A MONOGRAPH ON THE
POLYCHAETA OF
SOUTHERN AFRICA
PART 2. SEDENTARIA
Publication No.
656
A MONOGRAPH ON THE
POLYCHAETA OF
SOUTHERN AFRICA
PART 2. SEDENTARIA
BY
J. H. DAY,
Professor of Zoology,
University of Cape To\vn
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
LONDON: 1967
lisued October, 1967
© Trustees of the British Museum (Natural History) 19G7
Printed by Eyre and Spottiswoodc Limited at
Grosvenor Press Portsmouth
CONTENTS
PART 2
Page
Figures
Introduction
Notes on using the monograph . . . . . vii (2)
Map
Systematic Account
Classification xi (2)
Key to the famihes
xii(2)
0.4- 0.6
Spionidae ....
459
18.Z-18.9
Magelonidae ....
494
19.1
CirratuHdae and Heterospionidae
498
20.1-20.4
Trochochaetidae and .*\spitobranchidae
5'9
21.1
Chaetopteridae
522
22.1-22.2
Orbiniidae
533
23.1-23.6
Paraonidae
555
24.1-24.4
OpheUidae
570
25.1-25.2
Cossuridae
581
26.1
.Scahbregmidae .
583
27.1-27.2
Capitellidae
591
28.1-28.3
.'\renicoHdae
606
29.1-29.2
Maldanidae
613
30.1-30.8
Oweniidae and Sternaspidae
648
3I-I
Flabelligeridae .
652
32.1-32.4
Sabellariidae
667
33-I-33-3
Pectinariidae
678
34- 1-34-2
Anipharetidae
686
35-I-35-5
Terebellidae
706
36. 1-36. 10
Sabellidae
751
37-I-37-IO
Serpuhdae
79'
38.1-38.7
Glossary of Technical Terms
821
References
827
Index of Scientific Names in
Part
> 1 an
D 2 .
842
INTRODUCTION
NOTES ON USING THE MONOGRAPH
When identifying material from a particular area it is first necessary to know
what papers contain original records or descriptions of the fauna. These are not
easy to recognise in a long list of references such as that at the end of this mono-
graph. For this reason a complete chronological list is given below.
30. Seidler, H. J., 1923
31. Fauvel, P., 1923a
32. Mcintosh, W. C, 1925
33. Monro, C. C. A., 1930
34. Augener, H., 1931
35. Monro, C. C. A., 1933
36. Day, J. H., 1934
37. Monro, C. C. A., 1936
38. Monro, C. C. A., 1937
39. Treadwell, A. L., 1943
40. Day, J. H., 1951
41. Day, J. H., 1953
42. Tebble, N., 1953
43. Tebble, N., 1953a
44. Day, J. H., 1955
45- Day, J. H., 1957
46. Banse, K., 1957
47. Wilson, D. P., 1958
48. Kirkegaard,J. B., 1959
49. Tebble, N., i960
50. Day, J. H., i960
51. Day, J. H., 1961
52. Uschakov, P. V., 1962
53. Day, J. H., 1962
54. Laubier, L., 1962
55. Day, J. H., 1963
56. Day, J. H., 1963a
57. Bellan, G. and Picard, J., 1965
— Day, J. H., unpublished records
In biogeographical studies one must know what species are found in the area
covered by the monograph, where and at what depth they occur and what synonyms
have been used in earlier works. It is hoped that the species lists appended to each
I.
Peters, W. C, 1854
2.
Stimpson, W., 1856
3-
Kinberg,J. G., 1858-1910
4-
Schmarda, L. K., 1861
5-
Baird, W., 1865b
6.
Quatrcfages, A. de, 1863
7-
Kinberg,J. G., 1867
8.
Grube, E., 1867
9-
Grube, E., 1869
10.
Mcintosh, W. C., 1885
II.
Marenzeller, E. von, 1887
12.
VVilley, A., 1904
13-
Mcintosh, W. C., 1904
14.
Gravier, C., 1905c
15-
Ehlers, E., 1908a
16.
Ehlers, E., 1908
17-
Gravier, C., 1909
18.
Ashworth, J., 1910
19-
Ashworth, J., 191 1
20.
Pixell, H., 1913
21.
Ehlers, E., 191 3
22.
Ramsay, L., 1914
23-
Horst, R., 191 7
24.
Ehlers, E., 191 7
25-
Horst, R., 1918
26.
Augener, H., 1918
27-
Fauvel, P., 1919
28.
Fauvel, P., 1921
29-
Treadwell, A. L., 192 1
viii 12I
poi.vcHAi.rA or southern afric;a
family will sii]iply this informatii)n rapidly and that the "Records" and "Distri-
hutidn"' will siqiply the further details that may be required.
All the published records of polychaetes from southern Africa, Angola and
Madagascar have been extracted from the list of papers given above and other,
more recent unpublished records have been added. The complete list of valid species
is given familv bv family. Each list is arranged alphabetically for ease of reference
with synonyms and incorrect identifications preceded by the word "as". All species
names are aimotated by a code showing which workers used that name and the
province and depth in wliich the records were made. The code is explained as
follow s :
Anlhoritv for the record
.Sliown by a number whit !i
refers to the numbered
list of references given
above.
Provime whtrt' colhctcil
.\ = .Xngola
O = Cape Province
M= Madagascar
N = Natal
P = Portuguese East .Africa
(Mocambicjue)
■\V= Soulh West Africa
Depth range
a = abyssal (over 1000 metres)
d = deep (100-499 metres)
e = cstuarine
i = intertidal
p = planktonic
s — shallow (1-99 metres)
\d -- very deep (500-999 metres)
The use of the code is best shown by an example. Lejiidonotiis semilectus is listed
among the Polynoinae on p. 37 (Part i) as follows:
Lcpidonolus scmitecttis Stimpson 2C.i (and other code numbers)
as Lepidonotus ivahlhergi Kinberg 3C'iNi (and (ithcr code numbers)
as Pohnoe trochiscophora Schmarda 4Ci
The first record shows that the valid name is Lepidonoliis seiniiectus first used by
Stimpson 1856 (code number 2 in the literature list) and his record was made in
the Cape Province (code letter C) in the intertidal zone (code i). The same specific
name has been used by several other workers as shown by the other code numbers
against it. The first synonym is Lepidonotus icahlhergj Kinberg and the code 3CiNi
gives the reference to Kinberg's publication in 1858-1910 and the information that
these specimens were collected in the intertidal zone in the C^ape Province and Natal.
The second synonym is Polynoe trochiscophora Schmarda and the code letters 4Ci
show that Schmarda's name was published in 1861 and the specimens came from
the intertidal zone of the Cape Province. Other synonyms and records follow and
all ol them together show the full range of synonyms which appear in the polychaete
literature of southern Africa and that Lepidonotus semitectus is a common intertidal
and shallow water species which extends from South West Africa around the Cape
of Good Hope to Natal.
More detailed information is appended to the description of each species. If
there are only three or four locality records, all of them arc given, and if there are
many, a summary shows the limits of the geographical and bathymetric range.
After careful consideration, it was decided that place names wiiuld not be as helpful
as latitude and longitude since the names of many collecting stations would not be
found on ordinary maps and, in any case, dredged and plankton records would have
to be given in degrees of latitude and longitude. Minutes of latitude and longitude
ix(2)
•ader
)t be
[uare
:ient.
. east
' can
• the
>Jatal
I the
re is
1 the
Ilape
form
ition
Ided
:dly.
vhile
may
been
Tiists
^r- 130 14" 15- 16" 17° 18° 19
= 20' 21' 22° 23°
24° 25° 26
o 27° 28° 29° 30° 31° 32° 33° 34° 35"
36° 37"
12° 13° 14° 15° 16° 17° 18° I
9° 20° 21° 22° 23° 24° 25° 26° 27° 28° 29° 30° 31° 32° 33° 34° 35° 36" 37°
INTRODUCTION ix (2)
have also been omitted for the sake of brevity and this means that unless the reader
refers to the publication from vvhich the record has been extracted, he will not be
able to pinpoint the record more accurately than somewhere in the 60 mile square
formed by a degree of latitude and longitude. For most purposes this is sufficient.
Luckily the whole of southern .\frica is covered by degrees of south latitude and east
longitude so that the words "south" and "east" are omitted. Thus the locality can
be expressed in four figures and the depth range by a letter. For example the
records for Hermonia hystrix described in Part i are shown as Cape (31/15/d) ;? Natal
(29/3 1 /s). This means that the species has been recorded in Cape waters in the
latitude/longitude square 3rS/i5°E, in the depth range 100-499 metres; there is
also a doubtful record from Natal in the latitude/longitude square 29°S/3i °E, in the
depth range 1-99 metres. Reference to the map opposite will show that the Cape
record is off Lamberts Bay and the doubtful Natal record is close to Durban.
Distribution beyond the limits of southern Africa is given in the conventional form
and a code letter signifying the depth range has been added when this information
is available. It is urged that some indication of depth range should always be added
to summaries of distribution since the fauna at different depths may differ markedly.
For example tropical species are restricted to intertidal and shallow depths while
the very deep and abyssal bottoms may be colonised by cold water species. It may
also be noted that the summary of distribution given in this monograph has been
deliberately selected from twentieth century reports of well-known taxonomists
since earlier works are not always reliable.
THE SYSTEMATICS OF THE POLYCHAETA OF
SOUTHERN AFRICA
CLASSIFICATION
The phylum Annelida to which the Polychaeta and several other groups of worms
belong, is difficult to classify into classes and orders. Early workers included the
Archiannclida, Polychaeta, Myzostomida, Oligochaeta, Hirudinca, Echiura, Sipun-
cula and Priapulida. Recent workers including Dales (1963), regard the last three
groups as distinct and consider each of the other groups as a separate class of the
phylum Annelida.
The division of the class Polychaeta into orders has been attempted by Benham
(1896) and Dales (1963). They have based their divisions on the structure of the
head and the nature of the feeding organs, the regions of the body and the nature
oftheparapodia and setae. As shown in Part i the method of feeding and the habitat
whether it be planktonic, active crawling on the surface, burrowing in the mud or
tubicolous does have an important effect on the structure of the head and body.
However there are still many doubts regarding the homologies of the feeding organs
and there is still no general agreement as to which families should be included in the
various orders which have been erected. For this reason it is better to leave the
matter open and agree with Fauvel (1923) and many earlier workers that for practical
purposes the arbitrary grouping into Polychaeta Errantia and Polychaeta Sedenlaria should
be used.
The Polychaeta Errantia includes active carnivores and a few others while the
Polychaeta Sedentaria includes the remaining microphagous feeders. There are no
mutually exclusive characters which define these two groups and a summation of
characters is used in the following key.
Key to the Families of Polychaeta
Note Some families have such a wide range of characters that they appear twice in the key. In
these cases a number in brackets refers to the other couplet in which the family appears.
Most of the following characters:
Prostomium with sensory appendages. Pharynx armed with jaws or teeth. Parapodia
well deseloped and often bear compound setae . (Polych.veta Err.\ntia) Part i
Most of the following characters: p. 20
Prostomium seldom with sensory appendages and often fused to the peristome which
may bear grooved palps, buccal cirri or a branchial crown. No jaws or teeth.
Parapodia often reduced and compound setae very rare (Polych.\eta Sedentaria) Part 2
p.xii (2)
xii(a) l■()l.^ (;H.\r. r.\ C)l- SOUllir.KN aikica
Pi)i,yciiai:ta Sichentaria
I Hocly .'■iKirl aiul stcmt Willi .i lull nf rilaiiuiUoiis anal yills il'ig. (i. -,.!!) SrtRNAsriDAi (p. 648)
— Bocl>" rU'Us^atc. Xo anal gills .......... 2
■2 Head modilied by the development of frilly membranes (fig. 0.6. lb), buccal tentacles
;tig. o.ti.6) or a branchial crown aroimd the mouth (fig. o.b.y). Proslomiuni often
reduced and indistinguishable from the buccal segment ..... 3
— Head not greatly modified. Prostomium usually well de\"elopcd and c>b\ious. liutcal
segment sometimes with parapodia and may bear a ])air ofadiiesi\e palps {\\g. 0.4.1b)
or a few groo\'ed tentacles ^fig. 0.4.6b I . . . . . . . 17
3 Buccal segment \vith tentacles retractile into tlie mouth (fig. o.().3)
.\mph.\ri;tii:).\e (22) (p. C86)
— Buccal segment with a pair of adhcsi\'c palps (often broken ofi ) or several grooNctl
tentacles dorsally ............ 4
— Buccal segment without food-gathering ajspendagcs of any sort . .... 10
4 Hooded hooks (fig. 0.4.1.V) present in the posterior segments at least. Parapodia always
%vell dc\eloped ............ 5
— Hooded hooks entirely absent. Parapodia sometimes reduced to mere ridges . . b
5 Head flattened and spade-shaped. Gills absent. Palps papillose (fig. 0.4.2)
Magelonid.-^e (p. 494)
— Head not llattened. Gills ofien present. Palps grooved (fig. 0,4.1) . SpioNtoAE (p. 459)
G Long filamentous gills at least on anterior segments. Parapodia reduced to ridges . 7
— Gills not long and filamentous. Parapodia not in the form of ridges .... 8
7 Body divided into an anterior region of short segments and a posterior region of long
segments (fig, 0,4,5) ,.,,... Heterosfionidae (p. 318)
— Body not divided into regions; segments do not differ greatly in length (fig. 0.4.6)
Cirratulid."\e (p, 498)
8 Both rami of anterior parapodia well de\eloped and prox'itled with long setae (fig, 0,4,4)
Trochoch.^etidae (p, 519)
— Either the notopodiimi or the neuropodium of anterior segments reduced or absent . 9
9 ,\nterior segments uniramous having no neuropodia. Posterior segments biramous with
neurosetae in the form of minute uncini (fig, 0,4,3) ■ ■ Gh.'Vetopterid.^e (p, 322)
■ — .Anterior segments with notopodia reduced to cirriform lobes with an internal aciculum
but not setae (fig. 0.4.8) ,,...., Aspitobrancuid.^e (p, 321)
KEY TO THE FAMILIES
xiii (2)
Fig. 0.4. Illustrations of Family Characters, i, Spionidae. 2, Magelonidae. 3, Chaetopteridae.
4, Trochochaetidae. 5, Heterospionidae. 6, Cirratulidae. 7, Orbiniidae. 8, .\spito-
branchidae. (a) Entire worm, (b) Head, (ca) .\nterior foot, (cp) Posterior foot.
(d) Notoseta. (v) Xeuroseta.
x\\ i_-2) POI.VC:llAETA OF SOUTHERN AFRICA
10 Dcntatc-creslfd hooks (fig. o.').7\) present in posterior segments if not earlier . . 15
— No dentate-crested hooks ...........11
11 Capillary setae crenulate (fig. 0.4. 7d) ...... Orbiniid.xf. (p. 533)
— C'apillary setae not crenulate . . . . . . . . . . 12
1-2 \ single long filiform gill arising from the dorsum of setiger 2 or 3 (fig. 0.5.3)
CoSSfRIDAE (p. 581)
— Gills, if present, not single and median ......... 13
13 Capillary setae winged in anterior segments. .\ median antenna may be present (fig. 0.5. i )
P.AR.\o.MD.\E (p. 555)
— Capillary setae not winged. .V median antenna is never present .... 14
14 Prostomiimi a tapered cone. Body fusiform, often grooved \entrally (fig. 0.5.2)
Ophelhd.xe (p. 570)
— Prostomium notched or lobed. Body swollen anteriorly but not grooved ventrally
(fig. 0.5.4) ......... Sc.\LIBREGMID.\E (p. 583)
15 Dentate-crested hooks with hoods (fig. 0.5.5). Body resembling an oligochaete
C.\PITELLID.\E (p. 591)
— Dentate-crested hooks without hoods. Body not resembling an oligochaete ... 16
16 Middle segments greatly elongated but never annulated (fig. 0.5.7). Gills rare
Maldanid.\e (p. 613)
— Middle segments not greatly elongated but always annulated (fig. 0.5.G). Gills always
present ........... .'Xrenicolidae (p. 606)
KEY TO THE FAMILIES
XV (2)
Fig. 0.5. lUmlrations of Family Characters, i, Paraonidae. 2, Opheliidae. 3, Cossuridae.
4, Scalibregmidae. 5, Capitellidae. 6, Arenicolidae. 7, Maldanidac. 8, Sternaspidae.
(a) Entire worm, (b) Head, (ca) ,\nterior foot, (cp) Posterior foot, (d) Notoseta.
(v) Neuroseta.
xvi (-21 POLVc:HAKTA of southern AFRICIA
17 Head usvially witli a frilled food-galherint; membrane. Never any tentat les, palps or
liipimiaie radiolcs ^Hg. U.6.1) ....... 0\veniid.\e (p. 649)
— Head wiihovit a frilled food-gathering membrane but has either tentacles or palps or
hi])innale radiolcs ............ 18
18 Head widi stout setae ............ 19
Head without setae . . . . . . . . . . . .21
19 Capillary setae annulated. Xo marked body regions. Setae on head usually in the form
of a cephalic cage (fig. 0.6.2) ...... Flabelligf.ridae (p. 652)
— Capillan,' setae not annulated. Body regioiis well marked. Setae on head are paleac
which form part of an operculum ......... 20
20 Two to three rows of palcae. C!audal region lung an<.l cylindrical. Tube attached
(hg. 0.6.3) .......... Sabellarudae (p. 667)
— One row of paleae. Caudal region short and llaitcned. Tube free (fig. 0.G.4)
pECTINARnOAE (p. 678)
2 1 Head with soft tentacles for deposit feeding. Gills often present on the hrst few segments.
