SIPUNCULIDS OF MADAGASCAR
by
Edward
B. CUTLER
*
ABST RACT
RESUME
Cetarticle
Sipunculides
Madagascar.
pendant
nationale
sa participation
dans l’Océan
à I’Expédition
Indien,
du 24
10 septembre
1964. Neuf espèces
‘indicus,
: Sipun.culus
gistrées
Siphonosoma
losoma
This
est un compte-rendu
sur les
dans
la région
de Nosy - Bé,
par 1’ auteur
II s ont été réunis
culids
They
‘his
Interjuin au
pour la science,
nouvelles
pour
et Cleosiphan
espèces
n’est
are
total
thirteen
of
presented
présentée
en supplément
sion relative
à chaque
trée
des
13 espèces
commensales
à une brève discusespèce.
Une clé illusconnues
est
during
Indian
24 June to 10 Septemspecies
recorded
S. robustus,
new
to
this
:
Siphonosoma
and
species.
An
known
species
thirteen
now
niAspi-
making
known
of their
commensal
in addition
of each
country,
species
A discussion’
preference
est
author
coralficola,
and Cleasiphon
‘asperNone of these
are new to science
gillum.
but five
préféré
relations
the
sipun-
Madagascar.
Phascolosoma
S. australe,
P. scolops,
P. dentigerum,
Madagascar.
leurs
of the
Be,
International,
are nine
indicus,
dosiphon
mais cinq d’entre
elles
sont
cette
région,
faisant
un total
from
There
cumanense,
grescens,.
aspernouvelle
by
in the
Expedition
de 13 espèces
maintenant
connues
à Mada‘Une discussion
quant à leur habitat
gascar.
et
collected
Sipunculus
cumanense,
S. australe,
PhoscoP. scolops,
P. dentigerum,
Aspidosiphon
corallicola
gillunr.
Aucune
de ces
were
.ber 1964.
nigrescens,
is a report
of Nosy
participation
Ocean
y sont enreS. robustus,
paper
in the region
a
from
habitat
relationships
is
to a brief
discussion
illustrated
key
is
also
to the
included.
également
jointe.
INTRODUCTION
The
information
presented
in the U .S. Program
in
specimens
were collected
24 July
R/V
1964.
{(Anton
‘A
few
specimens
sipunculid
(Wesenberg-Lund
1959),
1958,
,1962),
South
1955,
1956).
added
two
more to this
mum
signifer)
was
of the
vicinity
were
Africa
In Herubel
(1908)
list,
in poor
Deportment University
each
is based
International
of Nosy Bé;
(Lanche-ster
from
Madagascar,
species
In the
surrounding
1905;
1942;
represented
condition.
I made
as a participant
Indian
Ocean
Expedition.
Most of the
Madagascar,
between
24 June 1964’and
at Tuléar,
described
(Stephens
six
on collections
during
a stopover
of the
1964.
bas’ been
Zanzibar
paper
collected
8-10 August
fauna
(Kalk
* Zoology
Biology
in the
Bruun B from
The
in this
are
Stephens
areqs
& Robertson
Wesenberg-Lund
listed
from
1963),
one specimen
material
l collected
1):
1952),
Mauritius
Mozambique
and Australia
Madagascar.
by only
of Rhode Island Kingston,
(Figure
(Edmonds
Hammerstein
(1915)
and one of.these(Dendrostowere
nine
species,
none
new
Rhode Island U.S.A.
51
to sci.ence
but five
in the U.S. National
previously.unreported
Museum,
Woshington,
LIST
OF
MADAGASCAR
Sipunculus
Sipunculus
PP
indicus
robustus
collection
has
been
deposited
SIPUNCULIDS
- He, PP
- pp
Siphonosoma
cumanense
Siphonosoma
Phasco/osoma
Phascolosoma
australe
- PP
nigrescens
- He, Ha, PP
scolops
- He, Ha, PP
- l-le,
Phascolosoma
dentigerum
Phascolosoma
Phasco/osoma
asser
- t-le
lobostomum
Dendrostomum
signifer
Aspidosiphon
truncatus
Aspidosiphon
corallicola
Cleosiphon
He
Ha
The
from Madagascar.
D.C.
PP
- PP
- Ha
- Ha
aspergillum
* l-le,
Ha
- pp
- pp
- Herubel
(1908)
- Hammerstein
(1915)
the
present,
paper
-
Fig.
1 - The
been
southwestern
Indian
Ocean
showing
localities
where
sipunculids
have
col lected.
THE AREAS AND STATIONS
The
480
15’ E
island
(Figure
I’Océanographie
et
Teahnique
Madagascar.
