Zoological Journal of the Linnean Society, 63: 275-287. With 3 figures
July 1978
Systematics of Dodecaceria (Annelida:
Polychaeta) and its relation to the
reproduction of its species
PETER H. GIBSON
Department of Zoology, The University, Newcastle upon Tyne, NEI 7R U, U.K.
Accepted for publication August 1976
The systematics of t h e genus Dodecaceria is confused due to t h e morphological similarity of
species. Eleven o u t of a total of fourteen species in the genus were compared by their
morphology, chaetal type and distribution, numbers of branchial cirri and body segments and
by their methods of reproduction. The genus is remarkable for showing sexual, asexual and
parthenogenetic reproduction which were briefly reviewed. Of the different methods of
comparison those using the types of reproduction were thought t o be the most useful. The
entirely descriptive methods were felt t o be entirely inadequate.
CONTENTS
. . . . . . . . . . . .
Introduction
Materials and methods
. . . . . . . . .
Results
. . . . . . . . . . . . .
Comparison of general morphology
. . . .
Comparsion of chaetal shape and distribution
Comparison of body length and number of cirri
Comparison of methods of reproduction
. .
Discussion
. . . . . . . . . . . .
Acknowledgements
. . . . . . . . . .
References
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275
276
277
277
278
278
279
284
286
286
INTRODUCTION
The systematics of the genus Dodecaceria comprising some 14 species
(Hartman, 1944; Knox, 1971) has never been fully revised and is at present in a
confused state. Much of the confusion stems from the morphological
homogenity of the group. The first species to be described was D. concharurn
by Oersted (1843) but in retrospect the description is inadequate in several
respects. It was added to by Grube (1855) and Quatrefages (1865) when they
described the same species only under different names. The early descriptions
were undoubtedly hampered by D. concharum invariably coexisting with D.
caulZeryi in the regions sampled. The presence of two species was first
suspected by Caullery & Mesnil (1898) although they did not establish this.
0024-4082/78/0063-0275/$02.00/0
0 1978 The Linnean Society of London
275
276
P. H . GIBSON
Separation of these two morphologically very similar species was definitely
established by Dehorne (193 3) on the basis of their reproductive biology.
Similar taxonomic problems still exist within the genus. For example the
North American species D. finibriata first described by Verrill (1879) under the
name of Heterocirrzrs jimbriatiis appears to be morphologically identical to D.
cuullerj*i (Martin, 1933). Also according to Reish (1952) the boulder building
species D. jewkesi is synonymous with D. fistulicola and this is accepted by
Knox (1971). Hartman (1959), however, catalogues both species separately.
The position is further complicated by Mesnil & Fauvel(1939) who thought D.
fistiilicola is undoubtedly synonymous with D. joubini. Previously Augener
(1914) suggested that D. juubirzi is identical with D. opulens. The similarity of
the three species t o each other as well as D. caulleryi and D. concharurn was
commented on by Fauvel (1930a, b ; 1953). In these an understanding of the
reproductive biology of the species might well help to establish their taxonomic
positions. This field, as it applies to other polychaetes, has recently been
reviewed by Clark (1977).
The aim of the present paper is to consider eleven species of Dodecaceria
and show how a knowledge of their reproductive biology may assist in their
systematics. The remaining 3 species of the 14 in the genus were not examined
but are briefly considered.
MATERIALS AND METHODS
All work was carried out on preserved specimens which were either from
museum collections or were obtained directly from field samples (Table 1).