Setal t^^pes not inverted in the posterior region ....... 22
— Head with a crown of bipinnate radioles (fig. 0.6.7) ^^^ suspeixsion feeding. No gills
behind the head. Setal t\pes inverted in the posterior region .... 23
22 Tentacles retractile into the mouth. They are cither grooved or papillose (fig. 0.6.5)
Ampharetidae {3) (p. 686)
— Tentacles not retractile into the mouth. They are grooved, never papillose (fig. 0.6.6)
Terebellidae (p. 706)
23 Tube sandy or muddy. .\n operculum is never present among the radioles (fig. 0.6.7)
Sabellid.\e (p. 751)
— Tube calcareous. A stalked operculum often present among the radioles (fig. 0.6.8)
Serpl'lidae (p. 791)
KEY TO THE FAMILIES
xvii (2)
Fig. 0.6. Illmlratiotu of Family Characlen. i,Oweniidae. 2, Flabelligeridae. 3, Sabellariidae.
4, Pectinariidae. 5, Ampharetidae. 6, Terebellidae. 7, Sabellidae. 8, Serpulidae.
(a) Entire worm, (b) Head, (ca) Anterior foot, (cp) Posterior foot, (d) Notoseta.
(h) Palea from operculum, (t) Tube. (\) Neuroseta.
SPIONIDAE
459
Fanuly SPIONTOAE Grube, 1850
Body vermiform and rather flattened consisting of numerous segments. Body
regions not marked except by the shape of the parapodia. Prostomium variable in
shape and sometimes with lateral projections but without true antennae. A pair of
long, grooved peristomial palps. Proboscis unarmed. Parapodia biramous. Branchiae
dorsal and often straplike or united to the notopodial lamellae. Neuropodia as lateral
ridges or lamellae. Sensory areas extend back from the prostomium over a variable
number of anterior segments. Pygidium with anal cirri or a glandular ventral
cushion. Xotosetae are winged capillaries anteriorly but hooks or specialised noto-
setae may be present posteriorly. Neurosetae are winged capillaries for the first
few feet but hooded hooks appear in later segments.
Records from southern Africa
Aonides oxycephala (Sars)
Boccardia ligerica Ferroniere .
Boccardia polybranchia (Haswell)
as Polydora polybranchia Haswell
Boccardia pseudonatrix Day
Dispio magna (Day)
as Spio magnus Day .
Laonice cirrata (Sars)
Malacoceros indica (Fauvel)
as Scolelepis indica Fauvel
Nerinides gilchristi Day .
Polydora antennata Claparede
Polydora armala Langerhans
Polydora caeca (Oersted)
Polydora capensis Day
Polydora ciliata (Johnston)
Polydora flam Claparede
Polydora hoplura Claparede
Polydora hoplura var. inhaca Day
Polydora cf. giardi Mesnil
Polydora kempi Southern
Polydora maculata Day .
Polydora norrnalis Day .
Prionospio bocki Soderstrom
Prionospio cirri/era Wiren
Prionospio cirrobranchiala Day
Prionospio ehlersi Fau\'el
Prionospio malmgreni Claparede
as ? Prionospio capensis Mcintosh
as Prionospio malmgreni var.
44Ci, 45Pi, 48CS, 51CS, — Ms
? 44Ce
4oNi, 44Ci
26WS
51CS
44Ci
48Cd, 5iCsd
4oNi, 45 PiNi
5iCsd '
44Ci, 51CS
32 ? Ci
45Ni
44Ci, 51CS
ayMi, 45?!
51CS
44Ci, 51CS
45Pi
44Ci, 51CS
45Pi
53CS
42NiPi
—Ms
51CS
5iCsd
48AS, — Xsd, Pd
15CS, 2iCi, 34Cd, 56Csd
loCs
15CS
460 I'()1.VC:HA1-. 1 A Ol' StlL'THlRN AIRK:A
Piiiirioipio piiiiiiilii VA\\vvs . . . a I AsCs, 31 As, 48\VsdCd,
f,iCsd, — Ns
Piinnospio sp. . . . . . f,6C'.d
Prionoipio saldaiiha Hay . . . 5iCsd
as Prionospio s]5. .... 44C'i
Priotwspio sexociilali! Augcncv . . . 26\\'s, 44Ca, 45X1, fjiCli
Prionospio sleenslrupi Malmgii-n . . 55C''i, 56C'sd, — Nsd
as Prionospio mnlmnreni \ar. iliihin Day . 5iC!sd
as Prionospio rhicrsi (nun Fainrl) . 48Csd
Pygospio elegans C'laparcde . . . S'Ca
Rhvnchospio glutaca (Ehlcrs) . . . 5 'tis
Scoleh'pis kjehvrei (Gravicr) . . . 53Mi
Scolelepis squamata (Muller)
as Nerine cirratidus (Delle Chiajr) . 44C'i, 45Pi-'^'ii r)3^Ii
as J\'erine cirratuliis var. capensis Mcintosh 3'-C'i
.S'/)/() filicornis (Mullor) .... 44C'i
Spiophanes homhyx (Cllaparcde) . . 48C;s, 5iCsd
Spiopiiants soederslromi WM-imM\ . . 5iCsd, 55Ca, — Xsd
ri()ix)(,ic:al xori'S
The spionids arc tyjiical deposit feeders. All oftheni have a jjair orions; peristomial
tentacles or "pal]5s" which they stretch out o\er the substratum to gather food
particles. These particles travel along a ciliated groove to the lips where they are
sorted and the smaller edible particles ingested ; however the sorting mechanism
does not seem to be \-ery efhcient for the gut contains many sand grains as well as
detritus.
All spionids live in burrows which they lint' with a fragile mucoid secretion.
Scolelepis (olim jXerinc) burrows in inlertidal sandbanks where there is sullicient water
movement to keip the detritus moving. Mahiciurros (olim Scolelepis) burrows in the
mud-filled crexices between stones ; ,sy(;i; often forms dense colonies on sandbanks
and when the current washes aw.iy the loose sand Iietween the tubes, they stand nut
like a host of tiny chimne\s. .Many other genera are common in dredgings on sandy
mud. The various species of Prionospio must be very abundant for tln'ir larvae are
present in enormous mmibers in neritic plankton samples. The lar\-ae oi Poljdora are
very common too but the worms of tliis genus burrow in calcareous materials using
die stout hooks on the fifth setiger for the purpose. Some species oi' Polrdora live in solt
hmestone, others in dead coral and Cjuite a number bore into living mollusc shells.
Most of the shells of abalonc [Haliolis midae) on C^ape coasts are riddled with Polvdora
and /'. ciliata is niiidrious fur foiming ""mud hlist( rs" in oyster shells. The burrow
is roughly pear-shaped and lined with bku k mud which is easily seen through the
nacreous lining of the shell. .Su( h nuul hlisteis wc.iken and may e\en lead to the
death of the oyster though Pohdnrd does not Iced on oyster flesh. Like all hiIk r
s]iionids it is a detritus feeder and its ]ialps may be seen projecting from the little mud
chimney which is iiinlinudus with llie nuid (if the blister.
Most spionids lay large eggs emlused in tough egg-capsules. 1 hese ma) be
SPIONIDAE 461
liberated directly into the sea water so that all development takes place in the
plankton, or, they may be protected inside the burrow during the early develop-
mental stages. Polydora does not release the larvae until they have reached the three-
setiger stage. Pygospio has a more complex life history. The individual ova are small
and contain little yolk. Groups of ova are enclosed in brood capsules and strings of
brood capsules are then attached to the lining of the mother's tube. Only a few of the
ova are fertilised and these grow rapidly and at an early stage develop an enormous
embryonic mouth. By this means the other ova are engulfed. The surviving embryos
then grow into ten-setiger larvae before they are liberated for a brief planktonic
existence.
THE MAIN DIAGNOSTIC CHARACTERS
Important studies on the Spionidae have been made by Mesnil (i8g6), Soder-
strom (1920) and Fauvel (1927).
The head. The prostomium may be pointed, rounded, notched or develop lateral
peaks anteriorly and is often produced back as a ridge with or without an occipital
tentacle. Eyes are of doubtful value as they may coalexce or be hidden beneath the
skin. The peristome supports and may enfold the sides of the prostomium or even
be produced upwards as lateral "wings" in some species. The peristome bears a
pair of grooved, food-gathering "palps" which are not homologous with the palps
of the Errantiate families ; they are often lost when the animal dies.
Body regions. These are not marked but there are changes in the setae, in the
shape of the parapodia and the development of gills. The first setiger is often small
or fused to the peristome and if overlooked will cause confusion.
Branchiae. The segmental arrangement and shape of the gills is of the first im-
portance, also the degree of fusion between the gill and the postsetal lobe of the
notopodium. In Prionospio the gills may develop lateral papillae (pennate gills), but
unfortunately the gills fall off easily.
Parapodia. The postsetal lamella of the notopodium is often well developed
anteriorly but low posteriorly. It may be joined to its fellow by a ridge across its
back in some segments. The neuropodium usually forms a low rounded postsetal
lamella and is of minor importance.
The setae. These include capillaries, hooded hooks and specialised forms whose
shape and segmental position is important. The capillaries have a flange or wing in
anterior segments but this is reduced or absent posteriorly. The hooded hooks
appear first in the neuropodia but may also be present in posterior notopodia. The
segmental arrangement and number of teeth above the main fang or rostrum is
surprisingly constant. Specialised setae provide very useful characters but it should
be noted that those in the posterior notopodia of Polydora only appear near the end
of the body and if this is absent the worm is unidentifiable.
The pygidium. This is glandular in the larva and may remain in the adult as a
ventral cushion or become flattened and saucer-like or be produced into two to
four anal cirri.
462 pc)I,vc;haeia ok solihi.rn aikica
Key to Genera
1 Fourth or fiflh segment with enlarged ai^d modified liooks (tig. 18.1. a) ... 2
- Fourth and fifth segment with normal setae only ....... 4
2 Fourth segment with enlarged hooks POLYDORELLA*
- Fifth segment with enlarged hooks ......... ;j
3 Branchiae start on setigcr 2 (lig. I B.I. k) ..... BOCCARDIA (p. ^62)
- Branchiae start after set iger 5 ....... POiFDOR/1 (p. 465)
4 Branchiae absent. (.\ sjjecialized neuroseta in the first foot (fig. 18. 5. d) )
SPIOPHANES (p. 474)
- .V single pair of large branchiae on setigcr i . . STREBLOSPIO*
- \ few branchiae starting well back or a pair on setigcr i and then a gap l)erorc the rest
PYGOSPIO (p. 475)
- Branchiae start on setiger 1 or 2 and continue over a few or many consecutive segments 3
5 Branchiae continue almost to the posterior end and may be partly or completely fused
to the notopodial lamellae .......... 6
- Branchiae absent ficuii ].iosterior segments. Branchiae never fused to the notopodial
lamellae .............II
6 Prostomium with laterally projecting ]:)caks ........ 7
- Prostomium without laterally projecting peaks ....... 8
7 Branchiae from setiger i MALACOCEROS (p. 477)
- Branchiae from setiger 2 RHYNCHOSPIO (p. 478)
8 Branchiae from setiger 1. No notopodial hooks. Either a ventral cushirm or anal cirri
below the anua ............ 9
- Branchiae from setiger 2. Notopodial hooks may be ])resent in ]>osteri(>r feet. /\ ventral
cushion below the anus (fig. iy.7.g) ......... to
9 Branchiae mainly fused to dorsal lamellae anteriorly. Small accessory branchiae
between the notopodia of middle segments. A ventral cushion below the anus
D/SP/O (p. 481)
- Brancliiae mainly free from dc)rsal lamellae anteriorly. No accessory branchiae. .\nal
cirri present ........... SPIO (|i. .\'V>)
10 Notopodial hooks present in posterior segments. Neuropotlial lamellae btlobed
posteriorly SCOLELEPSIS {|). 482)
— No notopodial hooks. Neuropodial lamellae not notched . . NERINIDES ip. 484)
1 1 Less than 12 pairs of branchiae which may be i>ennate or smooth and st.ut on setiger i
or 2 PRIONOSPIO (p. 485)
— More than 12 pairs of branchiae which are never pennate and never start on setigcr i . 12
12 Prostomium pointed. Notopodial hooks in posterior segments . . i40WD£S (p. 478)
— Prostomium rounded. Notopodial hooks absent .... LAONICE (p. 480)
BOCCARDIA C;arnzzi, 1895
Prostomium blunt to bilobcd anteriorly and with a tapering keel posteriorly.
Eyes usually present. Branchiae start on setiger 2, arc absent from setiger 5 and
then continue. Fifth segment enlarged and provided with stout burrowing hooks.
Anterior segments with winged capillaries in both rami. Posterior notopodia with
capillaries only. Middle and posterior ncuropodia with hooded bidentate hook.s.
Pygidium glandular and often saucer-shaped.
Typp: SPECIES : Po[ydora jm/jbiiuickia Has-wcW, i>']85.
Key to Species
1 .Special setae of fifth setiger are all simple hooks (fig. 18.1.J) .... B. ligerica
— .Special .setae of fifth setiger in two rows, only tlie second row being simple liooks . . 2
SPIOXIDAE 463
First row of special setae have swollen ends surmounted with a fibrillar brush (fig. iS.i.d)
No notosetae on setiger i ....... . B. polybranchia
First row of special setae have swollen ends with a central cone surrounded by a raised
rim (fig. 18. 1. m). No notosetae present on setiger i . . . B. pseudonatrix
Boccardia cf. ligerica Ferroniere, 1898
(fig. 18...J)
? Boccardia ligerica Ferroniere, 1898 : log, pi. 6 figs, a-i ; Fau\el, 1937 : 57, fig. 19 n-s.
Boccardia cf. ligerica; Day, 1955: 415.
Length about 25 mm. Prostomium notched anteriorly and with the keel extending
back to setiger 2. Four eyes. Notosetae present on setiger i. Branchiae from setiger
2, absent from 5 and continuing to setiger 30. Enlarged setae of setiger 5 (fig. iB.i.j)
are plain blunt hooks. Bidentate hooded hooks appear in the neuropodium of setiger
7. No specialised notosetae in posterior segments.
Type locality : Estuary of Loire, France.
Records: ? Cape (33/18/e).
Distribution: France (e).
Boccardia polybranchia (Haswell, 1885)
(fig. i8.i.a-h)
Polydora polybraru:hiaila.swe\], 1885: 275.
Polydora (Boccardia) polybranchia : Fauvel, 1927 ; 58, fig. 20 a-i.
A large species reaching 25 mm. for 80 segments (fig. i8.i.a). Prostomium
(fig. i8.i.b) outlined in black, notched anteriorly and with a keel extending back
to setiger 2. Nought to ten eye-spots. No notosetae on setiger i. Branchiae (fig.
i8.i.c) united to the dorsal lamellae and present on setigers 2-4, absent from setiger
5 and then continue to the middle of the body. Enlarged hooks of setiger 5 of two
types; front row (fig. i8.i.d) with expanded and truncate ends surmounted by a
dense brush of fibrils and second row of simple hooks (fig. i8.i.e). Bidentate hooded
hooks (fig. i8.i.f) appear in the neuropodium of setiger 7. No specialised notosetae
in posterior segments. Pygidium (fig. iB.i.h) small, concave posteriorly.
Type locality : S.E. Australia.
Records: South West .\frica (22 14 i and 26 15/s), Cape (from 33/181 and
34/2 1 /e, i to 28/32/e).
Distribution: English Channel (i, s), Bay of Biscay, Brazil, Subantarctic
(Magellan area), Kerguelen (i), Macquarie Is. (i). New Zealand (i, s), Medi-
terranean, Japan, .Australia.
464
I'CM.VCHAETA OK SOUIIIERN AFRKIA
Fig. iB.i. Bocainiui jiolvbrmuhia. (a) Entire worm (four times natural size), (b) Head.
(C and d) .'\nterior and posterior types of enlarged setae of setiger 5. (e) Tenth foot.
(fJ Hooded hook, (g) Posterior notoseta. (H) Pygidiiim. Boccardia ef. Ugericn. (j) En-
larged hook of setiger 5. Boccardia jneudonalrix. (k) .Anterior end. (l and m) Enlarged setae
of setiger 5. (n) Hooded hook, (o) Pygidium.
SPIONIDAE - 465
Boccardia pseudonatrix Day, 196 1
(fig. i8.i.k-o)
Boccardia pseudonatrix Day, ig6i : 493, fig. 5 e-j.