52
of Nosy Bé
Located
1).
et des Pêches
This
Outre-Mer.
lies off the northwestern
on the southeastern
which
served
toast
of Madagascar
corner
of the island
is operated
by the
as the hoadquartors
at 130 20’ Sand
is.the
Centre
de
Office
de la Recherche
Scientifique
for the American
IIOE program
on
season
The
climate
from
May
is tropical
with
a warm
The
total
to September.
in January
of 462 mm and a minimum
in November
to 15.00C in June. The
rainy
average
230
is 2244
E.
Along
the
shore
fringing
reef which
is partially
exposed
collections
were not extensive.
The stations
where
collections
is a broad
at low
were
muddy
tide.
made
and a cool
a maximum
ranges
25.20C
dry
recorded
from 34.80C
in August-
As
are
sand
time
swamps.
of the Tropic
flat
and off
and equipment
described
of Capricorn
shore
were
in Appendix
lies
limited
I and
at
a large
these
shown
in
2.
Fig.
48.O’E
2 - Map of Nosy
Be, Madagascar,
this study were
Appendix
1).
ECOLOGICAL
A clear-tut
species.
showing
made.,(For
stations
the descriptions
where
collections
of the stations
for
see
RELATIONSHIPS
pattern
of habitat
summarized
in Table
1 and Appendix
Most of the coral formations
four
to April
mm with
salinity
from 35.57 O/oo in August-October
to
is 2.2 m but may increase
to 4.4 m at‘spring
of habitats
within
easy reach of the Centre
Fis.2
of
October
from coral islands
through
sand and mud flats to mangrove
Tulear
is about 900 miles
south of Nosy
Be just north
20’ S and 430 40’
Figure
from
of 37 mm in July.
Th e air temperature
surface
water
temperature
ranges
from
in January-February
and the
September
to 31.30C
29.05 oho in January.
The average
tide differential
tides
(Centre
records).
There
is a wide variety
ranging
season
rainfall
The
most
numerous
preference
emerged
from
this
study.
1, I was able to make the following
(Stations
10,14,16,
17 and 20) were
and
cosmopoliton
was
From
the
information
observations.
inhabited
by a complex
Phascolosomo nigrescens. The
53
other
Phascolosoma
also
found
fact
that
the
in the
one
smaII
wheret
experiment
inches
of sand
the
the
second
four
leads
(which
sipunculids
(Rocellaria
and
A
at
the
later,
more
white
Harvard
numerous.
bivalves.
that
end,
it was
no experimentation
reasonable
idea
were
and the
seems
The
coral
they
were
was
was
members
relationship
in the Sand.
In addition
crustaceans,
present
two
theTubiporo
itself
to
pelecypods
at these
to be obligatory,
of habitat
was
were
low
tide
mixing
members
preference
and
exists
stations.
between
the
mollusc
is sometimes
reported
by Bovier
(1952).
(cf)
In this
rouseanus,
instance
the
forme,r
being
has
that
to both
to more
but
apparently.well
Young
being
given.
worm
the worm
(provides
length
inhabits
an
protection)
habitats).
of Sipunculus
at
at
The, most
subsequentlyabsorbed
desireable
species
points
discussed
been
the
the shell
two
been
answer
and probably
both
(S. ,indicus
and
(12 and
13b)
stations
oxygenated,
coarse
Sand.
There
two.populations.
of Siphonosoma
was
.by the
separated
white,
genus Jousseaumiella
suggestion
locomotion
preferred
clean,
into
by the coral,
at widely
of these
of the
to be.beneficial
type
two
buried
Tubiporo.
or
following
containing
way,into
away)
an indisputable
(1902)
seems
found
has
covered
They
a very
The
Sluiter’s
is then
cornes
nor
and provides
no apparent
s ubstrate
to ‘be
which
done
it upright
A third
there
been
(keeps
S. robustus).
rock-like
construction
in an aquarium
and Stephens
and Robertson
(1902),
(cf) geminale
and Heterocyanthus
produced
by the
in the
intide
from thetwo
a hard,
supplied
A bivalve
coral.
has been previously
The
indicates
have
own
its
were
5.
rocks
coral
prefer
their
and tunicates
appears
under
would
polychaetes,
holothurians,
which
in the
worked
located
numerous
found
was
placed
were
specimens
were
species
spp.
at Station
found
of
One worm
genera
of solitary
This
association
decided
has
shell
or dissolved.
four
as Cleosiphon
On the trunk of two of the worms withinHeteropsammiathere
were several
After
conferring
with Dr. W.J. Clench,
Museum
of Comparative
Zoology,
University,
emptygastropod
these
well
Sand)
collecting
5. or 6 inches
J. heteropsommia
(familly
Montacutidae).