When using field samples the best preserved specimens were obtained by
extracting the worms before fixing but this was only possible for Dodecaceria
caulEerj~iand D. concharzirn which were collected locally from a Northumberland beach (Gibson & Clark, 1976; Gibson, 1977). The majority of species
Table 1. Source of samples used in the present paper
Species
----___
D. berkeleyi
D. berkeleyi
D. capensis
D. caulleryi
U . concharum
D. coralii
D. fewkesi
D. f e w k e s i
D. f e w k e s i
D. fimbriata
D. fisrulicola
D. fistulicola
D. f'irtuficob
D. laddi
D. laddi
D. opulens
D. pulchra
D. pulchra
Location
-_______
New Zealand
Keikourar Peninsula
One Tree Island
Western Australia
False Bay
South Africa
Cullercoats Bay
Northumberland
Cullercoats Bay
Northumberland
Woods Hole
Massachusetts
Languna
California
Long Bay
British Columbia
British Columbia
Round Island
Newfoundland
Fosdick Point
Washington
Lime Kiln Light
Washington
Vina del Mare
Chile
Pulo Boenda
Sumatra
One Tree Island
Western Australia
Chile
Kalk Bay
South Africa
False Bay
South Africa
* On loan from the
United States National Museum.
t Australian Museum.
Date o f collection
24.2.7 3
-. 1 0 . 7 2
27.10.71
15. 8.69
15.8.69 24.9.69
17.7.53
19.11.39
-. 8.31
10.9.71
29.6.56
16.8.7 1
1951
-.11.63
-.10.72
21.9.64
1.10.71 27.7.72
28.5.75
Collector
G. Knox
P. Hutchingst
J . Day
P. Gibson
P. Gibson
M. Pettibone*
McGinitie*
-
E. & B. Berkeley*
D. Barnes
M. Pettibone*
J. Allen
F. Rivero-Ziniga*
A. Koln & M. Lloyd*
P. Hutchingst
J . Castilla
J . D a y L? C. Griffiths
C. Griffiths
S Y S T E M A T I C S OF DODECACERIA
277
from field samples were collected from different parts of the world (Table 1)
and sent after fixing the worms in their burrows. Formalin was found to give
the best preservation when the entire samples were fixed. The general
morphology of the worms was examined with special attention to the shape of
their nuchal organs and anus. Samples of chaetae were taken and the number of
body segments and branchial cirri counted. The methods of reproduction were
identified from the types of gametes in the coelom, the presence or absence of
epitokes and the existance of asexual reproduction indicated by regenerated
regions of the body.
RESULTS
In the atokous condition species have the following characters. In the
presegmental head region is a subterminal ventral mouth and a pair of nuchal
organs. The first segment is achaetous and bears a pair of grooved tentacles
above which is pair of filiform branchial cirri. The following anterior chaetose
segments each have at least one pair of dorsal branchial cirri. The body is
approximately cylindrical with the posterior region becoming slightly dorsoventrally flattened. The chaetae are capillary and simple or acicular with a
subterminal spoon-shaped depression which may have a slight proximal protruberance. Fresh individuals vary from brown to greeny black and there is usually
a prominant dorsal serpentine blood vessel running along the anterior region of
the body. Certain species have epitokes which are essentially the same as the
atokes. The differences are there is a large pair of orange to red eyes and the
body is more darkly pigmented and has long capillary swimming chaetae which
have replaced the acicular chaetae of the atoke. All species inhabit laterally
flattened, flask-shaped burrows which have a single opening in which the head
and anus lie. Species form dense colonies in, as a rule, a calcarious substratum
and are found either in intertidal rock pools or sublittorally.
Species are compared by their morphology with particular emphasis on
quantitative data as well as from a knowledge of their reproductive methods.
Comparison of general morphology
In preserved species the body shape of the 11 species examined is very
similar and no useful generalizations can be made. Comparison is made difficult
by distortion. This occurred a t fixation in extracted worms and was produced
by the contortions of burrows when the worms were fixed before extracting
them. An assessment of the colour of species is also a difficult criterion for
comparison because of its subjective nature and due to leaching of pigments
when specimens have been preserved. The most promising method of
comparing pigments is the spectrographic one used by Dales (1963). He
compared fresh pigments extracted from D. concharum, D. fimbriata and D.
fe wkesi.