Body about 10 mm. long with 70 segments. Prostomium (fig. iS.i.k) bilobed with
a posterior keel reaching setiger 2. Two eye-spots. First setiger with a well developed
notopodium and several notosctae. Branchiae on setigers 2, 3, 4, 6 and all subsequent
segments to setiger 40. Fifth setiger without notosetae and with two types of large
hooks. The three hooks of the first row (fig. iS.i.m) each end in a blunt cone
partially surrounded by a raised margin. The four hooks of the second row (fig.
1 8.1.1) are each simple and curved. Bidentate hooded hooks (fig. iS.i.n) appear in
the neuropodium of setiger 7. No special posterior notosetae. Pygidium (fig. 1 8.1.0)
very small.
Type locality : Knysna Estuary, South Africa.
Records: Cape (34/23/e).
Distribution : No other record.
POLYDORA Bosc, 1802
Small tubicolous worms burrowing in shell or limestone. Prostomium blunt or
bilobed anteriorly but extending back as a dorsal keel. Eyes present or absent.
Anterior parapodia with winged capillaries in both rami. Setiger 5 enlarged and
bears stout hooks for burrowing. Posterior notosetae may include speciaUsed setae
as well as capillaries. Bidentate and usually hooded hooks present in the neuropodia
from setiger 7-10 onwards. Branchiae start behind setiger 5 and are not united to
the notopodial lamellae. Pygidium swollen, glandular and often saucer-shaped.
Type species : Poljdora cormila Bosc, 1802.
Key to Species
1 Posterior neuropodial hooks without hoods and unidcntate (fig. 18. 2. f, g, h). (Posterior
notosetae include two to three straight needles) . . . .P. capensis (p. 466)
- Posterior neuropodial hooks hooded and bidentate ......
2 Posterior notosetae include specialized types ........ 3
- Posterior notosetae are all capillaries ......... 6
3 Enlarged hooks of setiger 5 bidentate. Posterior notosetae include one to two stout hook:
(fig 18.2.I, m) . '
- Enlarged hooks of setiger 5 hooded. Posterior notosetae include a cone of stout needles
(fig. i8.2.i, j) .......... P. armata (p. 466)
- Enlarged hooks of setiger 5 unidentate. Posterior notosetae include a bundle of fine
needles or three to four spines .......... 5
4 Enlarged fifth hooks with a minor spur twisted around the main terminal tooth (fig. 18. 2.1)
P. hoplura hoplura (p. 468)
- Enlarged fifth hooks end in a shelf from which two unequal teeth arise side by side
(fig. 18.2. n) ......... P. hoplura inhaca (p. 468)
5 Posterior notosetae include a bundle of minute needles (fig. i8.3.d) . . P.flava (p. 468)
- Posterior notosetae include three to four straight stout spines (fig. 18. 3. g) . P. caeca (p. 469)
466 POLVCHAETA OF SOUTHERN AFRICA
6 Hooded hooks start in the neuropodium of setiger 7. Enlarged hooks of setiger 5 cithrr
unidentate or bidentate. Branchiae may start on setiger 7, 8 or 9 . . . . 7
- Hooded hooks start after setiger 7. Enlarged hooks of setiger 5 always unidentate.
Branchiae always start on setiger 7 ........ . 8
7 Enlarged hooks of setiger 5 with a spur (tig. 18. 3. j). Branchiae from setiger 7 to posterior
end ........... P. ciliata (p. 469)
- Enlarged hooks of setiger 5 with a basal sheath simtilaling a spur (tig. i8.4.bj. Branchiae
from setiger 9 to setiger -^j ...... P. cf giardi (p. 471)
- Enlarged hooks of setiger 5 without a spur (fig. 18. 3. 1). Branchiae from setiger 8 to the
posterior end .......... P. normalis (p. ^-ji)
8 Hooded hooks start on setiger 8. Branchiae do not reach the posterior end. Prostomiiun
with one occipital tentacle ........... 9
- Hooded hooLs start on setiger 9. Branchiae reach the posterior end. Prostomium with
two occipital tentacles dig. 18.4.C) ...... P. maculata (p. 472)
9 Enlarged fifth hooks like pointed spoons accompanied by n')rmal winged cajjillaries
(fig. 18.4. f g) .......... P. antennata(p. 473)
- Enlarged fifth hooks are plain and blunt accompanied by setae like curved fangs with
a basal swelling (fig. 18.4.J, k} ...... . P. kempt (p. 473)
Polydora capensis Day, 1955
(fig. i8.2.a-h)
Polydora capensis Day, 1955: 416, fig. i k-s.
Body (fig. 18. 2. b) up to 20 mm. long with 100 segments. Picistomium (fig. 18. 2. b)
not notchetd anteriorly, posterior keel extending to setiger 3. Buccal segment forms
supporting lobes on cither side of prostomium. Notosctae present on setiger i.
Enlarged fifth setae (fig. i8.2.e) are plain blunt hooks. Branchiae (fig. i8.2.c) from
setiger 7 to 37. Xcuropodial hooks from setiger 7 ; thev number three to five and
are both hooded and bidentate anteriorly (fig. i8.2.f). Posteriorly they lose their
hoods (fig. 18. 2. g), become unidentate (fig. 18. 2. h) and are reduced to two to three.
Posterior notosetae include about three straight spines as well as capillaries. Pygid-
ium (fig. i8.2.d) small. Found boring in living gastropod shells.
Type loc.\lity: Simonstown, South Africa.
Records: Clape (from 32 171 and 34'i8;'i, s to 32/28/i).
DiSTRiBUTio.N' : .South African endemic.
Polydora arniata Langerhans, 1880
(fig. i8.2.i-j)
Pulydom armala Langerhans, 1880 : 93, pi. 4 fig. 5 ; Fauvel. 1927 : 53, fig. 19 a-e.
Body small, about 5 mm. long with 24-45 segments. Prostoinium bilobcd
anteriorly and produced back as a dorsal ridge reaching setiger 2. No eyes. Two to
three notosetae in the first foot. Branchiae from setiger 7 to 14. Enlarged setae of
fifth foot (fig. i8.2.i) number two to three, each bluntly hooked with a stout hood.
Three to four bidentate and hooded hooks from the neuropodium of setiger 7
SPIOMDAE
467
g h
Fig. 18.2. Polydora capemis. (a) Anterior end. (b) Entire worm (six times natural size).
(c) Ninth foot, (d) Pydigium. (e) Enlarged fifth hook, (f) Anterior bidentate hooded
hook, (g) Posterior bidentate hooded hook, (h) Far posterior hook from neuropodium.
Polydora armata (after Fauvel, 1927). (i) Enlarged fifth hook, (j) Cone of spines from a
posterior notopodium. Polydora hoplura. (k) Pygidium. (l) Enlarged fifth hook, (m)
Posterior notopodial hook. Polydora ho/jliira inhaca. (n) Enlarged fifth hook.
468 POl.VCHAETA OK SOUFHERX Al'RU'.A
onwards. Posterior notosetac include eight to t\\el\c spines which form a cone
when retracted (fig. 18.2.J) and a funnel when exerted.
Type locality: Madeira.
Records; One doubtfi^d record from an unknown locality.
DisTRiBtTlox: North Atlantic from the English Cihannel (i, s) to Morocco (i)
and Madeira Is.: Gulf of Mexico (d) ; Mediterranean; C^cylon ; North Pacific
(Japan).
Polydora hoplura hoplura CHaparedc, 1870
(fig. i8.2.k-m)
Pohdora hojilura C:laparede, 187U : -,8. pi. 22 fig. 2 ; Fauvel. 1927 : 50. tig. 17 a-g.
Large worms reaching 50 mm. in length \shich burrow in soft limestone. Pro-
stomium fairly deeply notched, often blackened but eyes few or absent. No noto-
sctae on first foot. Branchiae from setiger 7 to end of body. Enlarged hooks of
fifth foot (fig. 1 8.2.1) each With a lateral tw-istcd spur. About eight bidentatc and
hooded hooks from setiger 7 onwards. On the last few segments before the pygidium
the notosetae include one to two enlarged, yellow, reciHAed hooks (fig. i8.2.m).
Pygidium (fig. i8.2.k) saucer-like and often blackened.
Type loc;ai_itv : Naples.
Records : Clape (from 33 18 i. s to 34 23 i>.
DiSTRiBUTiox : North .Atlantic from Ireland to the English Clhannel (e, i, s) ;
Mediterranean.
Polydora hoplura inhaca Day, 1057*
(fig. i8.2.n)
Pohdora holplwti \'ar. inhaca Day, 1957 : 99, fig. 6 k, 1.
Generally similar to typical P. h. hoplura but prostomium rounded, eyes absent,
gills from setiger 7 to middle of body and enlarged hooks of setiger 5 distincti\e.
Each ends in a shelf from which a large and a small tooth arise side by side (fig.
iS.2.n).
Type locality: Inhaca Is., Dclagoa Bay.
RpfiORD : Mocambique (26 32 i).
DiSTRiBUTio.x - no other records.
Polydora flava CUaparedc, 1870
(fig. i8.3.a-d)
Pulyilnra flaia Ckiparede, 1870: 487; Faiivcl, 1927: 52. tig. 17 n-u.
Length up to 45 mm. for 150 segments. Prostomium (fig. 18. 3. a) notched
anteriorly and produced back as a ridge reaching setiger 2. No e\'es. A few noto-
setae on the first foot. Branchiae start on setiger 7-9 and extend back past the
*Pri>tiablv ( I ins[)(Tilic wiili P'llydara colotva Mmotc ki'i; fn'oi .Ma,saclnisfUs. L .S.A.
SPIOMDAE 469
middle of the body. Enlarged hooks of fifth foot (fig. 18. 3.0) have curved bkint tips
flattened on the concave side and are accompanied by fine straight spear-bladed
capillaries (c'). Three to four bidentate and hooded hooks from the neuropodium
of setiger 7 onwards. Posterior notopodia have packets of very fine needles embedded
near the base of the normal capillaries.
Type locality : Gulf of Naples.
Records: Cape (33/18/s and 34/18/s).
Distribution: English Channel (s) ; Mediterranean (i, s) ; Ceylon: Sumatra;
Japan and far northern Pacific.
Polydora caeca (Oersted, 1843)
(fig. i8.3.e-h)
Leucodorum caecum Oersted, 1843 : 39.
Polydora caeca: Fauvel, 1927 : 52, fig. 18 a-k.
Length reaching 40 mm. for 130 segments. Prostomium (fig. i8.3.e) deeply
notched anteriorly and produced back as a dorsal ridge reaching setiger 2. Eyes
four or absent. A few notosctae on the first foot. Branchiae start on setiger 7-9
and extend past the middle of the body. The special setae of setiger 5 include a
row of fine hastate setae and an oblique row of stout hooks excavated on the concave
side (fig. 18.3. f). Bidentate and hooded neuropodial hooks from setiger 7. Posterior
notopodia (fig. 1 8.3.g) with three to four stout straight spines as well as the capillaries.
Pygidium (fig. i8.3.h) saucer-shaped with a dorsal notch.
Type locality ; Oresund, Sweden.
Records: Natal (29/31 /s) ; Mocambique (26/32/i).
Distribution : Arctis (s) ; Nordi Atlantic from Greenland (s) to North Carolina (s)
and English Channel (i, s) to Morocco (i) ; Mediterranean ; tropical Indian Ocean.
Polydora ciliata (Johnston, 1838)
(fig. i8.3.i-j)
Leucodore cilialus ]cihnston, 1838: 67.
Polydora ciliata: Fauvel, 1927: 49, fig. 16 i-p.
Length up to 30 mm. for 100 segments. Prostomium (fig. i8.3.i) shghtly notched
anteriorly and produced back as a ridge reaching setiger 2. Four eyes in juveniles
but these may be lost later. No notosetac on the first foot. Branchiae from setiger
7 to near the end of the body. Enlarged hooks of fifth segment (fig. 18.3.J) are stout
hooks with an accessory tooth. Hooded and bidentate hooks start in the neuro-
podium of setiger 7. No specialised notosetae in posterior feet. Pygidium saucer-
470
I'OLVCHAETA OF SOUTHERN' AFRICA
C C
1
Fig. 18.3. Polydora flava. (a) Head,
and accompanying winged capillary
(•e) Head. <t) Enlarged fifth hook.
(ij Head, (j) Enlarged fifth hook.
IB) BranchiferoLis foot (ninth I. (c) Enlarged hook
front setiger 5. (Oj Far posterior foot. Polydora ciwca.
(<;) Far posterior foot. (n) Pygidinni. Polvdoia (ilinlii.
Polydora normalis. (k) Head, (l) Enlarged fifth hook.
SPIOMDAE 471
like and incised dorsally. Both the anterior and the posterior end often blackened.
Forms burrows in calcareous rock and "lithothamnion".
Type locality: Berwick, Scotland.
Records : Mocambique (23/35/e, s)
Distribution : Baltic Sea ; North Atlantic from Scotland (i) and the English
Channel (i, s) to Senegal (s) ; Falkland Is. (i) ; Mediterranean ; Red Sea (s) ;
India (i) ; Madagascar (i) ; N. W.Japan.
Polydora normalis Day, 1957
(fig. i8.3.k-l)
Polydora normalis Day, 1957 : 97, fig. 6 f-j.
Body rather small, seldom more than 10 mm. for 72 segments. Prostomium
(fig. i8.3.k) deeply notched anteriorly and continued back as a ridge to setiger 4.
No eyespots. One to two notosetae on the first foot. Branchiae start on setiger 8
and continue to near the end of the body. Enlarged setae of fifth foot as plain
hghtly curved hooks (fig. 18.3.I). Bidentate and hooded hooks appear in the
neuropodium of setiger 7. No obviously modified notosetae in posterior segments
but the shorter capillaries have blades that are more striated than usual.
Type locality : Inhaca Island, Delagoa Bay.
Records: Natal (30/30/s and 29/32/1); Mocambique (26/32/i and 25/35/e).
Distribution : S.^\'. Indian Ocean.
Polydora cf. giardi Mesnil, 1896
(fig. i8.4.a-b)
? Polydora giardi Mesnil, 1896 : 195, pi. 13 figs. 1-12 ; Fauvel, 1927 : 50, fig. 17 h-m.
Polydora cf. giardi: Day, 1961 : 493, fig. 5 k.
Body thread-like, about 12 mm. long. Prostomium (fig. 18. 4. a) deeply notched
anteriorly and continued back as a ridge to setiger 3. No eyes. One to two notosetae
on the first setiger. Branchiae start on setiger 9-10 and continue to the 25th or 30th.
Enlarged setae of fifth foot as three to four stout hooks (fig. i8.4.b) each with a
subtcrminal sheath which simulates an accessory tooth. Bidentate hooded hooks
appear in the neuropodium of setiger 7. No specialised notosetae in posterior
segments. Pygidium small, saucer-shaped.
Type locality : Saldanha Bay, South Africa.
Records: Cape (33,i8;'s to 32/28/i).
Distribution : (of P. giardi) Ireland ; English Clhannel (i) ; ? Mediterranean ;
Pacific from Alaska to western Mexico.
47--:
^cn.^■(:nAI-. r A oi' sornir.RX afric:\
■St. I'
Fi<:. iU.4. Pvlyilora cf. ;■/««//. (a) Head, (ni Enlars,'cd fiiih hooks. Pulydnui maculnln. (cl
Head, (dj Enlarged fifth hooks. Polydora anicnnala. lEiHcad. Ir, Lateral view of hciiyer 5
showiiiCT position of enlarged Itooks below notopodial capillaries, (cj Enlarged fifth hooks.
Pulnlara kanpi. (ill Head, (i) Lateral view of setiger -, to show distribution of setae.
(j) Fang-like seta from setigcr 5. (K) Enlarged hook I'ruin setiger -,.
Polydora maculata Day, 1063
(fig. i8.4.c-d)
Polydora macidata Day, 1963 : 417, fig. i) a -d.
Tubes occur in old gastmpocl slulls inhabiucl by hciniit ciabs. Vhk\\ up li> 20 mm.
long by I -o mm. for 130 segments. Palps bailed with blai k but no otlicr colour
markings. Prostomium (fig. 18. 4. c) bilobed anteriorly and continued back as a
ridge tC) setiger 2. Two small tentacles one behind the other on the prostomial ridge
between the bases of the palps. Branchiae from scliger 7 to the end of the body.
Xotosetae absent from the first foot. No specialised notosetae in posterior feet.
Bidrntale hooded hooks in tlie iieuropodia Imm setiu'i-r f), aulerioily v\'^\\y to ten
SPIOMDAE 473
in number, but only four posteriorly. Enlarged hooks of setiger 5 (fig. i8.4.d) have
plain, curved and slightly flattened ends. They alternate with a row of normal
winged capillaries but there is no separate tuft of setae on segment 5.
Type locality : Off East London, South Africa.
Records: Cape (32/28/s).
Distribution : No other record.
Polydora antennata Claparede, 1870
(fig. i8.4.e-g)
Polydora antennata Claparede, 1870: 320, pi. 21 fig. 3; Fauvel, 1927: 56, fig. 19 i-m ; Day, 1955:
4'5-
Length up to 30 mm. for 100 segments. Prostomium (fig. i8.4.e) deeply notched
anteriorly and produced back as a dorsal ridge with one free occipital tentacle.