The question
of how this relationship
but
was
5 were
individuals
relationship,
and two
‘complex.
specimens
extensive
as
Phoscolosoma
sixteen
either
and cracks,
Additional
evidence
other
there
(1894),
Boume c (1906),
Sluiter
the corals
were Heteropsammia
being
small
more
Station
ophiuroids,
commensal
Aspidosiphon
corallicola
a third
member
of this
A few
yielded
P. nigrescens
at one end.
both
mentioned
Lithophaga),
unique
muddy
holes
of Tubipora
was
hours
indurated,
to the conclusion
in
here
(hardened,
Possibly
4).
found
The
25 cm in diameter
of other
organisms.
P. kigrescens
from
six
were
in Millipora.
material.
Only
cari -live
piece
scolops)
found
3).
2 and
with’a
checked
about
in this
they
works
Two
:
clay
(Station
information
environment
When.
density
(Stations
forms.
This
abandoned
material
P.
were
rock-like
same’ area
islands
other
of
and
Cleosiphon
peculiar
high
in this
dentigerum
of the
mass
unusually
p.ools
and
(P.
Most
asperigi//“m.
were
also
segregated-from
one another,
but a different
apparent.
All of the S. cumanense
came from a rather
homogenous,
muddy
sand (Stations
‘6, 11, 13, 15, 19).
The 3. australe
were found
in a more heterogenous
substrate,
mostly
firm,
muddy
sand but also
containing
Iarger
particles.
such as pebbles,
mangrove
roots,
etc. (Stations
8, 9).
It would
have been illustrative
had. I made a transect
between
Stations
6 and 8 to see if there was a distinct
separation
or how. much overlap
species
occurred
These
were.exposed
and
54
Siphonosoma
with
Sipunculus)
laboratory
regard
at the border
almost
to at least
stations
three
one
all
IOWtide.
every
is interesting
experiments
of
area.
because
could
parameters
predict
differed
This
of the
that
: temperature,
from
the
information
T,ipuncu/us
about
physiological
the
two
in
are
and oxygen
that
the
former
genera(Jiphonosoma
implications.
Siphonosoma
salinity,
stations
these
much
(Figure
Without
doing
more tolerant
3) .
any
with
Fiq. 3 relafive
tolerance
Hypothetical
for Sipunculus
.
.
I.
. 9’
.
.
.
.
I.
.
.
.
.
.
Low
Low
Salinity
Oxygen
Low
This
The
also
has
does
have
evolutionary
genetic
is selection
mechanism
evolutionqry
of a capacity
significance
adaptation
a primary
interesting
more
of the
capacity
upon
(Kinne
implications
- phylogenetic
to acclimate(as
acting
of evolution
l)
l
High
.
.
.
.
.
.
I.
.
Temperature
(0
High
.
High
.
Siphonosomo
limits
) and Siphonosoma
(-
reflected
to acclimate
genetic
1963),
is based
Therefore,
variation.
i.e.
the
by wider
more
if
it
is
true
tolerance
on the assumptian
genetic
adaptation
capacity’to
acclimate,
that
limits).
the
that
is
more
genetic
adaptation
which
is, in a sense,
evolution.
.As yet, within
the sipunculids,
no attempt
has been made to establish
any taxa befween
genus and phylum.
.The foregoing
ideas maybe
helpful
when such an attempf
is made,
Habitats
which
consistantly
yielded
no specimens
were those areas composed
of soft,
fine,
silty
of any
mud.
This
probably
was
a very
unstable
situation
and net
suitable
for worm
burrows
sort.
P.S.,
ca
I* Coral,
rock,
hard
clay
Phascolosoma
scoleps,
P. nigrescens,
P. dentigerum
and Cleosiphon aspergillum
II, Within base of solitary
Aspidosiphon
III.
Clean,
corals
il
corallicola
white,
coarse
III
Sand; exposed
IV
rarely
Sipunculus
inbicus
IV. Homogenous,
muddy
V
and S. robustus
Sand; exposed
regu larly
Siphonosoma
V. Firm,
muddy
stones,
roots,
Siphonosoma
cumanense
sand with
PICTORIAL
SUMMARY
OF
,pebbles,
etc.
HABITAT
PREFERENCE
australe
Fig.
4 - Diagramatic,
pictoral
,summary
of habitat
preference
55
DISCUSSION
OF
Sipunculus
SPEÇIES
indices
PETERS,
1850
ranged
8 specimens.