The shape of the nuchal organs and anus were used by Caullery and Mesnil
(1898) and Dehorne (1933) in separating the species D. concharum and D.
caulleryi but have not been of use in separating other species. The shape of the
organ seen through the body wall as well as its opening onto the surface does
not noticeably vary within species due to fixation artifact. The nuchal organ of
278
P. H. GIBSON
D. caullepyi is unlike all other species except D. fimbriata in that it is small and
oval rather than long and slit-like. The nuchal organ in the species examined
reaches its greatest length in D.Zaddi.
Comparison of chaetal shape and distribution
In the general taxonomy of polychaetes, chaetal shape, number and
distribution along the body are important diagnostic characters as is indicated
by their use in numerous identification keys. However, in separating species of
Dodecaceria their use is doubtful. Caullery & Mesnil (1898) found the small
cone at the base of the spoon-shaped depression in the crotchets of D. caulleryi
is absent in D. concharum. However, throughout the species examined this
protruberance is seen to vary in size (Fig. 1).
The numerical distribution of the crotchets and capillary chaetae along the
body has been studied by Caullery & Mesnil (1898) for D. concharum and D.
caulleryi and by Martin (1934) for D. fimbriata and D. coralii. Using the data
from these studies and plotting it graphically (Fig. 2) the numbers of the two
types of chaetae are seen to compliment each other. The distributions for D.
caulleryi and D.fimbriata have three clear peaks in similar positions while there
are two clear peaks for D. concharum and two less distinct peaks for D. coralii.
The relative positions of the peaks can be expressed in absolute terms or as
percentages of body lengths (Table 2). The chaetal distribution might well
prove to be useful in characterizing different species but require a considerable
amount of work to obtain. The essential similarity of the distributions for
species becomes apparent, especially in the case of D. coralii, when the
percentages are used.
Comparison of body length and number o f cirri
In the genus Dodecaceriu the number of chaetose segments and branchial
cirri has been widely used as a diagnostic character probably because of the
absence of different morphological features. A great deal of faith appears to be
placed in the number of branchial cirri inspite of juveniles obviously having
fewer cirri and that the structures are easily lost through damage.
From examining many individuals the number of branchial cirri is seen to
be directly and significantly correlated with the size of the individual (Table 3).
Therefore the number of branchial cirri appears to be a poor criterion of
identification especially in view of the problem of distinguishing between
juveniles and adults. However, an index for the relationship between the
number of branchial cirri and chaetose segments can be calculated for different
species (Fig. 3). The modal point for large individuals in a size distribution of a
sample is taken as a measure of a mature individual (Table 3). The number of
branchial cirri for the individuals at this point is then calculated from a
significant (10% or less probability) regression for the numbers of branchial
cirri and size (Table 3). The data for a number of species lies along a curve
which suggests that the relationship between the size of the species and number
of branchial cirri is a physical one that may well be related to oxygen
consumption.
S Y S T E M A T I C S OF DODECACERIA
279
Figure 1. Photographs of crotchets from eleven species o f Dodecaceria. A, D. berkeleyi; B, D.
cupensis; C , D. cuulleryi; D , D. conchurum; E, D. coralii; P, D. fewkesi; G , D. fimbriutu; H , D.
fistulicolu; 1, D. luddi; J, D. opulens; K, D. pulchra.
Comparison of methods of reproduction
Reproduction in eleven species were investigated and found to be in keeping
with the observations of other authors (Table 4).In addition to the published
work D. berkeleyi was found to have both male and female as well as epitokous
individuals in two small samples (Table 1) of a total of 11 specimens.
D.opulens, in a sample of 82 specimens (Table l), was found to have both
male and female atokous individuals and showed no signs of asexual
reproduction. The possibility of D. opulens and D. fistzilicolu are the same
P. H. GIBSON
280
~1
i
m
.--.
'=I---
----_
0
d
x
w
0
0
s:
0
d
Ic)
0
w
e
. . . . . . . . . . . . . . . .
N
m
P
0 %
N
m
d
0
0
SYSTEMATICS O F DODECACERIA
18
-
-
4 D berkeleyi
I6
-
D coralii
A D fimbriata
D fistulicola
W D laddi
b D opulens
4 D pulchra
14
0
12
B D caulleryi
V D concharum
L
b.