Two to four eyes. No notosetae on the first foot. Branchiae start on setiger 7, are
poorly developed and end about setiger 40. The enlarged hooks of setiger 5 (fig.
i8.4.g) have ends like spoons and are arranged in a horse-shoe (fig. i8.4.f). Bidentate
hooded hooks start in the neuropodium of setiger 8. No specialised setae in posterior
notopodia. Pygidium saucer-shaped with dorsal and ventral notches.
Type locality : Italy.
Records: Cape (33/18/5 and 34/18/i).
Distribution : Mediterranean (i) ; Arabian Sea (d) ; Ceylon ; Gulf of Manaar ;
Japan.
Polydora kempt Southern, 192 1
(fig. i8.4.h-k)
Polydora kempi Southern, 1921 : 636, pi. 28 fig. 20 a-j ; Day, 1957 : 99.
Length about 12 mm. Prostomium (fig. i8.4.h) faintly notched anteriorly and
bears one occipital tentacle posteriorly. Four eyes. No notosetae on the first foot.
Branchiae from setiger 7 to 14. Setiger 5 (fig. 18.4.1) with normal notopodial
capillaries, a double row of specialised setae and normal neuropodial setae. The
specialised setae include an anterior row of recurved fang-hke forms with a swelling
at the base of the blade (fig. 18.4.J) and a posterior row of stout, simple, faintly
curved hooks (fig. i8.4.k). Bidentate hooded hooks start in the neuropodium of
setiger 8. No specialised notosetae in posterior segments. Pygidium small and
saucer-like.
Type locality : Chilka Lake, India.
Records: Mocambique (23/35/e).
Distribution: India (e).
474 POI.VCillAr, TA Ol' SOL'l IIKR N AIRICA
SPIOPHANES Gnibc, i860
Prostoniiuni with or without hitcral peaks, tapered posteriorly and bearing an
occipital tentacle. Eyes present or absent. No branchiae. Dorsal lamellae markedly
tapered. Notosetac arc winged capillaries throughout there being no notopodial
hooks. Neurosetae of first foot include numerous winged capillaries plus one to two
stout curved setae ; the next few neuropodia have winged capillaries only while
those further back bear hooded hooks plus an inferior sabre-like seta. Pygidium
with anal cirri.
Type species: Spiopliana kroyeri Grube, i860.
Key to Species
Prostoniiuni with jiuinteti lateral peaks. Hooks bidcntate . . . . . S. bonibyx
Prostomiuin broadly oval anteriorly. Hooks tridentate ...... 2
Genital setae smooth .......... .S. soederstromi
Genital setae furry . . . . . . . . . . . S. kroyeri*
Spiophanes bombyx (Claparcde, 1870)
(fig. i8.5.a-c)
Spio bombyx Claparede, 1870: 485, pi. 12 fig. 2.
Spiophanes bombyx \ Faiivel, 1927: 41, tig. 143-1.
Body up to 60 mm. in length with 180 segments. Prostomium (fig. 18. 5. a) broad
and straight in front witli large, pointed lateral peaks anteriorly and an occipital
tentacle posteriorly. Four eyes. A pair of interrupted dorsal sensory grooves from
prostomium back to setiger 4. Fibrous glands in the fifth to fifteenth foot. Noto-
podial lamella of first foot slender, succeeding ones (fig. 18. 5. b) broadened basally.
Low dorsal ridges join the notopodia from setiger 3 onwards. Neuropodial lamellae
of setigcrs one to lour trianguhu', succeeding ones (fig. 18.5.C) low and rounded.
Xotosetae of the first toot very long. Enlarged setae in neuropodium ol first foot
have recur\ed blades (fig. i8.5.d). From the fifteenth foot onwards the neurosetae
include bidcntate hooks (fig. i8.5.e) with a parti. il guard below tlie rostrum plus
an inferior sabre-like seta. Anal cirri cylindrical.
Type locality : Naples.
Records: C^ape (34/18/i, s and 34/j2/d) ; Natal (3o/30;'s).
Distribution-: Atlantic from Sweden (d), Scotland (s) and English C^hannel (i)
to North Clarolina (i) ; Mediterranean; Gulf of Mexico ; Gape Verde Is. (s) ;
Liberia (s) ; Angola (s) ; Falkland Is. ; N. Pacific from Japan and \V. Canada to
S. C'alifornia.
SPIONIDAE 475
Spiophanes soederstromi Hartman, 1953
(fig. i8.5.f-j)
Spiophanes idderslromiHartman, '953: 4I) fig. 14 a-c ; Day, 1961 : 484.
A slender species up to 25 mm. long with about 60 segments. Prostomium (fig.
i8.5.f) transversely oval anteriorly, supported by the buccal segment laterally and
narrowed with an occipital tentacle posteriorly. Two pairs of subdcrmal eyes may
be seen. A pair of raised nuchal ridges extend back to sedger 4. Postsetal lobes
of first four notopodia pointed, the next 5-8 broader and blunt and succeeding
notopodia (fig. iS.j.g) again pointed and slender. Well marked transverse ridges
across the dorsum from setiger 16 to 35. Postsetal lamellae of the first four neuro-
podia pointed but low and rounded on all subsequent feet. Notosetae are winged
capillaries throughout, those of the first foot are long but from setiger 4 they are much
shorter. A stout and strongly curved seta (fig. i8.5.h) in the neuropodium of the
first foot. Tridentate hooks (fig. 18.5.1) with a partial guard or none at all from
setiger 15. Sabre-hke genital setae (fig. 18.5.J) without fibrils become evident from
sedger 15 but may also be present in the anterior segments. Juveniles have brownish
parapodia from segment g-14.
Type locality : Uruguay and South Georgia.
Records: Cape (from 32/1 7/d, and 34/1 7/d, v.d, 34/23/s, d to 33/25/s) ; Natal
(31/29/s, d) and3o/3o/s).
Distribution : Sweden (d) ; Greenland (a) ; Uruguay (s) ; S. Georgeia (d) ;
? Okhotsk Sea ; ? Behring Sea.
PYGOSPIO Claparcde, 1863
Prostomium without lateral peaks but pointed posteriorly. Eyespots present.
In the female, branchiae start well back and are limited to a few segments but in
the male there is an additional single pair on setiger 2. Branchiae united to the
dorsal lamellae. Notosetae arc capillaries throughout. Neurosetac are winged
capillaries anteriorly and hooded hooks posteriorly. Pygidium with four glandular
lobes.
Type SPECIES : Prgnspio elegans CAaparede, 1863.
Pygospio elegans Claparede, 1863
(fig. i8.5.k-o)
Pygospio elfgans Claparede, 1863 : 137, pi. 14 figs. 27-31 ; Fauvel, 1927 : 46, fig. 16 a-h.
A small species 10-15 mm. long with about 60 segments. Prostomium (fig. 18. 5. k)
faintly bilobed in front and pointed posteriorly. Eyes four to eight, irregular.
Notopodial and neuropodial lamellae subequal. Branchiae (fig. 18.5.0) fused to
dorsal lamellae. In the female (fig. 18. 5. 1) they are limited to about eight pairs
starting on setiger 11. In the male (fig. i8.5.k) there are double this number and
in addition diere is a large pair on setiger 2 separate from the notopodial lamellae.
47G
I'Ol.VClllAElA Ol- SOL' lllKR.X AIRIUA
\ a
^<{^ /^"^
Fig. 18.5. Spiojihatus bumbyx. (a) Head, (b) Foiiilh fool, (c) Posterior fool, (d) Enlarged
hook from neuropodiuni or first foot, (e) Hfioded hook from a posterior neuropodium.
Sjiiophanes socderstrumi. (fJ Head, (g) Posterior foot, (h) Enlarged hook from neuropodium
of fir.st foot, (r) Hooded hook from a posterior neuropodium. (j) Sabrc-seta from same foot.
Pygoipio clegaiu (modified from Fauvel, 1927). (k) Head of male, (l) Head of female.
(m) Pygidium. (n) .Anterior foot, (o) Branchiferous foot. .Malncoceros indica. (f) Head.
(q) .Anterior foot, (ri Posterior foot, (s) Hooded hook, (t) Notopodial capillary,
(i:) Sabre-seta.
SPIONIDAE 477
Notosetae are winged capillaries throughout. Anterior neurosetae are capillaries but
four to five bidcntatc hooded hooks are present from setiger 8. Pygidium (fig.
i8.5.m) with four glandular lobes.
Type locality : St. Vasst la Hogue. France.
Records: Cape (34/1 8/i).
Distribution: Arctic; Baltic; Scotland (i) ; English Channel (i, e, s) ; ? Green-
land; Mediterranean; N. Pacific; Okhotsk Sea.
M^Z,^ COC£/?OS' Qiiatrefagcs, 1843
( = SCOLELEPIS auclnnim scd non Blainvillc, 1828)
Prostomium with lateral peaks and a tapered process posteriorly. Eyes may be
present. Branchiae start on setiger i and continue to near the end of the body ;
they arc incompletely united to the dorsal lamellae in anterior segments. Ventral
lamellae not notched. Notoseatae are capillaries throughout. Neurosetae are
capillaries in anterior segments and hooded hooks posteriorly. Pygidium with
anal cirri.
Type SPECIES : Spio vulgaris ]o\vaiion, iQ-ij
Key to Species
1 Hooded hooks bidentate, less than 20 per neuropodium ...... 2
- Hooded hooks tridentate, 20 or more per neuropodium .... M. girardi*
2 Posterior neuropodial lamellae smoothly rounded ..... M.fiiliginosa*
- Posterior neuropodial lamellae with a nipple-like projection .... M. indica
Malacoceros indicus (Fauvel, 1928)
(fig. 18.5.P-U)
Scolelepis indica Fauvel, 1928: 4, fig. 2 g-n ; Fauvel, 1930: 35, fig. 7 g-n.
Body up to 60 mm. long by i to i -5 mm. broad. Prostomium (fig. 18.5.P) broad
and shield-shaped with laterally projecting frontal peaks. Posterior end of cephalic
keel free. Numerous eye specks. Long cirriform branchiae from setiger i. They
are united to the bases of the lanceolate dorsal lamellae which are long anteriorly
(fig. i8.5.q) but are reduced to about one-third the length of the branchiae posteriorly
(fig. i8.5.r). Neuropdial lamellae rounded in anterior feet but develop a nipple-like
projection posteriorly. Notopodial capillaries (fig. i8.5.t) not dotted. Anterior
neurosetae include five to six large capillaries with abruptly pointed tips ; long
bidentate hooded hooks (fig. 18.5.S) appear in the neuropodia from the fordeth
or seventieth segment. Sabre-like setae (fig. 18.5.U) present in posterior neuropodia.
Type loc,\lity : Krusadai Is., Gulf of Manaar.
Records: ? Cape (34/22/d) ; Natal (29/31/i and 27/32/i) ; Mocambique (26/32/i
and 23/35/e).
Distribution : India (e, i), New Caledonia.
478 I'OI.VCHAETA OF SOUTHERN' AFRICA
RHYNCHOSPIO Hartman, 1936
Generally similar to Malacoceros but branchiae start on setiger 2. Prostoniium
w ith frontal peaks. Notosetac arc capillaries throughout. Neurosctae are capillaries
anteriorly and hooded hooks posteriorly. Anal cirri present.
Type SPECIKS : Rlirnchoipin arcnuicola Hartman, ii)36a.
Rhynchospio glutaea (Ehlers, 1897)
(fig. i8.6.a-c)
Scfllecolfpii i^lidtiea Ehlers, 1807 : 8;^, fig. 129-135.
Rhymhoipio glulaea : Day, 1961 : 491.
A small species only 10-18 mm. long. Prostomial peaks ffig. 18. 6. a) directed
obliquely forwards. No free occipital tentacle. A variable number of eye-spots.
First setiger small. Branchiae (fig. i8.6.b) from setiger 2 to the end of the body;
they are strap-like and free from the dorsal lamellae. Neiuopodial hooded hooks
(fig. 18.6.C) are tridentate and first appear in setiger 16. They have two small
teeth side by side abo\e the main fang.
Type locality: Pimta Arenas, South America.
Records: Gape (34/18/s) - ? introduced by ship.
Distribution: Antarctica (d) ; subantarctic (Magellan area (s) and S.
Georgia (s)).
AONIDES C;iaparcde, 1864
Prostoniium pointed anteriorly and with a tentacular process posteriorly. Eyes
may be present. Branchaie start on setiger 2 and are restricted to anterior segments.
They are separate from the notopodial lamellae. Neuropodial lamella not notched.
Gapillaries in both notopodia and neuropodia anteriorly and hooded hooks in both
posteriorly. Pygidium with anal cirri.
Ty'pe species : Aonidts auricularu Cilaparede, 1864 ( = Nerine nxycephala Sars, 1862).
Aonides oxycephala (Sars, 1862)
(fig. i8.6.d-g)
Nerine oxycephala Sars, 1862 : 24.
Aonides oxycephala : Fauvel, 1927: 39, fig. 13 a-e.
Body rounded in section, up to 100 mm. long with 200 well marked segments.
Prostomium (fig. i8.6.d) pointed anteriorly with an occipital tentacle posteriorly.
Four eyes. Peristomium not forming lateral lobes on the side of the prostomium.
Branchiae (fig. i8.6.e) cirriform and completely separate from the large triangular
notopodial lamellae and present from setiger 2 to 25. Notopodial and neuropodial
postsetal lobes flattened and triangular throughout but gradually decreasing in
size posteriorly (fig. ly.G.f). No genital pouches between the parapodia. Narrow
479
Fig. i8.6. Rhynchospio glutaea. (a) Head, (b) Middle loot, (c) Hooded hook, (c') Sabre-
seta. Aonides oxycephala. (d) Head, (e) Branchiferous foot (tenth), (f) Posterior foot.
(g) Hooded hook. Laonice cirrata. (h) Head, (il Branchiferous foot, (j) Lateral view of
three segments showing genital pockets, (k) Hooded hook. Spio filkomis. (l) Head.
(m) Anterior foot, (n) Posterior foot, (o) Hooded hook. Dispio magna, (p) Head.
(q) Anterior foot, (r) Posterior foot, (s) Speckled seta, (t) Hooded hook.
.[.".o 1'()I,m:hak lA ov soi'iHr.RX ArRic;\
bladcd capillaries in bulli rami of the parapodia anteriorly. Between three and four
bidentate hooded hooks (fig. i8.6.g) appear in the notopodia from sctiger 35-42
onwards and four to fi\e in the neuropodia from setigcr 32-35 onwards. Pygdium
with four to eight anal cirri.
Type locality : Floro, Xorway.
Records: Cape (from 33, 17's and 34/23/e to 33/25/s) ; Mocambique (26 32 i) ;
Madagascar (s).
DisTRliii-TiON : N. Atlantic from Norway, English Channel (e, i) to Morocco (s) ;
Mediterranean; ? Persi.m Gulf (s).
LAONICE Malmgren, 1867
Prostomium bioadly rounded in I'mnl and having .m erect occipital tentacle
posteriorly. Two eyes. Branchiae free from the notopodial lamellae and present
from sctiger 2 to the middle of the body. Neuropodial lamellae not notched. Genital
pouches between the parapodia in the middle of tlie isody. Xotosetae are capillaries
throughout. Xeurosetae arc capillaries anteriorly and hooded hooks posteriorly.
Pygidiimi witli anal cirri.
Type sPK(_:iES : .A'tiiiic ciiralu Sars, 1851.
e cirri
(fig. i8.6.h-k)
Laonice cirrata (Sars, 1851)
Serine citrata Sars, 1O51 : 64.
Laonice cirrala : Fauvel, 1927: 3R, fig- 12 a-e.
A large species reaching 100 mm. in length for 160 segments. Prostomium (lig.
i8.6.h) broadly rounded in fiont with two large eyes. An occipital tentacle posteriorly
followed by a long dorsal ciliated ridge extending over 28-40 segments. Mem-
braneous ridges across middle segments. Branchiae (fig. i8.6.i) long and cirriform
and present from setigcr 2 to 40. Notopodial lamellae large and auricular in the
branchial region. Lateral membraneous pockets (fig. 18.6.J) between the parapodia
from sctiger 25 in sexually mature specimens. Bidentate hooded hooks (fig. i8.6.k)
appear in the neuropodia from sctiger 45.
Type LOf:ALiTY : Northern Norway.
Records: C^ape (from 31/16/d and 34/18/3 to 36/21/d and 33/28/s).
Distribution : Cxismopolitan from the Arctic to the Antarctic between shallow
and abyssal depths.
SPIO Frabicius, 1785
Prostomium rounded ; lateral peaks never present. Eye spots often present.
Branchiae fmni sctiger i to near the end of the body. They are partly fused to the
nritcipodial lamellae anteriorly and ahnost separate posteriorly. No accessory
SPIO.MDAE 481
branchiae between notopodia of middle segments. Neuropodial lamellae not notched.
Notosetae are capillaries throughout. Ncurosctac arc capillaries anteriorly and
hooded hooks posteriorly. Pygidium with anal cirri.
Type SPECIES : J/ereis Jilicornis MiXWcr, 1766.
