine calido, rubro," the third and fourth " sanguine frigido,
rubro " and the last two " sanie frigida, albida." Insects were distinguished by the possession of antennae, Vermes by having " tontacula."

It follows from this mode of classification that any invertebrate not
having antennae, and therefore not referable to the class Insecta,^
would fall into the class Vermes, which therefore necessarily became
a very heterogeneous assembly.
In the twelfth edition (1766-8),
which was the last revised by Linnaeus, the class Vermes was
Insecta and

Vermes

the heart and the nature of the blood, the

subdivided

seven

into

orders

— Tardigrada,

Imperfecta,

Mollusca, Testacea, Lithophyta and Zoophyta

need be considered in this account.
these three orders in the following

— only

Intestina,

three of which

The genera were arranged under
manner
:

quondam dicta, ob summam simplicitatem
Animalia simplicia, absque artubus, nuda. Lumbrieus, Sipunculus, Fasciola, Gordius, Ascaris, Hirudo, Myxine.
Vermes Mollusca, nuda, brachiata, vagantur pleraque per maria, ....
Animalia simplicia, nuda (absque Testa inhabitata) artubus instructa. Actinia,
Ascidia, Limax, Aplysia, Doris, Tetliis, Holothuria, Terebella, Triton, Sejna,
Clio, Lernaea, Scyllaea, Aphrodita, Nereis, Medusa, Asterias, Ecliinus.
Vermes Testacea, mollusca, domiporta, calcarea*[ue domuncula nobilitata
.... Animalia Mollusca simplicia, domo, saepius calcarea, propria obtecta.
Chiton, Lepas, Pliolas, Mya, Solen, Tellina, Cardium, Mactra, Donax, Venus,
Spondijlus, Chavia, Area, Ostrea, Anoniia, Mytilus, Pinna, Argonauta,
Nautilus, Conus, Cypraea, Bulla, Volnta, Buccimim, Stromhus, Murex,
Trochus, Turbo, Helix, Nerita, Haliotis, Patella, Dentalium, Serpula, Teredo,
Vermes Intestina,

teiTena

corporis, terebrant omnia.

Sabella.
It is evident

from this commingling

of

worms and members

other phyla that the work of previous naturalists and his
vations

were not

sufficient

characteristic features of the

to

indicate

worms

clearly

wdiich

to

own

Linnaeus

we now group

of

obser-

the

together as

Annelids, the seven genera of which he placed in three different
classes, or even to enable him to separate worms from Mollusca and

Echinoderma.
affinities

of

Linnaeus had, however, some conception of the
both Serpula and Sabella, for, after the name of the

former genus he added " Animal Terebella," and after the name of
the genus Sabella Awimsil Nereis" (12th edition, pp. 1264, 1268),
and under the species Sabella alveolata he stated " Genus hoc multa
''

habet communia

cum

Nereidibus sed

&

os

&

tentacula oris diver-

In both the tenth and twelfth editions of the " Systema " (pp. 639, 1064,
respectively), Linnaeus placed Scolopendra marina in the class Insecta.
In
the twelfth edition he stated that this animal was used as bait for herrings.
'

There can be no doubt that one or more species

of Nereis

were thus indicated.


:

Historical

7

But the presence of the "test" evidently outweighed these many common characters, so that Sahella was not

sissima

"

;

placed near Nereis but

among

the shelled Mollusca.

Gmelin, in his (the thirteenth) edition of the "Systema" (1788),
made no advance in regard to the separation of Worms from Mollusca.
The only changes from the twelfth edition which call for comment
here are (1) the addition to the Vermes ^lollusca of the genera
Amphitrite Miiller, Spio Fabricius, and Nais Miiller and (2) the
removal of the species Scolopendra marina from the Insecta, and its

—

;

synonym under Nereis versicolor and noctiluca. Gmelin
seems to have taken practically no notice of the suggestive work of
Pallas on Serpula (vide infra), for he still retained this worm among
the shelled Mollusca placed in the Vermes Testacea, and did not
adopt the improved arrangement of some of the worms suggested by

reduction to a

Miiller (see p. 8).
Tlie stimulus given to the systematic study of animals by the
appearance of the " Systema Naturae " soon produced a marked
effect

on the growth

of

knowledge in regard

to the class of

Vermes.