10
28 1
-
8-
6-
4-
2-
o 30
*
1
4''
0
I.....
1
50
60
70
80
90
100
110
120
130
Chaetose segments
Figure 3. Relationship between the number of branchial cirri and body segments for nine
species of Dodecacena. (Data from Table 3 ; data for D. fewkesi included with that for D.
fistulicola; standard deviations indicated.)
species was suggested by Fauvel (1930a, b; 1953) but is, on the basis of their
reproduction (Table 4), out of the question. D. opulens shows no signs of
asexual reproduction while D. fistulicola does. Also D. fistulicola builds
boulder colonies while D. opulens does not appear to.
The existence of asexual reproduction is deduced from recently produced
fragments of individuals in field samples (Dehorne, 193 3 ; Martin, 19 3 3 ;
Berkeley & Berkeley, 1954; Gibson & Clark, 1976; Gibson, 1977). The
regenerated regions are distinguished from the original region or fragment by its
lighter pigmentation. There are three types of individual: those regenerated
from the original anterior region, posterior or pygidial region and one or more
segments from the central region of the body. A fourth type of asexually
produced individual not showing signs of regeneration is derived from the
autotomized regenerate of the single segmental fragments. This individual is
indistinguishable from prefragmentary individuals.
Further fragmentation in the same or subsequent years following regeneration occurs in D. caulleryi (Dehorne, 1933; Gibson & Clark, 1976) and D.
pulchra (Gibson, 1977) and may well occur in other species. These postfragmentary individuals, for the purposes of the present classification are
morphologically identical to those resulting from the first fragmentation.
From the proportion of the various types of individual (Table 5 ) the species
%
t One specimen.
%
Mean
%
Mean
%
Mean
' Two specimens.
n =4
D. concharum
n=4
D. fimbria ta
n=4
D. coralii
n = l
Type of chaetae
Peaks
Axes
_________
D. caulleryi
Mean
11.8
100.0
13.5
100.0
13.0
100.0
18.0
100.0
y
1
11.4
6.0
13.9
5.0
9.5
11.0
9.7
5.0
X
8.0
68.1
5.8
42.6
7.2
55.8
16.0
88.9
Y
2
16.8
38.1
19.5
45.1
18.5
35.1
42.0
37.2
X
Y
3
X
__-__
Y
4
6.0
46.2
7.8
66.0
29.0
55.0
54.5
24.0
2.3
19.1
0.8'
11.1'
2.3
17.3
I
i
Capillary
44.0
100.0
42.0'
100.0'
52.8
100.0
68.0
100.0
X
--
10.3
87.2
10.5
77.8
10.5
80.8
14.0
77.8
Y
1
9.0
20.5
9.0
20.8
11.8
22.3
24.0
21.2
X
5.3
44.7
9.7
72.2
4.7
36.5
Y
2
~
59.7
31.5
19.8
44.9
17.8
40.3
X
Crotchets
i
_
4.7
40.4
5.0t
35.7t
3.8
22.8
_
Y
_
3
_
X
_
29.0
65.9
40.0t
100.0t
46.3
87.7
_
Table 2. Coordinates for the peaks of the curves for the chaetal distributions (see Fig. 1 ) for four species of Dodecaceria
Z
_
.cr
_
02
N
N
8
4
169
39
40
5
( 69
86
14
82
102
n
3.5
4.3
2.9
2.9
8.2
3.7
4.2
5.7
2.4
12.2
4.3
P
-
80
107
42
40
96
59
56
96
49
106
79
X
1.25
0.43
1.07
1.31
2.44
0.40
1.19
1.31
1.57
3.84
0.61
OY
19.8
6.2
10.9
13.8
20.8
4.6
13.3
17.1
12.2
26.6
10.6
ux
0.274
2.678
-0.356
-0.441
-1.057
2.363
1.056
2.114
-2.425
-0.814
2.965
b
m
0.041
0.015
0.077
0.083
0.096
0.023
0.056
0.037
0.098
0.122
0.017
~~~
0.656
0.210
0.791
0.882
0.823
0.262
0.625
0.486
0.758
0.834
0.296
r
0.1
1 .o
0.1
1 .o
0.1
10.0
Not sig.