Key to Species
I Neuropodial hooded hooks bidentate and start from setiger lo-i i. Prostomium not brown
S. filicomis
- Neuropodial hooded hooks tridentate and start from setiger ii. Prostomium brown . S. sp.*
Spio sp.*
(NAD.29.W)
Spio filicomis (Miiller, 1776)
(fig. 18.6.I-0)
J^'ereis fdicomis yiuWer , 1776: 218.
Spio filicomis : Fauvel, 1927: 43, fig. 15 a-g.
A small worm reaching 30 mm. for 90 segments, but usually smaller. Prostomium
(fig. 1 8.6.1) blunt anteriorly and pointed posteriorly. Branchiae (fig. iB.S.m)
attached to the bases of the notopodial lamellae anteriorly but free posteriorly
(fig. iB.G.n). Notopodial lamellae auricular. Neuropodial lamellae broadly oval
anteriorly but narrower posteriorly. Neuropodial hooded hooks (fig. 18.6.0) bi-
dentate, about eight to ten per neuropodium, and start from setiger lo-ii. Two
inferior sabre setae in posterior neuropodia.
Type locality : Denmark.
Records: Cape (33/18/i and 34/18/s).
Distribution : Arctic ; North Atlantic from Greenland (s, d) and the North
Sea (i, s) to the Bay of Biscay ; Pacific from the Behring Sea and Japan to California
(i).
DISPIO Hartman, 1951
Prostomium fusiform, pointed anteriorly and without frontal horns but with an
occipital ridge extending posteriorly. Peristome enfolding the sides of the pro-
stomium. Branchiae from setiger i to near the posterior end ; they are partly or
completely fused to the notopodial lamellae anteriorly but are mainly free posteriorly.
Accessory branchiae on the posterior part of the notopodia of middle and posterior
parapodia. No hooded hooks in the notopodia but present in the neuropodia.
Pygidium as a ventral cushion below the anus.
Type species : Dispio uncinata Hartman, 1951a.
*Only one broken specimen. Prostomium blunt and brown. Peristome brown. Hooded hooks tridentate
with the third tooth much smaller than the second. Hooks from setiger 1 1 onwards, about eight per neuro-
podium and accompanied by two to three inferior sabre setae with speckled blades.
482 l'OI,\C:H.\FTA OF SOUrilF.RX AFRICA
Key to Species
1 Accessory braiKliiac as minute digitiforni branches. Anterior branchiae serrated /). uncinata*
- Accessory branchiae lamellar, not branched. Anterior branchiae not serrated . D. magna
Dispio magna (Day, 1955)
(fiR. i8.6.p-t)
Spiu mcgriJis Day. 1933 : 413, hg. 2 a-g.
A large species reaching 100 mm. by 3 mm. for 200 segments. Prostomium (fig.
i8.6.p) smoothly conical with a ridge extending back to sctiger 2. No eyes. Peri-
stome closely applied to the sides of the prostomium. Branchiae from setiger i to
the posterior end. They are completely fused to the notopodial lamellae anteriorly
(fig. i8.6.q) but from the middle of the body the distal part of the notopodium
separates as a triangular lobe (fig. i8.6.r). Accessory branchiae as minute lamellae
(not digitiform lobes) behind the neuropodia from about sctiger 35 onwards.
Neuropodia with a small oval preselal lobe and a dorsoventrally longer postsetal
lobe. This develops a free superior projection in later segments which project upwards
towards the notopodium. Some notopodial and neuropodial capillaries with speckled
axes (fig. 18.6.S). About nine hooded hooks in the neuropodia from setiger 30-44
onwards according to size. In juveniles they have faintly bilobed ends (fig. 18. 6. t)
but in adults they are bluntly unidentate. One to two sabre setae at the inferior
edge of the neuropodia from setiger 14 onwards.
Type locality : Simonstown, South Africa.
Records: Clape (34/18/i).
Distribution' : No other records.
SCOLELEPIS Blainville, 1828
( = NERINE ]oh.ni\.on, 1838 et auctoriim*)
Prostomium often pointed anteriorly and with an occipital tentacle or keel
posteriorly. Eyes present or absent. Branchiae partly or completely fused to the
notopodial lamellae ; they start on setiger 2 and continue to near the posterior
end. Neuropodial lamellae simple anteriorly but more or less notched posteriorly.
Winged capillary setae in both rami of the parapodia anteriorly but hooded hooks
appear in both rami posteriorly. Pygidium as a ventral glandular cushion.
Type SPECIES : Liimhricus squamaiiis Muller, 1806.
Kly to Species
I Hooded hooks unidentate ......... S. lefebi<rei
— Hooded hooks bidentate ......... S. squamata
*Ttic synomy oi Scolelfpii was discussed by Pettibone (1963a).
SPIONIDAE 483
Scolelepis lefebvrei (Gravier, 1905)
(fig. i8.7.a-b)
J^erine Lefebvrei Gravier, 1905: 43, pi. 2 fig. 185, text-figs. 322-326; Day, 1962 : 648.
Body large, up to 80 mm. long by 3 mm. broad for 150 segments. Prostomium
(fig. 18.7.3) pointed anteriorly and extending posteriorly as a tapered keel which
reaches setiger 2. No ft'ee occipital tentacle. Four eyes in a transverse row. Lateral
folds of the peristomium enclose the sides of the prostomium. Branchiae from setiger
2 to the last few segments. Neuropodial lamellae smoothly oval anteriorly but deeply
notched posteriorly. Notopodial lamellae almost completely fused to the branchiae
anteriorly but become more and more separate posteriorly and by the sixtieth
segment they are joined only at the base. The two lamellae become joined across
the back by a dorsal fold from the middle of the body onwards. Anterior notosetae
are numerous winged capillaries ^vith punctuations towards the distal end. Posterior
notosetae few, fine and elongated. No hooded hooks in the notopodia of posterior
segments. Anterior neurosetae are winged capillaries but unidentate hooded hooks
(fig. i8.7.b) appear about setiger 40 and increase to a maximum of si.x posteriorly ;
meanwhile the capillaries decrease to a small tuft of fine setae. Pygidium with a
ventral cushion below the anus.
Type locality : MaskaU Is. (Red Sea).
Records : Not recorded from southern Africa.
Distribution: Red Sea (i), Madagascar (i).
Scolelepis squamata (Miiller, 1806)
(fig. i8.7.c-h)
Lumbricus squamatus Muller, 1806 : 39.
Lumbricus cirrattilus Delle Chiaje, 1825 : 196.
Merine cirTalulus: Fauvel, 1927: 36, fig. 11 g-n ; Day, 1955: 412, fig. 1, j.
Length up to 80 mm. for 200 segments. Prostomium (fig. 18.7.C) pointed anteriorly
with four to six eyes in a row and a well marked occipital ridge reaching setiger 2.
Notopodial lamellae fused to the branchiae anteriorly (fig. i8.7.e) but auricular
and mainly free posteriorly (fig. i8.7.f) with the inferior margin produced dowii-
wards towards the neuropodium. Bidentate hooded hooks (fig. i8.7.h) in the
neuropodia from setiger 30-35 onwards and in the notopodia from setiger 60. A
maximum of twelve neuropodial hooks. Pygidial cushion (fig. i8.7.g) small,
broader than long.
Ty'pe locality : Denmark.
Records: South-west Africa (26/15/i) ; Cape (from 32/18/e and 34/18/e, i to
33/25/e) ; Natal (29/3 1 /i, 27/32/e) ; Mocambique (26/32/i, 23/35/e).
Distribution : Atlantic from Scotland (i) and English Channel (i) to Senegal (i)
and North Carolina (i) ; Mediterranean ; ? India, Madagascar (i) ; Pacific from W.
Canada to Southern California.
4»4
I'OI.VCHAKTA OF SOL THERX AIRICA
4
sk\
I-i.;. 18.7. ScoltUpii hftbvid. (Ai Hcatl.
'D' Entire worm i 1.5 tiines natural size
diiim. ill) Hooded hook,
foot. :l Middle loot.
j Hooded h'nik. Siolelifii^ \quninaia. ic.) Head.
IE) .\nierior tciot. (t-i l*osterior Riot. u;,i Pygi-
.\'ifi'ii(it'\ giUitr!\fi. Ill Heati. (j) Hootled hook. (K) .Anterior
till' ii.irapodia
segments. .\o
NERINIDES McsnW, 1896
Prostomiiim bkiiU nr priiiited anteiiorly and with an occipital tentacle posteriorly.
Eyes present or absent. Branchiae present from setiger 2 and ]iartlv or coinplctely
fused to the notopodial lamellae : they continue over most of the bodv. Xmiropodial
lamellae not notched. \Vingcd capillary setae in botli rami of
anteriorly, but hooded hooks appear in the ncuropodia of later
notopodial hooks. Pygidium as a ventral cushion.
Typk spkciks : .\Ial(icocno\ longiiinti'n Qiiatrcfages, 1843.
Ki V lo .Species
I Hooded hooks witli one denticle above the main fang ...... 2
Hooded hooks with two denticles above the main fang . . . . A^. tridentata*
Hooded hooks with three dentic les above the main fang. (P<;)Sterior branchiae tlag-shai3ed,i
N. gilchristi
SPIOXIDAE 485
2 Branchiae united to the posterior lamellae by a pleated web. Neuropodial lamellae not
notched ........... . N, cantabra*
- Branchiae separated from notopodia lamellae and tipped with two to six clavate papillae
in middle segments. Neuropodial lamellae notched . . . . A^. williami*
Nerinides gilchristi Day, 1961
(fig. i8.7.i-l)
Nerinides gilchristi Day, 1961 : 491, fig. 5 a-d.
Length about 25 mm. for 100 segments. Prostomium (fig. 18.7.1) sharply pointed
anteriorly and with an erect occipital tentacle posteriorly. Two pairs of subdermal
eyes. Short nuchal bands extend back from the prostomium to setiger 2. First
setiger small but possesses both notosetae and neurosetae. Branchiae from setiger 2 ;
the anterior ones (fig. iB.y.k) are large, strap-like organs completely fused to the
notopodial lamellae ; from setiger 16 onwards a notch appears between the branchia
and the lamella and deepens until the two are mainly separate. The branchia then
becomes a narrow stalk with a flag-hke end (fig. 18.7.I) and the lamella becomes
broadly triangular and lateral in position. Neuropodia ear-shaped throughout.
Anterior notosetae and neurosetae are winged capillaries. Punctate setae absent.
Posterior notosetae unknown. Neuropodial hooded hooks appear on setiger 16-18
and soon replace the winged capillaries. Individual hooks (fig. 18.7.J) show three
long teeth above the main fang. No inferior sabre setae. Body usually broken in
the middle and the characters of the posterior segments are unknown.
Type locality : Saldanha Bay, South Africa.
Records: South-west Africa (26/15 's); Cape (from 32/17/d and 32,'18/s to
34/22/s) ; Natal (30/30/s).
Distribution : Endemic.
PRIONOSPIO Malmgren, 1867
Prostomium rounded or pointed anteriorly and supported by the buccal segment
which may develop wing-like lateral expansions. No posterior occipital tentacle.
Eyes present or absent. Notopodial lamellae often large and erect in anterior
segments. Branchiae free from the notopodial lamellae and restricted to a few
anterior segments. They are often pennate and start on setiger i or 2. Neuropodial
lamellae not notched. ^V■inged capillaries present in both rami of the parapodia
anteriorly but are partly replaced by hooded hooks in both rami posteriorly.
Pygidium with a median cirrus.
Type species : Prionospio steenstrupi Malmgren 1867.
Key to Species
I Gills start on setiger 2 and always exceed four pairs ; none pinnate .... 2
- Gills start on setiger i or 2 and never exceed four pairs ; at least one pinnate ... 3
4Sb P()l.\ (.iHAETA OF .SOUlUliRN AKRICA
2 Gills increase in size to sctigcr 4 and then decrease. Four eyes (fig. 18. 8. a)
P. cirri/era (p. 486)
- Gills all unil'orm in size and about equal to the notopodial lamellae. No eyes (tig. 18. 8. e]
P. cirrobranchiata (p. 488)
3 Gills start on setigcr i and number three pairs, all pinnate. Setiger i not small. (Wing-
like peristomial expansions enfold the sides of the prostomium) (fig. 18.8. i)
P. pinnata (p. 48!!)
- Gills start on setiger 2 ; setiger i small ......... 4
4 Gills tuo to three pairs, pinnate (fig. 18. 8. m) .... P. sexoculata (p. 48(1)
- Gills four pairs, not pinnate .......... 5
5 First pair of gills pinnate, the rest smooth. (Fourth pair of gills slender and tapering, i
Lateral pockets between anterior neuropodia (fig. 18. 9. d) . . P. ehlersi (p. 490)
- Large fourth pair of gills, pinnate, the rest smooth (fig. 18.9.JI. (Hooks w ith a single tooth
above the main fang.) No lateral pockets . . . . P. saldanha (p. 492)
- Both first and fourth pair of gills pinnate, the other two smooth ..... 6
6 Lateral pockets between anterior neuropodia. .A low ridge across setiger 7 P. bocki (p. 490)
- No lateral pockets ............ 7
7 .\ membraneous ridge across setiger 7. Eyes obvious . . P. malmgrcni fp. 492)
- Xo membraneous ridges across anterior segments. Eyes indistinct or absent
P. steenstrupi (p. 489)
Prionospio cirrifera Wiren, 1883
(fig. i8.8.a-d)
Prionospio cirrifera Wiren, 1883: 409; .Soderstroni. 1920; 237, fig. 134 a-b, fig. 146; Fauvel,
1927: 62, fig. 21 k-n ; Day, 19G1 : 487.
Length 30 mm. for ijo .segments. Prostomium (fig. i8.a) bluntly pointed in fi'ont
with a keel posteriorly which reaches setiger 2. Four subdermal eyes not easily seen
in large specimens. Buccal segment more or less fiised to setiger i which is reduced.
Long, smooth, tapered gills from setiger 2 to 10. Longest gill on setiger 2-4. It is
three times the length of the corresponding notopodial lamella. Anterior nrjtopodial
lamellae (fig. 18. 8. b) pointed; the first one small but the second and later ones
increase in size to the si.xth and then decrease. Posterior lamellae (fig. 18. 8. c) sub-
equal. No dorsal ridges. Ncuropodial lamellae small and triangular with lateral
pouches between them from about the 5th foot in mature .specimens. Hooded hooks
in the neuropodia from setiger 15-18 and in tlie notopodia after setiger 30. Hooks
\vith two series of four long teeth above the main fm in northern forms (fig. iB.S.d')
but only two paired teeth in .Soutfi African forms (fig. ll).8.d).
Type loc;ai.itv : Behring Sea.
Reciords : Cape (from 34 i8,s to 34'23/s).
DiSTRiBL'Tio.N : Arctic; North Atlantic from Greenland (a). North and South
America (d, vd, a) and Sweden (d, vd) to the North Sea (d), English C'.hannel and
PiirtiiL;al ; ? India; North Pacific from the Behring Sea to southern California.
SPIOXIDAE
4»7
Fig. i8.8. Prionospio cirrifera. (a) Head and branchiferous region, (b) Branchiferous foot.
(c) Posterior foot, (d) Hooded hook of Cape specimen, (d') Hooded hook according to
Soderstrom, 1920. Prionospio cirrobranchiata. (e) ,\nterior end. (f) Branchiferous foot.
(g) Posterior foot, (h) Hooded hook. Prionospio pinnata. (i) .\nterior end. (j) Posterior
foot, (k) Notopodial capillary, (l) Hooded hook. Prionospio sexoculata. (m) .\nterior end.
(n) Middle foot, (o) Hooded hook.
488 I'OIA'CHAI. lA <)1' SOL rHKRN AIRIC.A
Priottospio cirrobranchiata Day, 1061
(tig. I !!.!!. c-h)
Priotwsjiio cirtiihjiiru'fiialii Day, 1961 : 481^, fig. 4 a-d.
B(id\' small and thread-like, only 15 mm. long with 60 segments. Prostomium
(fig. io.8.e) depressed, square in front and produced back as an inconspicuous
plate btit not keeled posteriorly. Two pairs of small i-yes not \isible in adults. Aisout
elcN'cn to twehe pairs of smooth gills starting on setiger 2. Gills (fig. iS.B.f) uniform
in length and two to three times as long as the notopodial lamellae. Notopodial
lamellae triangular, pointed and subequal alter the first one which is small. Ncuro-
podial lamellae small but similar in shape to the notopodial ones, even in posterior
segments (fig. 18. 8. g). About five heeded liooks in the neuropodia from setiger 18-19
onwards. Each hook (fig. 18. 8. h) \\ilh a single tooth above the main fang. Four
subequal and cirri.
Type lociality ; Ofl" Saldanha Bay, Soulh Africa.
Records : Cape (from 32, 17 d to 36,21 d and 34/21,'s to 34 '22,'d).
DisTRlBUTlo.x : North C^areilina (si.
Prionospio pinnata Ehlers, 190 1
(fig. i8.8.i-l)
Prwu'isf'io jnnnala Ehlers, KjOi, iG^; Hartman, if)fj(»; 114. pi. q figs. i-~,.