Length
These
animals
were
There
are many
interesting
from
very
165-510mm
abundcint
simi larities
at
;. diameter
this
between
7-l 1 ‘mm;
station
but
the situation
color,
white.
apparently
quite
restricted.
here and that reported
on Zanzibar
by Stephen
the animats
and Robertson
(1?52).
for bait by inserting
When we arrived
at Antsakoabe,
there was a fisherman
collecting
straight
stick,
which
was (tburreda
on the end, into
a square,
the
and
proper
burrow
exerting
just
the
introvert
it pulled
the stick
Sand next to the worm with
pulled
it out more or less.
this.manner,
Sipunculos
finally
robustus
1 specimen.
This
very
gave
low
KEFERSTEIN,
Size 185mm
individual
was
to a better
I was
unable
After
returning
produce,
This
Siphonosoma
cumanense
These
(1955)
uses
epidermal
out that
are
((skin
present
these
as well-defined.(<warts
off.
S. angasi
the
have
misleading
to o person
species
I suggest
skin
bodies,
An
dug
out,
to
thot
or epidermot
interesting
pinched
look
and
frein
)) that
but
but
was
the
dark
One
brown,
damaged
noteworthy
others
specimen
remain
after
layer
them
on the
bicalve
the
of autotomy
posterior
the
underlying
looke,d
was
causing
the
.It did
postefior’
organs
color,
quite
muscle
is removed
there
smooth-skinned
feoture
mottled,
occurred
damaged
near
pinkish
well.
the
Edmonds
It should
be
which
protrude
are
stripped
layers
is left
when
multicellulor
only
CI thin
piece
themselves
remoin ottached
protuberances,
it could be
Siphonosomo.
glands
when
quarter
be used
Therefore,
rother
than
in
papillae,
individual,
trunk
within
but no other
damaged
whi.le
being
24 hours
after
it wos
organs.
I have
been
1865)
from
45-210mm
sf this group
some u,niform
almost
by byssal
very
one
of the
port of the digestive
tract
of this for the sipunculids.
Length
ronged
rows of hooks.
had’on
Attached
which
tide
active.
from
in the Phascolosoma
epidermal
white
trunk
is the
yellow
similar
threads
to
(overoge
the
color,
yellow
(strow)
the
brown
introvert.
On examination
probably
due to the onimols’
environment.
bluish-grey.color
similar
to Siphonosomo
posteri.or
to the Jousseaumielle
105mm);
voriability
of color.
Some were a uniform
with
a darker
introvert,
and one,
smoll,
but retoined
this coior proved to be o tronsient,
superficial
loyer,
This
color cari be rubbed
off exposing
an irridescent
Umonense.
the
very
tufted
12Omm);
descriptions
the.papillae
multicellular
oustra/e(KEFERSTEIN,
17 specimens.
brown;
50-20
to dark
the
orgons.
collected.
This portion
contoined
unable to.find
any previous
record
Siphonosomo
not
coming
lacks
(average
published
the muscle
for
t&m
example
off
As
ihtolerable
)) and (tpapillae))
as synonyms
to describle
the
Gerould
(‘l9’13) uses (tepidermal
organs w.
form.
when
the
Sand.
bec.ome
of skin with holes which
were the pores of the glands.
The glands
to the muscle
‘loyer.
As the term papillae
suggests
well-developed
this
retracted
dug down in the
a firm grasp cind
to obtain
any in
1866)
d,ifferent
however,
Baird,
fit
quite
of
might
of it apparently
common
in this
surface
in an aquarium
much
35-245mm
)) or «knobs
In Siphonosomo,
the
micro-habitat
ranged
from
preserved.
bodies
are
grey.
a.long
placed
resembles
individuals
glands
pointed
as the animai
from the fisherman.
color,
of .mucus,
(KEFERSTEIN,
the terms
SO that
Dr. Humes had seen theni there on previous
occasions,
found,
it was covered
with a layer of Sand and mucus.
it was
amounts
closely
its
Although
When
Length
54 specimens.
dirty white to pale yellow
alive,
day,
laboratory
copius
species
movinb
hot that
any’more.
to the
of pressure
some
1865
X 13 mm ;
found
location.
to locate
however,
bulb.
rectum.
up, and.purchased
and the sun was
it to move
amount
down inside
immobilizing
itself.
The-native
then
his hand, loosened
the, Sand around
the worm, got
Even though
it looked
easy- I was unable
intact.
end
found
of one
specimen
was
on the Aspidosiphon(page5).
o small
Phascolosomo
nigrescens
KEFERSTEIN
sensu
STEPHEN
Because
of the varied
states
of contraction
44 specimens.
The
opproximote
range
is
from
25-120mm,
not very
meaningful.
The
color
varied
length.
alternating
dark and
easi ly removed.