0.1
0.1
0.1
Not sig.
% sip.
3.8
3.9
4.1
8.1
4.1)
5.5
2.0
14.4
4.2
55
55
95
55
95
45
125
75
PC
85
ss
n, Number of specimens examined;.y,, mean number of branchial cirri; 2, mean number of body segments; uy, standard
deviation for the number of branchial cirri; ux, standard deviation for the number of body segments; b, slope of the regression
line; m, intercept of the regression line; r, correlation coefficient; sig., the significance of r; ss, number of body segments found
from the modal point for the largest individuals in their size distribution; pc, the number of paired branchial cirri calculated for
ss from the regression.
t Including D. fewkesi.
* Data for figures in brackets taken from Martin (1934).
D. fistulicolat
D. laddi
D. opulens
D. pulchra
D. berkeleyi
D. capensis
D. caulleryi
D. concharum
D. coralii
D. fimbriata *
Species
Table 3. Relationship between the number of branchial cirri and body segments for ten species of
Dodoecuceria. Linear regression and correlation for the number of branchial cirri and body
segments calculated. Size of mature individuals found from the frequency distributions of the
number of body segments
N
W
m
284
P. H. GIBSON
Table 4.Types of reproduction in nine species of Dodecuceriu examined in the
present paper (time of collecting and location of samples given in Table 1)
'pecies
D. laddi
D. berkeleyi
D. caulleryi
D. f im briata
D. opulens
D. f e wkesi
D. coralii
D. puichra
D. concharum
Epitoke
'E
$toke
Parthenogenetic ( P ) Asexual
+
+'
+
+
+
+
+
+
+
+
+
+
+
+
+
Published work
Reish, 1968
Knox, 1971
Caullery & Mesnil, 1898
Gibson & Clark, 1976
Martin, 1 9 3 3
Present work
Berkeley & Berkeley, 1954
Martin, 1933
Gibson, 1976
Caullery & Mesnil, 1977
Gibson & Clark, 1976
' Reported here for the first time.
Table 5 . Types of individual found in seven asexually reproducing species of
Dodecaceria examined
Species
D. berkeleyi
D. capensis
D. caulleryi'
D. f e wkesi
D. fimbriata
D. fistulicola
D. pulchra
Total number
11
7
180
37
5
23
268
% posterior
X anterior
% segmental % no. apparent asex. reprod.
18
71
36
14
47
78
9
15
23
13
65
46
60
19
22
5
18
45
30
40
11
14
Posterior individuals not found at the time of year sampled (Gibson & Clark, 1976); time o f
collecting and location of samples given in Table 1 .
can be divided into two groups. The first comprises D. fewkesi, D. capensis and
D. fistulicolu which have less than 20% anterior individuals and appear to be
atokous. The second comprises D. pulchra, D. fimbriata, D. caullervi and D.
berkeleyi which have greater than 20% anterior individuals and, with the
exception of D. pulchra, all have epitokes. The two groups are not necessarily
fundamentally different but may merely reflect the variation in asexual
reproduction in the genus. Though the proportions of the different types of
individual within the genus (Table 5 ) do not give conclusive evidence of their
taxonomic relationships they do indicate how a more detailed study might do
so.
DISCUSSION
Original descriptions of the general morphology of species of Dodecuceriu
are of little diagnostic use because the species are morphologically so similar.