A large species reaching 60 mm. Prostomiimi (fig. 18. 8. i) bhmtly pointed and
produced back as a ridge enfolded by large, wing-like lateral expansions of the
peristome. Eyes seldom visible in adults. l\ memljraneous ridge across the dorsum
immediately behind the origin ol the palps. First setiger well developed. Three
pairs of large piimate gills on setigers 1-3 but one or more are often lost. Postsctal
lamellae of notopodia large and pointed on the first five setigers and smaller and
more rounded thereafter; from setiger 21 onwards to the middle of the body they
unite across tlie dorsum to form low ridges. Xeuropodial lamellae prominent and
pointed anteriorly, low and rounded posteriorly (fig. iB.S.jj. \Vinged capillary set.ic
(fig. i8.8.k) in both rami anteriorly. Hooded hooks appear in die neuropodia from
setiger 9 ; they attain a maximum of 12-15 P""'" neuropodium and are accompanied
by an inferior sabre set.i. Individual hooks (fig. 18. 8.1) with f lur pairs of teeth above
the main fang.
Type locality: Taleahuano, Clhile.
Records: South \Vest Africa (23/14/s, d to 26/15/s) ; Cape (from 32,i7:d and
33, i8,s, d to 35,20,(1 and 34,26 clj ; Natal (30 30/3 to 29'3i,s).
Distribution : Atlantic from North Clarolina (s, d) and Morocco (s, d) along
tropical west Africa (s, d) to S. Africa (s, d) ; tropical Indian Ocean (s, d) ; Pacific
from \V. Canada and Japan to C'liile (s) ; New Zealand.
SPIOXIDAE 489
Prionospio sexoculata Augencr, 1918
(fig. i8.8.m-o)
Prionospio sexoculata Augener, 1918: 405, pi. 6 figs. 159, 172; te.Kt-fig. 52 (partim.) ; Day, 1955:
414 (partim.).
A small species 10-20 mm. long. Prostomium (fig. i8.8.m) rounded in fi-ont
and supported by small pcristomial ridges on either side. Six eyes. Setiger i very
small and widiout branchiae or notosetae. Two pairs of pinnate branchiae on setigers
2 and 3. Triangular notopodial lamellae on setigers 3-5 and smaller, more rounded
ones later (fig. i8.8.n). No membraneous dorsal crests. Tridentate hooded hooks
(fig. 18.8.0) appear in the neuropodia from setiger 14-16 and in the notopodia from
setiger 30. Inferior sabre setae in the neuropodia from setiger 12.
Type loc.'\lity : Walvis Bay, South 'West Africa.
Records: South West Africa (22/14/s and 26/1 5/s) ; Cape (from 33/17/s and
33/18/i, s) ; Natal (27/32/e).
Distribution : Endemic.
Prionospio steenstrupi Malmgren, 1867
(fig. 18.9.0-r)
Prionospio steenstrupi Malmgren, 1867: 202, pi. 10 fig. 55; Fauvel, 1927: 60, fig. 21 f-i.
Prionospio malmgreni var. duhia Day, 1961 : 489, fig. 3 j-n.
Length up to 45 mm. for 100 segments. Prostomium (fig. iS.g.o) rounded in
front and continued as a raised keel which reaches setiger 2. Erect lamellae of the
peristome are fused to the notopodia of setiger i. Eyes indistinct or absent. Four
pais of gills on setigers 2-5 ; the first pair pinnate and much larger dian the rest,
the second and third pairs smooth, stout and no longer than the notopodial lamellae,
the fourth pair (fig. i8.g.r) again pinnate but short. Notopodial lamellae triangular
and pointed ; they increase in length to the fourth but then decrease and later ones
are low and rounded (fig. iS.g.p). No membraneous ridges across the dorsum.
No genital pockets between anterior neuropodia. Neuropodial lamellae low and
oval after the first few. Hooded hooks in the neuropodia from setiger 18-19 and in
the notopodia from setiger 50. Each hook (fig. i8.9.q) with four double rows of
teedi above the main fang. An inferior sabre seta in the neuropodia from sedger 1 8.
Type locality : Iceland.
Records: South West Africa (28/14/d) ; Cape (from 32/1 7/d and 34/i8,s, d to
36/2 1 /d and 34/26/d ; Natal (29/3 i/s, d).
Distribution : North Atlantic from Greenland and Iceland to Norway and the
Nordi Sea (s) ; North Carolina (s).
490 POLVOHAETA OF SOUTHERN AFRICA
Prionospio bocki Scklcrstroni, 1920
Prionospio bocki S'dderslTom, 1920: 234, figs. 142-143.
LcngtJi up to 20 mm. Prostomium blunt in front and tapering back as a kcrl
reaching setigcr 2. Two pairs of eyes, the anterior pair small, the posterior pair
large and comma-shaped. Sctiger i small, but with broad notopodial and neuro-
podial lobes. Four pairs of gills on setigers 2-5, the first pair large and beset with
numerous short papillae, the second and third pairs short and smooth and the fourth
pair tapered and fringed with a row of short papillae. Notopodia of setigers 3 and 4
arc the largest and subsequent notopodia decrease in size. From sedger 6, they are
united across the dorsum by low, inconspicuous ridges. Ncuropodial lamellae all low
and oval ; from setiger 2 onwards they arc united by membranus genital pockets
which become conspicuous after the first 10 segments. Hooded hooks with 4-5 pairs
of long denticles above the main fang, appear in tlic neuropodia from setigers 16-18
onwards.
Type locality : Kobe Bay, Japan.
Records: Madagascar (s).
DisTRiBiTiox ; Japan (s).
Prionospio ehlersi Fauvel, 1936
(fig. i8.9.d-f)
Prionospio ihlersi Fauvel, 1936: 61, fig. 4 a-e.
Length up to 20 mm. Prostomium (fig. i8.9.d) expanded anteriorly and narrowed
posteriorly forming an elevated keel between the peristomial folds. Two to four
small eyes and black pigment flecks. First setiger with small notopodial and ncuro-
podial lobes. Four pairs of gills on setigers 2-5 ; tJie first pinnate, the second and
third short and smooth and the fourth long, smoodi and tapered. Notopodial
lamellae triangular and pointed ; they reach maximum size on setigers 3-5 and then
decrease. A low membraneous ridge unites the notopodial lamellae for 20-30 seg-
ments starting on setiger 5 or 6. Hooded hooks appear in the neuropodia on setiger
19-21 and in the notopodia from sctiger 37. A punctate sabre seta appears in the
inferior neuropodium from setiger 19-23 onwards. Genital pockets (fig. iS.g.e)
between the neuropodia of adult specimens from setiger 2 for about 20 segments.
Hooks (fig. l8.9.f) with numerous teeth. Neuropodia lamellae rounded, at first
longer than broad, later oval.
Type loc.\lity' : Dredged off Morocco.
Records: Natal (30/30,'s, 29/31/^) ; Mocambicjue (26/33/d).
Distribution: Morocco (s, d).
491
Fig. 18.9. Prionospio malmgreni. (a) Anterior end. (b) Posterior foot, (c) Hooded hook.
Prionospio ehlcrsi. (d) Anterior end. (e) Lateral view of middle segments to show genital
pockets, (f) Hooded hook. Prionospio saldanha. (g) Notopodial capillary, (h) Sabre-seta.
(i) Neuropodial capillary, (j) .interior end. (k) Sixth foot, (l) Middle foot, (n) Hooded
hook. Prionospio steenstrupi. (o) Anterior end. (p) Middle foot, (q) Hooded hook.
(r) Fourth gill on fifth foot.
492 P0I.VC:HAF,TA Ol' SOUTHERN AFRIC:A
Prionospio saldanha Day, 1961
(iig. i!;.9.g-n)
Piionosjiio ialdanha Day, 1961 : 485, fig. 3 a-h.
Body thread-like, up to sf, mm. Ions for 80 segments. Prostomium (fig. 18.9.J)
rounded anteriorly and produced back as a keel reaching setiger 2. Four eyes.
Setigcr I small and fused to the peristomium. Four pairs of gills on sctigers 2-5;
slie first three pairs smooth and the iburth, larger pair pinnate. Anterior notopodial
lamellae pointed and face outwards (fig. iB.g.k) ; they reach maximum size on
setigers 6-9 and then decrease and become rounded. From setiger 20 onwards the
lamellae arc united by dorsal ridges (fig. 18.9.I). A maximum of six hooded hooks
in the neuropodia from setiger 15 onwards and in the notopodia from setiger 25-35.
Each hook (fig. 18. 9. n) with a single tooth above the main fang. An inferior sabre
seta (fig. 18. 9. h) in the neuropodium from setiger 12.
Xvtc : A closely related form P. caspcisi Laubier (19611 from the Mediterranean
differs in having a square jirostomium, a dorsal membraneous ridge on segment 7
only, and notopodial hooks nc\er before segment 411.
TyPK loi_:ai,itv : Saldanha Bay. South Africa.
Records: South \Vest Africa (2G/i5,sj ; (Cape 33,18,1, s to 34/2 2 /d ) ; Xatal
(30/30/5)-
Distribution : Endemic.
Prionospio sp.
Prionospio sp. Day, 196313: 418.
Only jiuenilcs measuring 4 mm. are known. Prostomium sho\el-shapcd with
four eyes. No wing-like peristomial expansions. Setiger i small and without gills.
Very long cirriform branchiae on setigers 2-10. Anterior notopodial lobes small,
later ones inconspicuous. No dorsal crests. Neuropodial hooded hooks seen on
setiger 15 but possibly occiu' earlier.
Prionospio nialmgreni Cllaparedc, 1870
(fig. l8.9.a-c)
Priotioijiio malmgreni Claparede, 1870: 333, pi. 22 fig. 3: Fauvcl, 1927: Gi, fig. 21 a-c : Day,
!g63a : 418.
A thread-like species about 25 nmi. long for 60 segments. Prostomium (fig.
1 8. 9. a) bluntly rounded in front and produced back as a ridge which reaches
setiger 2. Four eyes, the second pair large and elongate. Peristome with small lateral
folds fused to setiger i. First setiger small. Four pairs of gills on setigers 2-5 ; the
first and fourth pairs arc botJi long and pinnate and tJic second and third pairs are
both short and smooth. Anterior notopodial lamellae triangular ; the largest is on
.setiger 4 Init later ones tlecrease and from setiger 8 onwards they are low and
SPIONIDAE 493
rounded (fig. iS.g.b). A well marked membraneous ridge unites the lamellae across
setiger 7 and smaller incomplete ridges are present on the next few segments.
Ventral lamellae all low and roimded. No genital pockets. Pluridentate hooded
hooks (fig. iS.g.c) appear in die neuropodia from setiger 12-14 and in the notopodia
after setiger 40. An inferior sabre seta in the neuropodium from setiger 12.
Type locality : Gulf of Naples.
Records: Cape (33/1 7/s, d, 34/25/s) ; Natal (30/30/s).
Distribution : Atlantic from North Carolina (s) and the North Sea (s) to Madeira
(s) ; Mediterranean (s) ; S. California (s) ; Japan.
494 i'oi.vi:ii \i; r A ok souther.x ai-rica
Family MAGELONIDAE Cunningham and Ramagc, 1888
Body slender and divided into two distinct regions. Prostomium flattened,
roughly oval in outline and lacks eyes. Two very long palps which arc papillose
distally. Proboscis large and eversible but unarmed. No branchiae. Parapodia
biramous with lamellar lobes. Thoracic region of nine segments with capillary
setae in the first eight segments but sometimes speciahsed setae in the ninth. Ab-
domin.il setae are mainly hooded hooks. Pygidium \vith anal cirri.
Records from southern Africa
Ma ad on a capensis Day . . . . 5'C's, — Nsd
Magelona cincta Ehlers ..... 15CS, 45 PiNi,
46CS, Pi, 51CS
Mas,t'lona papiUicornii Miillcr .... 44Ci, 51CS
Remarks: This family contains only one valid genus, Maatiuna. Jones (1963)
gave a key to the 21 known species.
The magelonids burrow in muddy sand and use the spade-like head and large
distensible proboscis to force their way through the substrate. Like the spionids
they are detritus feeders and the food particles are gathered by a pair of very elon-
gated palps which bear sticky papillae. There is no food groove along the palp
instead the particles are brought to the mouth by muscular contraction. The gut is
usually full of sand grains so there appears to be little selection of what is ingested.
MAGELONA Muller, 1858
Body di\ided into an anterior region with nine setigers and a posterior region of
nuinerous setigers. Prostomium large and flattened. Two long papillose palps.
Parapodia biramous with lamellar notopodia and neuropodia. No branchiae. Setae
are simple capillaries anteriorly, mainly hooded hooks posteriorly and on the ninth
setiger, sometimes special setae. Anal cirri present.
Type SPECIES : Magelona papillicunili MuUer, 1858.
Key to Species
1 Setiger 9 with specialised sclao ha\'ing a subtcrminal expansion (fitf. ifj.i.d). Prostoniiiun
spathiilate and smoothly oval. Hooded hooks with two tcetli side by side above tiie
main (an^ ......... AI. papillicornis
- .Setiger 9 without speciahsed setae. I'rostDmiiim with antero-lateral angles or projections.
Hooded hooks with two teeth side by side above the main fang . .... 2
2 .\ red band on setigers 5-8. Parapodial lamelhie of abdomen ligulate . . M. cincta
— -Xo red band. I*arai>odi.il lamellae of abdomen o\-al and rc^tiistricted at the base Af. capensis
MAGELONIDAE 495
Magelona papillicornis Miiller, 1 858
(fig. ig.i.a-d)
Magelona papillicomis MiiUer, 1858: 216; Fauvel, 1927: 64, fig. 22 a-h.
Length up to 170 mm. with 150 segments and uniformly white in colour. Pro-
stomium (fig. ig.i.a) bluntly spear-shaped with a median thickening. Anterior
end smoothly rounded. In the anterior region the notopodia (fig. ig.i.b) have
low, rounded presetal hps and much larger tongue-shaped lamellae. In addition
sctigers 1-8 have a small, erect digitiform lobe above the notosetae. The anterior
neuropodia have smaller lamellae than the notopodia. The presetal lamella of the
neuropodium is produced inferiorly below the neurosetae. The postsetal lip is
small except on setigers 7 and 8. Setiger 9 is short with subcqual presetal and post-
setal lobes in both rami. Abdominal segments from setiger 10 onwards (fig. ig.i.c)
have equal, tongue-shaped postsetal lamellae in both rami curving towards one
another. Lateral pockets are present on several segments after setiger 25.
Setigers 1-8 have narrow-winged capillaries in both rami. Setiger 9 has a few
capillaries and numerous setae each with a subterminal expansion preceding the fine
capillaries and numerous setae each with a subterminal expansion preceding the fine
tip (fig. 19. 1. d). Abdominal setae are rows of hooks each with two teeth side by
side above the main fang.
Type locality : St. Catherine Is., Brazil.
Records: South West Africa (26/15/s and 28/18/s) ; Cape from 33/18/s and
34/23/e, s to 33/27/s.
Distribution : Atlantic from Scotland (i, s, d) and English Channel (e, i) to North
Carolina (s) and Brazil and tropical \V. Africa (s) ; Mediterranean ; Madagascar (i).
Magelona cincta Ehlcrs, 1908
(fig. ig.i.e-k)
Magelona cincla Ehlers, 1908 : i 1 1, pi. 15 figs. 9-12 ; Wilson, 1958 : G20, figs. 2-3.
Body (fig. ig.i.e) up to 30 mm. long with a reddish or purple band from setiger
5 to 8. Prostomium (fig. ig.i.f) a truncate triangle with a pair of muscular strands
diverging anteriorly to end in antero-lateral projections. The whole prostomium
about as broad as long. The anterior region has shghtly larger lamellae on the
neuropodia. The notopodial postsetal lamellae (fig. ig.i.g) are ligulate and there
is no erect superior lobe or "dorsal cirrus". The neuropodial lamellae arc more
infrasetal than postsetal and that on setiger 8 is distinctly larger than the notopodial
lamella. Presetal lamellae absent. Setiger g short. Abdominal parapodia from
setiger 10 onwards (fig. ig.i.h) have subequal, tongue-shaped postsetal lamellae
on both rami which cur\'e towards one another and end in pointed tips. Lateral
pocket-like folds are present in front of some posterior parapodia in adults.
Setigers 1-8 have simple capillary setae in both rami (fig. ig.i.i). Setae of setiger
4'j6
POLVCHAErA OF SOUTIIERX AFRICA
Iif.. I'j.i. M<i[^floru! papilliiornl^. (a) Head, (ii) Third foot. (( ) Abdttniinal IViot. (r>) XoUi-
seta from ninth foot. Alagfhma cincla. (e) Entire worm (three times natural size), (f) Head.
(g) Thoracic foot, in) Abdominal f(>ot. (i) Thoracic notoseta. (j) Hooded hook.
(k) Notoseta from ninth foot. Magclona cajicmis. (l) Tlioracic foot, (m) Part of palp.
fx) Anterior end and part of abdomen, (o) Ninth foot, (p) Abdominal foot. (Q) Hooded
hook, (r) Notoseta from ninth fciot.