This
species
There
authors.
distinction
light
between
as the hooks,
two
species
are
cross
dark
exist,
in this
P. ,nigrescens
same
very
pale
region
and
deol
ond
but
the
has
of variation
organs,
(Edmonds
1956,
page
289;
was
been
in the
are
40-80mm
some
skin
P. puntorenoe
is not olways
etc. are quite variable
segmental
measurements
between
with,
In o few
of the world
a good
size
most
most
brown;
on the. introvert.
however,
popillae,
the
to
bonds
is common
does
such
from
the
in
without,
very
reported
the
loose
and
by several
descriptions,
and
the
cleor..
Many structures
ond it may be that these
Wesenberg-Lund
1963,
poge
129 and
Wesenberg-Lund’s
(1963)
description
of P. nigrescens
table
1 ; Fisher
,1952, page 430).
particulorly
confusing
for several
reosons and seems to more closely
agree -with other descriptions
of P. puntorenoe
‘based
on Dr.
(e.g.
caecum
Stephen’s
and controctile
criteria,
vesse1
Selenka’s
(1883)
both
species
from the United States
National
One charocteristic
which
is sometimes
tion
that
this
might
be important
villi).
The
f’gI ure ,130,
moterial
and
Museum as identified
reported
is the hook
l meosured
the
height
I hove
is P. nigrescens
comparison
to specimens
by W.K. Fisher.
dimensions.
On the
ond width
of
is
152 hooks
of
assump-
from
thirteen
P. nigrescens
first
it looked
ond fifteen
hooks’ from the one P. puntorenoe
(Figure
as though there was a positive
correlation
between
body
as I measured
anything
more
more. hooks,
the correlation
wos not always
consistent.
It is’ difficult
to say
thon:
as the hook size of P. nigrescens
is variable
it should
not be used as a
taxonomie
width
character.
of the hook
ln rrcost
(at the base)
of the
specimens,
is between
which
are
between
5 ond Appendix
3). At
size and hook size,
but
40 ond
80mm
long,
the
70 ond 80 p.
tjtt.
E
;o
58
90
SIZE
OFANIMAIS
- mm
Fig.
where
These
anolysis
distinction
form.
Africa
reported,
size
the two
should
of Phoscolosomo
ond
be made
nigrescens
P. puntorenoe
are o good
the world
should
be carried
or the two
should
be combined
scolops(SELENKA
ond DE MAN,
1883)
two specimens
(60.and 15 mm respectively)
represent
It is perhaps
coost
hook
two forms,
P. nigrescens
of specimens
from.around
between
PhoFcolosomo
These
5 - The
they
however,
noteworthy
generally
from
thot
comprise
Mauritius.
only
two
a large
This
were
found.
percentage
apparent
decrease
onother
example
out.
of a situation
Eithcr
a better
as one species.
common
Indo-Pocific
On .the Zanzibar-Mozambique-South
of the total.
They
have not
in frequency
with
distance
been
from
57
This
species,
together
with
be significant.
complex
of Phoscolosomo
with very similor
This
species
differs
from
minor differences.
P. ogossizi,
the continent
may
others
form another
exhibiting
only
a rectal
caecum.
The’noture
of the papillae
Phoscolosomo
dentigerum(SELENKA
20 specimens.
The trunks
characterize
this
Aspidosiphon
corollicola
SLUITER,
I am indebted
Th e anal
of the solitary
corals.
dark
red-brown
and
granules
anteriorly.
somewhat
becoming
in size
Cleosiphon
end
trunk
Figure
216.
difficult
KEY
TQ
THE
1 a. Horny
1 b. Above
6 b.
the
Stephen’s
(1952)
The
The
tenter.
the
the
size
were
their
which
to obtoin
Skin
weak
1865)
on ivory,color
.Hooks
introvert
description
‘very
well.
protrudes.
They
-on the
hook
ranged
from
30-75mm.
were
most
obundant
as they
BF
a, few
were
cap
a rather
or knob
streak
Introvert
muscle
skin
on introvert
is even
SIPUNCULIDS
OF
dark
The
average
in very
was
......
merged
of the trunk
Edmonds
in Selenka
48mm.
...............
reddish
bands
Introvert
locking
hooks.
pineapple-like
2
10
triangular
and tall,
cleor
conical,
...........
Grooves
shields
of anal
at both
ends ,of trunk.
............................
shield
run from
ventral
spoce
preanal
6
(Figure
(Figure
papilloe
Phosyolosomo
6-4).
.........
caudal
to margin
osser
Phoscolosomo
nigrescens
6-5 & 6). .......
8
oriented
somewhat
posteri-
scolo
Phoscolosomo
dentigerum
Phoscolosomo
Siphonosomo
australe
Siphonosoma
end of trunk
edge
indicus
.........
9
spaie
; ............
The
4
; ..
5
robustus
7
....................
..........................
cap at the anterior
3
12
...........................
(Figure
.........................................