Observations on worms, in some cases accompanied by fairly adequate
descriptions of their

characters,

appeared in treatises of natural

and in faunistic works, and there were
also accounts or memoirs pu])lished on single genera or species.
Seba ^ figured Millepeda marina [two nereidiform worms], Eruca sen
Scolopendra marina \Apliroditc\ Penicillum mariiium [a Sabellid]
and a cluster of slender worm tubes. Baster^ prepared a series
history, in records of travel

of plates with good figures of the external

characters of Nereids,

" Scolopendi'ae

plumosae " [Sabellids], Serpulae [including a fragment of a Lanice and its tube], " Hirndo piscium" [Pontobdella^,
Aphrodita acidcata, A. squamata [a Polynoid] and Nereis pelagica ;
in the case of the last three some details of the feet were given.
Pennant's "British Zoology" (vol. IV, 1777) contains figures of
Lumhricus marinns, terrcsiris and minor, leeches, a Sipunculid,
Aphroditidae and Polynoids, Nereids and the tubes of Serpula and
Spirorhis.
Pallas"" puldishod in 1776 an account of his admirable researches
on the anatomy, both external and internal, of Aphrodita [including
under this name Aphrodite, Polynoids and Amphinomids], Nereis

'

•*

lleium Nat. Thes.

i,
Anistelaedami (1734), tab. Ixxxi, 7,
(1758), tab. iv, 7, 8: tab. xvi, 7a, 7b: tab. c. 8.
Op. Subseciva, i, Harlemi (1759-60), tab. iv, v, ix, x ; ii (1765), tab. vi.
Miscell. Zool., Hagae Comitiuiu (1766), tab. vii-xi.

Locupl.

tab. xc, 1-3

;

iii

8


—

.

.

.;

.

Catalogue of Chaeiopoda

8

cylindraria [Fectinaria\,
[a Sabellid],

Fierpula,

Nereis

and

concliilega

Lv/inhricns

[Lanicc], Nereis
\Ec]iiiLrus\,

ccliivrns

lutaria

and

in

on various
"Nereis," including "Nereis Inmhricoides" [Arenicola marina], and

1788 there appeared a further

some tnbicolous worms

Up

series

of

his

studies

^

liucludiniui; Spirorhis\.

to the time of Pallas, and, indeed, for a considerable period

subsequently, the tul>es of Serpvla were described and figured in
works on conchology - along with the more loosely coiled gastropod

but Pallas showed in his earlier paper ^ that the shell of
Serpula is different from that of the true Testacea, that Serpula
agrees fundamentally in structure with Nereis and Aphrodite, with
shells

which

;

it

placed

should be imited to form an order, in which should also be

genera Lvmhricus, Hirudo, Ascaris, Gordius, and CA'en
was thus the first to recognise some of the essential

the

Pallas

Taenia.

differences

Otto

between worms and molluscs.
Miiller *
and Otto

Friedrich

elaborated the systematic details regarding

much

Fabricius

worms

^

successfully

of various kinds,

and
did
Nais,
and
founded
genera
Amphitrite
and
Miiller
the
species.
of
genera
previously
established
much to clarify the diagnoses of
having
class
the
His
classification
of
the
Vermes
worms.
(1776)
He

the limits defined by Linnaeus
is well worthy of notice.
devoting

care to the distinction and definition of genera

—

—

divided the class as follows

:

1.

Infusoria [animals living in infusions]

2.

Helminthica.