Also, the use of the number of branchial cirri and the point at which the
crotchets first appear along the body can be seen from a brief literature review
by Knox (1971) to be of little descriptive use. In fact the general confusion in
the literature is good evidence of the difficulty of giving satisfactory
descriptions. There is still uncertainty over whether the following species are
SYSTEMATICS O F DODECACERIA
285
identical: D. cuulleryilD. fimbriutu (Fauvel, 1927; Dehorne, 1933 ; Martin,
1933), D. fistulicolulD. fewkesi (Reish, 1952; Hartman, 1959; Knox, 1971)
and D. fistulicolulD. joubinilD. opulens (Augener, 1914; Mesnil & Fauvel,
1939).
In the present work general morphological descriptions of Dodecuceriu are
difficult to make because specimens became badly contorted at death. The use
of chaetae is limited because they are structurally so similar. Reliable data on
numbers of chaetae, segments and branchial cirri is ambiguous as well as
difficult to obtain since many specimens are required. However, though the
physical aspects of species are so similar there are many marked differences in
their methods of reproduction.
Differences in reproduction between the morphological similar species D.
conchurum and D. cuulleryi were used by Dehorne (1933) to separate them.
These two species are exceptions in that they coexist in the same niche and
therefore the original assumption was that they are the same species. The
reverse is true for D. cuulleryi and D. fimbriutu which occur on opposite sides
of the Atlantic and therefore are assumed to be different species. However,
morphologically as well as reproductively they appear to be the same. Probably
this is true for D. fewkesi and D. fistulicola which have a cosmopolitan
distribution. An aspect of the morphological similarity of D. cuulleryi and D.
fimbriutu and to a lesser extent D. conchurum is seen from the similarity in
numbers of body segments and branchial cirri in the species (Fig. 3).
The two species D. diceriu and D. multifiligeriu and the subspecies D. luddi
oculutu were not examined in the present work but need to be commented on.
From their descriptions they do not, on morphological grounds alone, fit into
the genus. Both D. luddi oculutu (Hartmann-Schroder, 1962a) and D.
multifiligeria (Hartmann-Schroder, 1962b) are peculiar in having eyes in the
atokous condition. D. diceria (Hartman, 1951) does not have typical
spoon-shaped crotchets although, as noted, the crotchets are not an infallible
guide. D. multifiligeriu, however, is atypical in having a large number (over 20)
of branchial cirri and the paired tentacles appear to arise dorsally instead of
ventrally to the first pair of branchial cirri. Also, the stout chaetae are pectinate
and in no way resemble the crotchets of other species of Dodecuceriu. The
existence of eyes in D. luddi oculutu and its small size suggests that the species
is a juvenile of D. luddi since juveniles of D. conchurum have eyes which are
lost with further growth.
An explanation for the morphological similarity of the species of Dodecuceriu
is difficult to find. The ostensible reason would appear to be that the habitat is
physically undemanding. If the majority of species are secondary borers, and
take over already existing burrows of a variety of primary borers, as appears to
be so for D. cuulleryi (Gibson & Clark, 1976), a comparatively unspecialized
condition may be an advantage. The reproductive phase of the species has
become specialized for colonizing niches largely through cloning by asexual
reproduction. Another factor contributing to the similarity of species may well
be evolution of the genus through polyploidy.
The possible role played by polyploidy is illustrated by an explanation of
how D. conchurum and D. cuulleryi come to inhabit the same niche. I t may
well be that D. concharum arose as a polyploid of D. cuulleryi for asexual
reproduction in D. cuulleryi and parthenogenetic reproduction in D. con-
286
P. H. GIBSON
charum support the idea. Parthenogenetic reproduction is unusual in animals
except in the case of polyploids (Muldal, 1952). Asexual reproduction in D.
caulleryi would undoubtedly assist in propagating any parthenogenetic mutant.
Since asexual reproduction is common in the genus and at least two species are
known to be parthenogenetic polyploidy may also exist and account for the
lack of morphological diversity.
ACKNOWLEDGEMENTS
I wish to thank the numerous people who have been so kind as to send me
samples of Dodecaceria and Marian Pettibone in particular for encouragement.
I also wish to thank Liz Fording for typing the manuscript.
REFERENCES
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