10 b. Corrugoted
6-2 & 3)
..........
from
These
Mi//iporo.
hard
Sipuncu/us
(Figure
.: ..................................
; ........................
with basal,
triongular
clear
..‘..................L
of hook separate
with
absent.
brown.
anterror
MADAGASCAR
bonds
......................
up of numerous
platelets
papilloe
...
hooks
of hook
are
is quite
at the
The hooks differ
slightlyfrom
more strongly developed as
of the trunk
made
and complex
without
bases
changing
to rounded
but a rather
pale tan
tooth
ocorn-like
tooth
SPECIES
complex;
present
Introvert
10 a. A white
58
out to me in the
to the.anus
however,
but
compound,
white,
orly (Figure
6-7).
..............................
8 b. Papillae
smoller
and not os above
9 a. Introvert
with hooks ....................
11 a.
these
secondary
secondory
intact
longitudinal
papillae
..........
7 b. Cleor streak
9 6.
fit
pointing
.......................
Longitudinal
muscles
gathered
into a number of bands .......................
Longitudinal
muscles
form a continuous
sheath
..............
.’ ...........
Skin divided
into numerous
rectangles
or squares
(Figure
6-l).
...............
Skin not divided
into rectangles
or squares
.........................
24-30 strong
longitudinal
muscle
bonds
Sipunculus
....................
7 a. Clear
8 a.
towards
with
in that
KNOWN
5 b. Protruding
6a.
shields
for
or colcoreous
shields
absent
from both ends
shields
present
at one or both ends of trunk
4 b. 41-43
5 a.
P. joponicum.
DE MAN,
in length
1902
to Dr. M.. Pichon
(QUATREFAGES
Most of these
through
21)
very
3 a;
3 b.
4 a.
it from
present.
animals
Figure
were
2 a.
2 b.
darker
but usuolly
unique
of the
(1883)
separates
composed
of longitudinal
furrows
next
The caudal
shield
is not deep red-brown,
ospergillum
15 specimens.
are
(1956,
hooks and the internol
anotomy
P. ogossizi
by the absence
of
species.
11 specimens.
variable
ond
ranged
and
1883)
from 20-52mm.
The expanded
introverts
The lorgest
individu01
was 77mm
at Iéost half as long and some as long as the body.
Most of these individuats
hod a very pole body with the tall,
conical,
preanal
papillae
posteriorly
and the introvert
showing
the reddishbrown
bonds
or blotches
which
were all
overoll.
oriented
These
platelets
P. joponicum
shield
cumonense
6-8)
...............
Cleosiphon
asperigillum
is pale with,rodiating
grooves
.: ..................
Aspidosiphon
. : ......
11
truncofus
1. Generalized
2-3.
appearance
Sipunculus.
Two views
covered
.4. Hook
merged with
5-6. Phascolosoma
complex
Fig.
6 - Morphological
the known
features
sipunculids
surface
,in
numerous
small
triangular
hook
platelets.
nigrescens;
clear
clear space.
with
separate
streak
triangular
space (P. scolops
and P. dentigerum).
of introvert
of Phascolosoma
dentigerum
showing
8. Anterior
9. Anal
body
ofa complexf~ascolosoma~papillae
with
of Phascolosoma
clear
7. Base
of
talli
conical,
pre-anal
papillae.
end of Cleosiphon
aspergillum
showing
“cap”.
shield
to assist
of Aspidosiphon
corallicola.
in identifying
of Madagascar.
59
longitudinal
grooves
posteriorly,
11 b. Anal
shield
with
two distinct
areas;
introvert
(Figure
6-9) . . . . . . . . . . . ~ . . . . . . . . . . . . . . . . Aspidosiphon
12 a. Tentacles
dendritic.
12.b.
filiform
Tentacles
granules
near
corallicola
. . . . . . . . . , . . , . . . . . . . . . . . . . . . . . . . . Themiste
signifer
(formerly
Dendrostomum)
or digitiform;
complicated
hooks
present.
. . . : Fisherana
lobostomum
(formerly
Phascolosoma)
ACKNOWLEDGMENTS
The
author
is especially
indebted
Edinburgh,
for several
Museum,
the director
and Dr. M, Pichon
American
Univers ity,
the National
Island,
liason
Science
kinds
for
their
officer
Foundation
for assistance
at
for
in preparing
to the
following:
of assistance;
cooperation
Nosy
financial
Be,
Dr.
the
for
A.C.
§tephen,
personnel
at the
and
facilities;
Dr.
his
suggestions
support;
and
Dr.
A.G.
and
D.J.
Royal
COPN,
H.umes,
help
Zinn,
Scottish
particularly
Boston
in collecting;
University
of Rhode
the manuscript.