MoUusca

[similar to the order Mollusca of Linnaeus, except that the
genera Aphrodita and Nereis were transferred to the division
Helminthica]
4. Testacea [corresponding to the Testacea of Linnaeus]
5. Cellularia [corresponding to the Lithophyta and Zoophyta of Linnaeus]
The Helminthica found in Denmark and Norway were subdivided into two

groups
I.
Mutica, containing Gordius, Ascaris, Echinorhynchus,
Hirudo, Taenia ; II. Setosa, containing Lumbricus, Nereis, Amphitrite, Nais, Aphrodita.
3.

:

This classification marks a distinct advance on that of Linnaeus,
showing good progress in the direction of a separation of worms and
molluscs.
JMiiller's order Helminthica contains worms only.
There
'

^

Nova Acta Acad.

Sci. Imper. ii, 1784, Petropoli (1788).
See, for instance, d'Argenville's Conchjdiologie, Paris, 3 Edit., vi (1780),

tab. Ixviii.
=»

Op.

cit. (1766), p. 74.

Vermium

Terr, et Fluv., Havniae,
Zool. Dan. Prodr. (1776).
° Fauna Groenlandiae, Hafniae
(1780).
*

i

(1773), pars

i

:

pars altera (1774)


Historical
are,

9

however, three worms not inchided with the rest, namely,
T-., tlio newly defined genus rhniarui Miiller, and Serpvia

FftHnola
L.

;

lull

it

knowledge

must be a(hnitted

that, lakint; into

at tliat time, these

scheme

correctly in the

of

aecount the state of

tiuee were diflicult genera to place
Miiller referred Fasciola

classification.

and Scrpula to the Testacea. He does
not appear to have known tlie work of Pallas on Serjmla ; at any
Miiller was the first to
rate, he did not refer to it in his synonymy.

use the presence of chaetae as the distinguishing cliaracter of a group
and rianaria

to the Mollusca,

of worms.

Blumeuhach pointed out that Vermes differ from Insecta not
only in the aljsence of antennae hut also of jointed locomotor organs.
He was the first to state and emphasise this fundamental difference
^

hetween the jointed appendages of " Insecta " \i.c. Arthropoda] and
the feet of worms.
His classification closely follows that of
Linnaeus.
Barhut,Ijased

errors.

its

Bruguiere^ and others, produced systematic memoirs
Linnaean system and reproducing many of
The majority of ti-eatises on natural history published

largely on the

during the last third of the eighteentli century held tenaciously
to the Linnaean classification of tlie Vermes, and in the hands

most workers this class was still in the same unsatisfactory
The work of Pallas on
it had heen left by Linnaeus.
Scrjmla and the outline classification given l)y Miiller were the first
of

condition as

indications of the

dawn

of order, which, in the closing years of the

upon the chaotic assemblage of Vermes.
In 1795 Cuvier communicated to tlie Societe d'Histoire naturelle of
Paris a memoir* on the circulation in " animaux a sang blanc," in
eiglitcenth century, broke

which he described the heart and blood-vessels of various molluscs,
and also gave a Table showing the nature of these organs in various
The work done in preparation for this memoir
classes of animals.
brought clearly before him the characters which distinguish worms
from molluscs, and from this time for\vards Cuvier separated these
two classes of animals. In his next memoii- " Tableau elementaire
the two
de I'histoii-e naturelle des animaux " (Paris, An T), = 1798)

—

—

Handb. dcr Naturg., Gottingen (1799), 6 Aufl., p. 401. [1st Edit., 1779.]
Genera Vermium, London (1783).
^ Hist. nat.
Brugui^rc
des Vers in Encyclop. method., Parin (1791).
established a new order in the class Vermes Vers I^chinodermes to contain
the star-fishes, sea-urchins, etc. The other Vermes were left in the same
arrangement as in the Systema.
1

-

—

—

*

Bull, des Sci. par la Soc. Philom., Paris,

i,

An

iii

[1795], p. 91.