MAGELOMDAE 497
9 (fig. ig.i.k) similar to those of setigcr 8. Abdominal setae are rows of hooks with
two teeth side by side above the main fang (fig. ig.i.j).
Type locality : Algoa Bay, South Africa.
Records: Cape (from 34/18/3 to 33/25/s) ; Natal (30/30/s and 29/31/i) ;
Mocambique (26/32/i and 23/35/e).
Distribution : Morocco (s) and tropical western Africa (s, d).
Magelona capensis Day, 1961
(fig. 19.1.1-r)
Alagelona capensis Day, 1961 : 495, fig. 6 a-h.
Body about 35 mm. long for 40 segments, uniformly pale in colour without a
red band on setigers 5-8. Prostomium (fig. ig.i.n) 1-3 times as long as broad with
antero-lateral corners. Notopodia of setigers 1-8 (fig. 19.1.I) with a small superior
lobe united to a much larger inferior lobe. Neuropodia with a tapered inferior
lobe. Setiger g without a superior lobe to the notopodiimi but with a small pro-
jection below the neuropodium (fig. 1 9.1.0). Abdominal parapodia (fig. ig.i.p)
with subequal notopodial and neuropodial postsetal lamellae which are oval in
outline with constricted bases. Setiger g with broad-winged capillaries (fig. ig.i.r)
similar to those of setigers 1-8. Abdominal hooded hooks with two teeth side by
side above the main fang.
Type locality : Agulhas Bank, South Africa.
Records: South West Africa (26/15/s) ; Cape (from 32/18/s to 34/23/s) ; Natal
(29/3 1 /s, d).
Distribution : Endemic.
498
POI.VCHAICTA OF SOUTHERN AKRIOA
Family CIRRATULIDAE Carus, 1863
Body cylindrical with numerous similar segments and tapered at biith ends.
Prostomium small and usually without projections. Proboscis unarmed and not
evaginable. Pcristomial segment triannulate and achactous. A pair of grooved
palps or several grooved tentacular filaments are inserted at the posterior end of the
pcristomium which may extend back above the anterior sctigcrs. Long cylindrical
branchial filaments arise above the notopodia of the first and a variable number of
succeeding setigers. Para podia biramous but the parapodial lobes are not developed
so that the two bundles of setae arise directly from the sides of the body. Sctal types
include simple capillaries, acicular hooks or in a lew cases compound hooks. No
anal cirri.
Records from southern Africa
Caiilleiiiili! ackula Day ....
Caidleriella bioculata (Kefcrstcin)
as Helerocirrus fiavo-viridis St. Joseph
Caulleriella capensii (Monro)
as Heteiocbnis capiit-eiocis var. capensis Monro
as Hticrnciirin capensis Monro
? as Dodecaceiia afra Augener
Chaetoznne setosa Malmgren
(.'irrdtiiliis africanus Gra\'icr
tArratulus chrvsodermn Olaparede
Cirratulus cirratus (Miiller)
Cirratulus concinnus Ehlers
Cirratulus filiformis Keferstein .
Cirratulus gi/ihristi Day ....
Cirratulus sp. .....
Cirriformia capensis (Schmarda)
as Cirratulus capensis Schmarda
asCirratulus australis Stimpson (non Blanchard)
as Cirratulus cirratus (non Miiller'
as Audouinia filiaera \'ar. capensis
as Audouinia australis (Stimpson)
Cirrijormia afer (Ehlers) .
as Cirratulus afer Ehlers
Cirriformia filigera (Delia C'.hiaje)
as Audouinia fdii^cra Dclle Clhiajc
Cirriformia jninclata (Gruhe)
as Audouinia punctata (Grube)
ns Aud(niinia Jiligera var. capensis (p
Cirriformia saxatilis (Grax'ier) .
as Audouinia saxalilii Gravier
f,7Cs, — Ns
26.Ai
51CS
33CS
44Ca, 48GS
26\Vi
51CS
44Ci
44Ci, ?45Pi
15GS
48Ad, 51CS
5 1 Csd
51GS
S'Cs
4Ci, loGi, I iGi,
liGi, i6\Vi, 26\Vis
2Ci
i3Ci
33CS, 35Gi, 36Ci
44Ci
1 f, As
45NiPi
4oNi, 45Pi
36X1
45Pi
CIRRATULIDAE AND HETEROSPIOMDAE
499
Cirriformia tentaculata (Montagu)
as Audouinia tentaculata (Montagu)
as Cirratulus tentaculus Montagu
as Cirratulus atrocollaris Grube
asCirratulus tentaculus var. meridionalis (Marenz.
as Audouinia filigera var. meridionalis (Marenz.)
Dodecaceris capensis Day .
Dodecaceria laddi Hartman
Dodecaceria pulchra Day ....
as Dodecaceria futulicola (non Ehlcrs)
Tharyx annulosus Hartman
Tharyx dorsobranchialis (Kirkegaard)
as Cirratulus dorsobranchialis Kirkegaard .
Tharyx filibranchia Day ....
Tharyx marioni (St. Joseph)
as Tharyx multijilis Moore .
5>C5
4oNi, 44Ci,
45Pi.\i, 48\VsCs
iiWi, i3Ci, 32Ci
laCi
26\Vis
35Ci, 36.\iCi
5iCs
4iCs, 45\i
44Ci
38Ci
— Xs
5iCsd, — Nsd
48,'\sCs
5iCs
5 1 Cs, 56\Vs
48CS
BIOLOGICAL .NOTES
Cirratulids are deposit feeders and gather food particles from the sea bottom by
means of a single pair of grooved palps or numerous grooved tentacular filaments
which appear to arise from the first few setigerous segments. The homologies of these
structures will be described later.
Cirratulids are sluggish worms and commonly bury their bodies just below the
surface of the sea bottom so that only the long gills and tentacular filaments are
visible. These are usually bright red or orange. Cirriformia capensis is common among
closely packed mussels on rocky shores, in the hold-fasts of kelp and among massed
colonies oi Pyura. This species seems to be limited to well oxygenated habitats. Cirri-
formia tentaculata on the other hand is found under stones lying on black mud and
presumably tolerates low oxygen tensions. Cirratulus chrysoderma seems to have similar
habits and is often gregarious.
Dodecaceria is an interesting genus which burrows in thick encrustations of litho-
thamnion, Dodecaceria pulchra being very common on Cape shores at the Patella
cochlear level. If the hthothamnion in a shallow rock pool is examined carefully,
many minute holes will be seen through which Dodecaceria extrudes its palps and gills.
If undisturbed, tiiese spread out around the head like the petals of a tiny black
flower. How the worm burrows in the dense matrix of the coralline is unknown but
the worm appears to do little damage to the plant which grows up around the hole
so that the whole surface is covered with volcano-like knobs.
While Dodecaceria produces eggs and sperm Uke other polychaetes it is also capable
of asexual reproduction by fragmentation. It is said that even a single isolated
segment may regenerate into a small worm.
500 PO[.vc;iiAr.iA or southern ai'rica
THE MAIN" DIAG.XOSrtC CiHARACI'ERS
riic body consists of numerous rounded segments witJiout parapoilial projeeti<ins
but with two bimdlcs of setae and long filiform branchiae. The main diagnostic
features include llic licad and feeding appendages, the number and disposition of the
branchial filaments and the nature of the setae.
The head and foedins, a/ipeiidanes. The prostcunium is a well developed conical lobe,
antero-dorsal to the mouth. The eye-spots are seldom well developed and always
subdermal but their number is important in certain species. Prostomial projections
are absent in all genera except Aerncirrm and Macrochaeta which possess a pair of
fronto-lateral lobes or "palpes frontal". The buccal segment or peristomium is
vaguely triannulate and has been regarded by some workers as representing three
fused and achactous "mctastomial segments". Dissection shows that there is in fact
only one elongated buccal segment. The last peristomial annulus bears either one
pair of large grooved palps or several pairs of smaller tentacular cirri. These
appendages usually originate close to the intersegmental constriction which separates
the last peristomial annulus from the first sctiger. In the genera Cirraliilm and
Cirriformia however, the last part of the peristome bearing the tentacular filaments
may extend back over the first few setigerous segments so that the tentacular fila-
ments seem to arise from the dorsal surface of these seginents ; in Cirriformia tenta-
culata for example the tentacular filaments arise from the dorsal surface above setigers
6 trj 7. Dissection, however, shows that the tentacular filaments are truly peristomial
in origin.
In several species the distinction between the tentacular filaments and the branch-
ial filaments of the setigerous segments is not immediately obvious. However the
tentacular filaments are always slightly stouter and more dorsal in origin and careful
examinadon will show that they are always grooved whereas the branchial filaments
are cylindrical ; finally the tentacular filaments have a single blood vessel while
the branchial filaments have two.
The arrannemeni nj the hranehial filaments. Apart from the genus Timarele which possesses
several brancliial filaments per segment there is only a single branchial filament
per parapodium. In the species of Dodeeaeeria branchial filaments are restricted to
the first few setigers, in CaiiUerieUa there are more and in Cirraliilu':, branchial fila-
ments usually extend to the posterior end. Typically, the branchial filaments arise
just above the notopodial setae but in some species the branchiae of middle segments
arise further above the notosetae than the distance which separates the notosetae
and neurosetae. It is emphasised that this applies to the middle segments for the
branchiae of the first few segments always arise close to the notosetae. The extreme
cases occar in Tharr\ dor\nhranchiali^ where the branchiae of the middle segments
arise liTjni the mid-dnrsal line and Tharrx jdahram'hia where they arise between the
notosetae and neurosetae.
Setae. Compound acicular setae occur only in the neuropodia of Acrocirriis and
.Macrochaeta. In other genera the setae are all simple and are either acicular or
capillary or both. The shape of the acicular hooks v.uies iVom simple spines to
CIRRATULIDAE AND HETEROSPIOMDAE 501
forms with excavated spoon-shaped ends or even bidentatc hooks approaching those
of the Spionidac. The capillaries are usually somewhat flattened distally with one
margin very finely spinulose or hispid. In Tharyx these capillaries are the only type
of setae ; in other genera, however, where both capillaries and acicular hooks
occur, the capillaries tend to be more numerous in the notopodia and in the anterior
segments while the hooks are more numerous in the ncuropodia and in the posterior
segments. Thus two or three hooks may appear in the middle notopodia and six
or seven hooks in the anterior neuropodia. The exact segment in which hooks
first appear seems to change with the size of the individual so that only major
differences in the distribution of hooks and capillaries are of systematic importance.
SUBDI\lSIO.\S OF THE FAMILY
The family CirratuUdae has been used as a dumping ground for a variety of
genera. Streblospio has now been referred to the Spionidae and other aberrant
genera have been placed in the new families Hetcrospionidae and Cossuridae. The
genera that remain may be grouped into two sub-families. All the large and common
genera with branchiae are included in the CirratuUnae and three small genera
without branchiae are included in the Ctenodrihnae. No member of the latter has
been reported from southern Africa.
Key to Subfamilies and Gener.\
1 Branchial filaments absent ....... Subfamily Ctenodrili.n.ae*
- Branchial filaments present (subfamily Cirr.\tulin.\e) ...... 2
2 Prostomium with fronto-lateral lobes. Compound hooks in the neuropodia ... 3
- Prostomium without projections. Compound setae absent ...... 4
3 One to two pairs of branchiae on the first metastomial segment . . . ACROCIRRUS*
- Branchiae absent from the first metastomial segment . . . MACROCHAETA*
4 More than one pair of branchial filaments on each parapodium . . TIMARETE*
- One pair of branchial filaments on each parapodium ...... 5
5 Two large grooved palps (fig. 20. i .g) ........ . 6
- Several grooved tentacular filaments (fig. 20. 3. 1) ....... 9
6 Seven or less pairs of gills (fig. 20. 1 .c) ..... DODECACERIA {p. ^o\)
- 12 or more pairs of gills (fig. 20.1.1) ......... 7
7 Capillary setae only present ........ THARYX (p. 504)
- .Acicular hooks and capillaries present ......... 8
8 Acicular setae at the posterior end of the body form a continuous dorso-ventral arc of
spines (fig. 20. 1. n) CHAETOZONE ip. 509)
- Setae remain in two distinct bundles to the posterior end of the body
CAULLERIELLA (p. 506)
9 The first branchiae appear on the same segment as the tentacular filaments :fig. 20. 3. g)
CIRRATULUS (p. 510)
- The first branchiae appear anterior to the tentacular filaments (fig. 20.4.3)
CIRRIFORMIA (p. 514)
DODECACERIA Oersted, 1843
Small worms commonly found burrowing in calcareous encrusdng algae. Pro-
stomium blunt and forming a hood over the mouth. Buccal segment long and
achactous but bearing two stout grooved palps at its junction with setiger i. Two
502 POI.VtlHAF.TA OF SOUTHERN AFRICA
to eiglit pairs of branchial filaniouts on the buccal segment and the first few setigers.
Setae are all simple and include capillaries and stout acicular hooks with spoon-
shaped ends.
Type SPECIKS : Dodccaceria conchamm Oersted, 181.3.
Key to Species
1 Two pairs of gills, the first on the buccal segment and the second on the first sctigcr
(fig. 20.1. g) D.laddi
- Four to five pairs of gills ........... 2
2 Gills arranged in an inverted V with the first pair close together on the buccal segment.
Setiger i with notosetae (fig. 2o.!.c) ....... D. pulchra
- Gills not in an inverted V. The first two pairs on the buccal segment, one above and one
below the palps. Setiger i without notosetae (fig. 20. i.j) . . . D. capensis
Dodecaceria laddi Hartman, 1954
(fig. 20.1.g-i)
Dodiccueria Imldi Hartm^in, i<)-,{: 638, figs lybCand 177D-H; Day. I9'i7: 102.
A small species about 7 mm. long. Palps long and lateral in origin (fig. 20. 1. g).
Two pairs of gills both smaller than the palps, the first pair arising dorso-latcrally
on the posterior margin of the buccal segment and the second, more dorsal pair on
the first setiger. Capillaries in both rami except in the neuropodia of middle
segments. Hooks in both rami from setiger 1 1-15 onwards, the anterior ones (fig.
20. 1. h) with slightly flattened tips, later ones spoon-shaped and posterior neuro-
podial ones (fig. 20.i.i) with a boss below the excavation.
Type locality : North Marshall Islands.
Records: Cape (34/1 8,'s ; 34/22/s).
Distribution': Marshall Is. (Pacific).
Dodecaceria pulchra Day, 1955
(fig. 20.i.a-f)
Dodecaceria pulchra Day, 19^,5 : .jiS, fig. 2 h-m.
Black gregarious worms up to 20 mm. long boring in encrusting calcareous algae
(fig. 20. 1. a). Prostomium (fig. 20.1.C) a stout hood overhanging the mouth. Palps
lateral in origin. Four pairs of gills which are longer and more slender than the
palps, the first pair arising from the dorsal surface of the buccal segment above the
palps, the second pair wider apart and on setiger 1, and the third and fourth pairs
still wider apart and on setigers 2 and 3 so that the whole group forms an inverted V.
.Saw-edged t ,i])iihiiies (fig. 2o.i.c) present in the notopodia of setigers 1-12, then
CIRRATULIDAE AND HETEROSPIONIDAE
503
Fig. 20.1. Dodecaceria jmUhra. (a) Coralline alga with burrows of Dodecaceria. (b) Entire
worm (three times natural size), (c) Head, (d) Posterior foot, (e) Capillary seta.
(f) Spoon-shaped hook. Dodecaceria laddi. (g) Head, (h) Anterior spoon-shaped hook.
(i) Posterior hook. Dodecaceria capemis. (j) Head, (k) Hook. Chaelozone selosa. (l)
Anterior end. (m) Middle foot, (n) Posterior end showing spines and pygidium. (o)
Anterior hook, (p) Posterior spine.
504 I'OI.VC'.HAKTA OF SOUTHERN AFRKIA
often missing from middle segments only to reappear and continue to the posterior
end (fig. 20. 1. d). Capillaries absent from the neuropodia after the first 12. Spoon-
shaped hooks (fig. 20. 1. f) present in the neinopodia from setiger 10 and in the
notopodia from setiger 12. Posterior hooks with a boss below the excavation.
Type localmv : False Bay, South Africa.
Records; South West Africa (26/15/!); Cape (from 29/16/i to 32/28/i) in
encrusting coralline algae.
Distribution : Endemic.
Dodecaceria capetisis Day, 1961
(figl. 20.1.j-k)
Dodecaceria cajiensi^ Day, 1961 : 504, fit;. 7 j-k.
Gregarious \vorms boring in calcareous algae. I^Migth up to 15 mm. for 60
segments. Prostoniium (fig. 20.1.J) a stout hood o\erhanging the elongate mouth.
Palps short and arise laterally just in front of setiger i . Four pairs of gills longer than
the palps ; the first two pairs arise immediately above and immediately below the
palps, the third pair on the dorso-lateral surface of setiger i and the fourth pair on
the sides of setiger 2. Setiger i with ncuiosetae only. Notosetae arc all capillaries
up to setiger 12 where they are replaced h\ acicular hooks for about 10 segments ;
later notopodia with both hooks and ca]5illaries. Neurosetae arc all capillaries for
the first seven segments, then both capillaries and hooks from setiger 8 to 11, then
hooks only for the next few segments and then capillaries reappear again and both
types of setae continue to the posterior end. The acicular hooks (fig. 20.i.k) have
the usual spoi in-shaped ends.