BIBLIOGRAPHY
BOURNE,
G.C.1906. On Jousseoumia
in Report
on the
Manaar
(Ceylon),
edited
by W.A. Herdman.
The Royal
1907.
Note
to correct
the name Jousseoumia.
BOVIER.
1894. Un nouveau
polypiers
madreporaires
EDMONDS,
S.J,-
1955.
Siphonosoma.
Sipunculoidea,
J. Mar.
and
1956. 1 Australion
Sipunculoidea,.ll.
Aspidosiphon,
and Cleosiphon.
21 textfigs.
On Sipunculus
1961.
4: 217.
FISHER,
Nat.
aeneus
Austral.
Baird
W.K.1952. ‘The Sipunculid
Mus. 192 : 371-450,
22 pls.
Res.
E.
and
Kiobenhavn
OERSTED,
for
AS.-
(Sipunculoidea).
worms
1859.
of California.
eastern
Annulata
Madagascar,
HERUBEL,
les
M.A.-
KALK,
M.14:
60
1958.
189.
London.
genera
,6:
Sipunculus,
82-97,
1 pl.,
260.
avec
CX.IX
des
: 95.
Xenosiphon,
and
9 textfigs.
Dendrostomum,
Golfingia,
Res.
7:’ 281-315,
3 pls.,
Ann.
and
Mag.
and
Baja
California.
toast
7:
of North
Oerstediana.
Vid.
,Nat.
Hist.
America.
Medd.
Ser.
13;
Proc.
U. S.
Proc.
U. S.
naturh.
Foren.
1858.
HAMMERSTEIN,
O.- 1915.
Gephy reen van
Arkiv
for ZOO~. 9 (10):
l-3,
1 fig.
20 : 107-418,
Soc.
*Th e genera Phascolosoma
J. Mar. and Freshwater
GEROULD,
J.H.1913.
The sipunculids
of the
Nat. Mus. 44:
373-437,
5 pis.,
16 figs.
GRUBE,
Malacological
1. The
Freshwater
fisheries
of the Gulf of
Part 5 : 243-266,
3 pls.
d’Aspidosiphon
: association
C.R.
Acad.
SC. Paris
cas de commensalisme
et un Mollusque
bivalve.
Australian
Austral.
Proc.
Pearl oyster
Soc. London.
1908.
Recherches
pls.
5-10.
The
intertidal
fauna
sur
of rocks
sipunculides.
at Inhaca
gesammelt
von
Mem.
Island,
Soc.
W. Kaudern
ZOO~.
Mozambique.
191111912.
France.
Ann.
Natal
1907.
Mus.
W.- 1865.
B er‘t rage zur anatomischen
Zeitschr.
f. wiss.
Zool.. 15:‘404-445.
KEFERSTEIN,
culoiden.
KINNE,
O.- 1963.
Theeffectsoftemperature
In 0 ceanography
1. Temperature.
LANCHESTER,W.E.tions
L:
1905.
The
made by C. Crossland
28-35,
pl.
marine
SLUITER,
STEPHEN,
Natal
fauna
in the years
of Zanzibar
1902.
- and ROBERTSON,
Zanzibar.
Proc.
Naturhist.
East
Gephyrea.
Die
and flora
of sand
Proc.
C.- 1883.
Die
d. Philippinen
Sipunculiden
und
Intertidal
der Sipun-
water
editor.
Africa
animals.
from
Zool.Soc.
collecLondon.
Island,
eine systematische
14 pls.
Siboga-Expedition
of Cape
Mozam-
Provin*ce
XXV
:
and Natal.
l-53.
Ann.
report on the Echiuridae
and Sipunculidae
I.D.- 1952. A p re l iminary
Royal Soc. Edinburgh.
Sect. B 64.: 426-444,
1 chart,
1 pl.
African
Foren.
at lnhaca
Sipunculiden,
II. 4: 1-131,
Echiuriden.
Sipunculids
flats
and Echiuroidea
E.- 1959.
S i p unculoidea
.
Foren.
1121 : 55-73,
6 figs.
WESENBERG-LUND,
Dsk; Naturhist.
South
and British
1901 and 1902.
I.G., and BULOW,
Reisen
im Archipel.
A.C.-, 1942.
N o t es on the
Mus.
10 : 245-256,
1 pl.
1963.
Kenntnis
1..
DEMAN,
Semper,
C.Ph.-
systematischen
’
and salinity
on marine and brackish
and Marine
Biology,
Vol. $1. H. Barnes,
MACNAE,
W. and KALK,
M.- 1962.
The fauna
bique,
J. Animal
Ecol.
31 (1) : ‘93-128.
SELENKA,
E.,
Monographie.
und
sipunculids
125:
and
echiuroids
from
coastal
from
Mauritius.
waters.
Vid.
.
Vid.
Medd.
of
Medd.
Dsk.
101-146.
61
TABLE
DISTRIBUTION
OF
SPECIES
I
SIPUNCULID
.
.
.
SPECIES
.
.
.
AT
.
.
.
EACH
STATION
.
.
each
SD
Sipunculus
indicus’
8
Sipunculus
robusfus
1
Siphonosoma
cumanense
54
Siphonosoma
australe
17
Phascolosoma
nigrescens
Phascolosoma
scolops
2
Phascolosoma
20
denfigerum
Aspidosiphon
11
corallicola
Cleosiphon
15
aspergillvm
Total
no.
of
4
individuals
at each ,station
5
3
22
7
1.1 16
1
3
14
APPENDIX
Station
2.
Nosy
3.
Shore
Ambariobe;
south
4.
Nosy
Tanikely;
5.
Shore
south
appeared
25 June
1964;
of laboratory;
26 June
1964;
of Idboratory;
like
rock
1964;
coral
poois
coral,
clay.
7.
Southwest
of Nosy Komba;
13O 30’ S 48O 20’
Sand, shell fragments,
and sol,itary
corals.
8.
A Kirintsa,
mangrove
stones,
and roots.
9.
Ampasindava,
with underlying
62
east side
pebbles
of Ambanoro
and stones.
1964;
E;
of Ambanoro
Bay;
11
8
18
172
5
6 July
rocks.
and
of hard
Ambanoro
Boy;
30 J une 1964, 1 & 9 July
bay; muddy sand flats with some rocks.
end
dead
1964;
6.
at north
3
masses.
under
and 21 July
the consistency
20
List
tide
2.3 m tide;
28 June
but with
1964;
12
&l
I
2.3 m tide;
25 June
8
living.
h ar d ened,
3.1~ m, 3.5
2 July
Bay;
1964;
mostly
sand
m, 1.3 m tides;
dredged
5 & 19 July
.1964;
muddy
north
in 20 m;
1964;
rocks,
(indurated)
firm
few
end of
bottom
s.andy
sandy
mud,
patches
-
10.
Ambatoloaka;
11.
Antafianambitry;
muddy
12.
13.
8 July
1964;
16 July
1.9 m tide;
1964;‘0.9
coral,
sandstone.
m tide;
collected
1964;
0.9 m tide;
clean,
white,
Antsakoabe
; 12 July
Ankifi,
east of coffee
factory;
11 & 14 July 1964;
((grasses
)) and some organic
material;
13 a. sand with
clean,
‘barren
Sand.
13b.
14.
Lokobe
15.
Ambatozavavy;
Point;
13, 16 & 17 July
16 July
1964;
16.
Ilot
17.
Crater
Lake;
19.
1964;
Tulear;
3,4 m tide;
‘coraf.
8 & 9 August
1964;
20.
Tulear;
Ambariotsimaramara;
flooded
10August
1964;
20 July
extinct
1.964;
1.4 m, 4.0m,
coarse,
muddy
1964;
volcanic
sandy
coral
coral
crater
collecting
methods
and (2) snorkeling
were
in the coral
fragmenting
these with a hammer
Several
types
of relaxing
best
results.
slow
addition
larger
ones
black,
Other
substances
of ethanol.
were
first
All
iniected
Sand.
coral.
somewhat
open
to sea;
22 July
of pier.
II
Materials
generaf
types
:
(1) digging
breaking
off
large
pieces
the enclosed
organisms.
tiere
tried,
but menthol
tried less successfully
were
were preserved
th e s p ecimens
formalin
to prevent
APPENDIX
Data
m tides;
one side
south
and
of two
areas,
to obtain
agents
witli
well-oxygenated
.heads.
tiith
mud flots
4.7
coarse,
reef.
Methods
The
extensive
Sand.
APPENDIX
mud;
by R. Maddocks;
sandflats.
and sifting
of coraI,‘and
crystals
sand
generally
gave
on Phascolosoma
deterioration
nigrescens
Size
(in mm)
Number of
Hooks
measured
of the
interna1
structures.
Hook
Size
Average
width
at base
(in microns)
s-12
25
12
48
51
t-l
35
12
40
14
74
66
73
s-13
USNM-1
45
5
72
76’
.50
60
9
74
70
s-15
9
74’
79
73
s-9
65
10
71
69
s-157
70
10
67
72
72
s-165
80
10
62
s-5
8.0
23
68
80
s-150
85
16
87
86
s-8
95
10
84
85
100
12
92
96
s-l
the
propylene
phenoxetol,
MgCl,
and
in 70 per cent ethanol,
but the
III
Animal
s-124
and/or
subsequently
63