Type locality : Dredged in False Bay, South Africa.
Rfx;ords : South ^\'esl Africa (26/15/s) ; C^ape (34/18/s).
Distribution ; Endemic.
THARYXWeh^ie-y and Benedict, 1R87
Body elongate with numerous segments. Prostomium conical. Buccal segment
elongate and achaetous, with a pair of long grooved palps arising at its junction with
setiger i. Branchiae from setiger i at least to the middle of the body. Hooks
absent, the setae being all capillaries.
1'ype species: Iharjx aculm Webster and Benedict.
Kl Y TO SpECItS
1 Setal blades short and ccjarsily toothed ....... T. aiinulosiis
- Setal blades long and finely spinulosc to smooth ....... 2
2 Eyes present. Posterior branchLil filaments arise between the notosetae and neurosetae
(fig. 20. 2. b) ........... T. filibranchia
- Eyes absent. Branchial filaments arise abo\e the notosetae tlirouglioiit ... ^j
CIRRATULIDAE AND HETEROSPIONIDAE 505
3 Branchial filamenls in the middle of the body arise laterally just above the notosetae (fig.
20. 2. e) ............ T. marioni
- Branchial filaments in the middle of the body arise from the mid-dorsal line (fig. 20. 2. g)
T, dorsobratichiata
Tharyx filibranchia Day, 1961
(fig. 20.2. a-d)
Tharyx filibranchia Day, 1961 : 503, fig. 7 e-h.
Body elongate, up to 20 mm. in length with 180 very short segments and filamen-
tous branchiae. Prostomium (fig. 20. 2. a) pointed and with a single pair of subdcrmal
eyes. A pair of large grooved palps arises from the junction of the buccal segment
and the first setiger. Very fine branchial filaments from setiger i to near the posterior
end. At first they arise above the notosetae but later from between notosetae and
neurosetae (fig. 20. 2. b). Capillaries (fig. 20. 2. d) present in both rami throughout,
but in posterior neuropodia there are pointed setae (fig. 20.2.0) intermediate between
capillaries and acicular setae.
Type locality : False Bay, South Africa.
Records : (Cape 34/18/s and 34/23/s, d) ; Natal (30/30/s, 29/31/s, d).
Distribution : Only three records.
Tharyx marioni (Saint-Joseph, 1894)
(fig. 20. 2. c)
Heterocirrus marioni Saint-Joseph, 1894: 56, pi. 3 figs. 62-64.
Tharyx marioni: Fauvel, 1927 : 100, fig. 35 a-b ; Day, 1961 : 503.
Body up to 100 mm. long by i mm. but swollen anteriorly and tapered at both
ends. Prostomium broadly conical and without eyes. Buccal segment triannulate
with the third annulus distinct and bearing a pair of very stout grooved palps at the
junction with setiger i. Anterior segments short, about 12 times as broad as long
but posterior ones longer, only two to three times as broad as long. Long branchial
filaments from setiger i to near the end of the body. They arise immediately above
the notosetae throughout (fig. 20. 2. e). Notosetae and neurosetae are all smootJi
capillaries, the notosetae being longer than the neurosetae and in the posterior region
some are as long as the body is broad.
Type locality : France.
Records: South West Africa (26/14/d and 28/16/s) ; Cape (33/1 8/s and 34/21 /s) ;
Natal (29/3 1 /d).
DisTRiBUTio.N : N. Atlantic from North Carolina (s) and Scotland (s, d) via
Enghsh Channel (i, s) to Morocco ; Mediterranean (s). Also recorded as T. rmdtifilis
from Persian Gulf (s) and Madras (i, s, d).
E
506 rOLVCIIAr.TA or SOUTHERX AFRUIA
Tharyx dorsobranchialis (Kirkigaard, if)-)9)
(fig. ■-•o.-j.r-h)
Cinatulus iiorsobrdncfiialis Kirkcgaard, 1959: 34, figs. 2-3.
Tharyx doTsobramhiaUs : Day, 1 9(3 1 : 501.
A thread-like species about 35 mm. long (fig. 20.2.f). Prostomium sharply conical.
No eyes. Buccal segment expanded and triannulatc, with two large caducous
grooved palps arising from the junction of the third annulus and sctigcr i. Very
slender branchial filaments on all setigers from the first onwards ; a fusiform inter-
branchial dorsal area extends from the first sctiger to about the twentieth and
behind this the branchiae arise from the mid-dorsal line (fig. 20. 2. g). .\nterior
segments short, posterior ones about as broad as long. Setae are all saw-edged
capillaries (fig. 20. 2. h) ; acicular hooks entirely absent.
Type locality : Dredged in 20 fth. ofl' Angola.
Records: South "West Africa (26,15/s); Cape (from 32/17/d and 34'18/s to
33/25 s; Natal (2g'3i/s, d).
Distribution: Angola (s).
Tharyx annulostis Hartman, 1965
TJtttryx annuloitis Hartman, 196;,: 167, p\. 34.
Body up to 18 mm. long, cylindrical anteriorly but with marked segmental
constrictions giving a beaded effect in tlie middle of the body and the final segments
arc short and somewhat flattened. Prostomium bluntly conical and without eyes ;
buccal segment elongate without annuli. A pair of grooved palps mounted dorso-
laterally at the junction of the buccal segment and the first setiger. Branchial
filaments inserted immediately above the notosetae. The first few pairs are almost as
long as the palps but later ones are progressively slinrter and they disappear before
the middle of the body. The setae are all serrated capillaries ; some of the anterior
ones are fairly long but the rest are short and about a third the segmental width.
They have thick, expanded blades which taper abruptly to fihform tips and under
high power the blade is seen to have well marked teeth like the edge of a saw.
T^PE LOCALITY : Continental slope oft Massachusetts.
Records: Natal (29/31/s).
Distribution : Massachusetts (d, vd, a) to N.E. South ,\merica (vd, a).
CAVLLERIELLA Chambcrlin, 1919
Body \ermilnrm and rounded in section. Prostomium conical. Buccal segment
elongate and triannulatc. A pair of large grooved palps and the first pair of branchial
filaments at the junction of the third annulus of the buccal segment and setiger i.
Brandiiai- from setiger i .md extend o\er the first half 1 if the body. Nolopodi.d and
CIRRATULIDAE AND HETEROSPIOMDAE 507
neuropodial setae not widely separated but never form a continuous arc. Capillary
setae and hooks both present in some part of the body.
Type species: Cirratulus bioculalus Keferstein, 1862.
Key to Species
1 Acicular hooks unidentate. Capillaries and hooks present in both rami of posterior feet
C capensis
- Acicular hooks bidentate. Capillaries may be absent from posterior neuropodia . . 2
2 Acicular hooks with narrow dorsal hood (fig. 20. 2. p). (Posterior neuropodia without
capillaries) ............ C. acicula
- Acicular hooks without hoods .......... 3
3 Middle and posterior neuropodia with capillaries. Notopodial hooks absent C. zetlandica\
- Middle and posterior neuropodia without capillaries. Hooks present in the notopodia
from setiger 6-9 onwards ......... C, bioculatus
Caulleriella capensis (Monro, 1930)
(fig. 20.2.i-l)
Heterocirrus capulesocis var. capensis Monro, 1930 : 156, fig. 61.
Body yellow, rather swollen, about 20-30 mm. long, with rather stout gills and
rather long segments. Prostomium (fig. 20. 2. i) short, conical and depressed with
a pair of dark ocular bars formed of three to four eyes across the posterior margin.
Buccal segment swollen, its third annulus bearing a pair of stout palps much thicker
than the branchial filaments. Branchial filaments restricted to the anterior part of
the body, seldom more than 15 pairs, each filament arising close above the notosetae
(fig. 20.2.J). Parapodial tori prominent anteriorly. Capillary setae with finely
spinulose margins (fig. 20. 2. k) and present in both rami tliroughout. Unidentate
acicular hooks (fig. 20. 2. 1) from setiger 15-20 onwards.
Type locality : Simonstown, South Africa.
Records: ? South West Africa (26/15/s) ; Cape (from 33/18/s and 34/18/i to
34/22/i, s and 34/25/s).
Distribution : ? Endemic.
Caulleriella acicula Day, 1 961
(fig. 20.2.m-p)
Caulleriella acicula Day, 1961 : 501, fig. 7 a-d.
Body up to 20 mm. long with 200 segments. Prostomium (fig. 20. 2. m) a sharply
pointed cone with a pair of small subdermal eyes. Buccal segment swollen with
a pair of palps at its junction with setiger i. Body segments short, rather flattened,
with prominent parapodial ridges on either side. Slender branchial filaments
arise just above the notosetae (fig. 20.2. n) and extend at least to the middle of the
body. Parapodial rami well separated. Very few capillaries (fig. 20.2.0) in the
tA doubtlul record for South .-V^rica.
5o8
I'OIAtiHAl. lA OF SOUrm'.RN A1'RK:A
Fi'i. 20.2. lliarw jdibramhia. ix) .Vntcrior end. (b) Middle segnienl. (c:) Intermediate
form of seta. (D) Clapillary .seta. Tharvx warinni. (E) Three-quarter view of three middle
set^ments. Thiirvx (lnTSobram}uaUs. (f) Entire worm fthrce times natural size), (g) Middle
.segment. (n| Clapillary seta. Caullmdla cafienui. Hi Anterior end. (j) Middle segment.
(k) Capillary seta, (l) .Sigmoid hook. CauHcriilla aciciila. (m) Anterior end. (n) Middle
.segment, (o) Capillary seta, (p) Bidcntate hook. Caidltiielta biocidalus (after Fauvel;.
(q) Bidentale hook.
CIRRATULIDAE AND HETEROSPIOXIDAE 509
notopodia and none in the neuropodia. Acicular hooks in both rami throughout.
Individual hooks (fig. 20. 2. p) bidentatc with a small secondary tooth and a narrow
hood on the convex side of the shaft.
Type locality : Dredged in False Bay, South Africa.
Records: Cape (from 32/18/s to 34/22/s) ; Natal (29/31/s).
Distribution : Endemic.
Caulleriella zetlandica (Mcintosh, 191 1)
Chaetozone zellandka Mcintosh, 191 1 : 16; Southern , 1914; 115, pi. 12 and 13 figs 29 a-k.
Heterocirnis zetlandica : Fauvel, 1927 : 99, fig. 34 i-n.
Body thread-like, up to 25 mm. long, pale in alcohol. Prostomium a long cone
without visible eyes. Peristome stout and triannulate. A pair of stout grooved palps
at the junction of the peristome and setiger i. Body rounded in section with short,
crowded segments. Branchial filaments arise immediately above the notosetae, are
always slender and become shorter posteriorly. Notosetae and neurosetae close
together. Notosetae are all capillaries, some longer with more slender blades, some
shorter with broader, curved blades. Neurosetae include short broad-bladed
capillaries similar to those in the notopodia plus two to four stout sigmoid hooks.
In juveniles the hooks are obviously bidcntate but in adults the teeth are very blunt
and the end of the hook is almost truncate. The hooks have no hoods.
Type locality : Ireland.
Records : ? ? South Africa.
Caulleriella bioculatus (Keferstein, 1862)
(fig. 20.2.q)
Cirraluliu bioculatus Keferstein, 1862; 121, pi. 10 figs. 23-27.
Heterocirrus bioculatus: Fauvel, 1927 : 96, fig. 33 i.
Body up to 40 mm. long with 140 segments. Prostomium pointed and with two
eye-spots. A pair of long grooved palps inserted at the junction of the buccal
segment and setiger i. Branchial filaments from setiger i to the middle of the body.
Notopodia with capillaries throughout and one to three acicular hooks from setiger
six to nine onwards. Neuropodia with capillaries in setigers i and 2 but only hooks
thereafter. Hooks (fig. 20. 2. q) bidentatc with a minute secondary tooth but without
hoods.
Type locality : France.
Records : Not recorded from southern Africa.
Distribution: English Channel (s) ; ? .\ngola (1).
CHAETOZONE Malmgren, 1867
Body elongate, rounded in section and tapered at both ends. Prostomium
conical. Buccal segment triannulate. A pair of elongate, grooved palps at the
5IO roLvcuiAErA OF sourm.RX afrk;a
junction between the third annulus and scliger i. Branchial filaments on numerous
segments. Setae include slender capillaries and simple acicular hooks ; the latter
form a continuous dorso-ventral arc on posterior segments.
Type SPECIES : C'kaiiozone stiusa Malmgrcn, 1867.
Chaetozone setosa Malmgrcn, 1867
(fig. 20.1.1-p)
Chaetozone setosa Malmgrcn, 1867 : 1206, pi. 13 fig. 4 ; Fau\cl, iO-7 : loi » '~ig- 33 d-k.
An elongate worm about 20-25 rnm. in length with 70-90 segments. Prostomium
(fig. 20.1.1) conical. No eyes. Two stout palps at the anterior margin of setiger i.
Branchial filaments from setiger i to the middle of the body ; they arise close above
the notosetae. Capillary setae from the first setiger to the posterior end. Notosetae
up to four times the segmental width. Simple sigmoid acicular hooks (fig. 20.1.0)
appear in the notopodia from setiger 3 onwards and in the ncuropodia from setiger i .
The notopodial and neuropodial setae are separate bundles over most of the body
(fig. 20. 1. m) but form a condnuous dorso-\cntral arc of spines (fig. 20.i.p) in
posterior segments (fig. 2o.i.n). Pygidium with a dorsal anus.
Type loc.\lity : Sweden.
Records : Cape (34*22 s and 34'23/s).
I)iSTRiBi_TiON : .\rctic ; Greenland (s, d) ; North Clarolina (s) ; Sweden id) and
Scotland (s) to Morocco (s, d) ; and tropical West Africa to Angola (d) ; Mediter-
ranean fs) ; Aden (s) : subantarctic (Heard Is. (s) ) ; N. Pacific from Behring Sea
and Japan to C^alifornia (i, s).
CIRR ATULUS LAmavck, 1801
Body elongate, rounded in section and tapered at both ends. Prostomium
conical. Buccal segment triannulaic. A few grooved tentacular filaments above
one of the anterior setigers. Branchial filaments appear on the same segment as the
tentacular filaments and extend over the greater part of the body. Setae are slender
capillaries usually with acicular hooks as well.
Type species : Lumbiicus cinalm }^lu\lcv, 1776.
Key to Species
1 .Xcicular hooks absent, only capillary setae present e\'en in i^osterior segments . . 2
- Both acicular hooks and capillaries present --------- 'i^
2 'Fentacular filaments arise above setigers 4-7 (fig. 20.3.3) . . C chrysodernia 'p. 311)
- 'tentacular filaments arise at the anterior margin of setiger I (fig. 20.3. e) C. filifortnis (p. 511)
3 Tentacular filaments arise on the anterior margin of setiger i. Prostomitim with or
without eyes ............. 4
- Tentacular filaments arise above setigers 2-3. Prostomium without eyes (fig. 20. 3. g)
C. africanus (p. 311)
4 .Middle and posterior neuropodia \vith hooks only. Two to four pairs of evis
C. gilchristi (p. 513)
- C'apillaries present in both rami throughout. Eyes present or absent .... 5
CIRR.'KTULIDAE AXD HETEROSPIONIDAE 511
5 Aclcular hooks present in both rami after setiger 12. Two to four pairs of eyes
C. cirratus (p. 513)
— Acicular hooks absent from the notopodia. Eyes absent . . . C. concinnus (p. 514)
Cirratulus chrysoderma CHaparede, 1868*
(fig. 20.3.a-d)
Cirratulus chrysoderma Claparede, 1868 : 262, pi. 23 fig. 4 ; Fauvel, 1927 : 95.
Body (fig. 20. 3. b) slender, up to 25 mm. in length, greenish brown in colour.
Segments much broader than long. Prostomium (fig. 20. 3. a) bluntly conical with
two indistinct eyes. Two to four pairs of tentacular filaments above setigers 4-7.
Branchial filaments start on the same segment and extend over the anterior half of
the body ; they are fairly stout and arise closer to the notosetae than the distance
between notosetae and neurosctae (fig. 20.3.0). Acicular setae absent and only
long capillaries (fig. 20.3. d) in both rami of the parapodia.
Type locality : Italy.
Records: Cape (from 29/16,1 and 34/18/i to 34/21/i).
Distribution : Mediterranean (s) ; Gulf of Manaar ; India ; Malaya ; Japan.
Cirratulus filiforntis Kcferstein, 1862
(fig. 20.3.e-f)
Cirratulus Jiliformis Keferstein, 1862: 122, pi. 10 fig. 28-31 ; Fauvel, 1927: 94, fig. 33 h; Day,
1961 : 499.
Body very slender, up to 50 mm. long and greenish brown when alive. Prostomium
(fig. 20. 3. e) pointed and without eyes. Buccal segment swollen. About four tenta-
cular filaments arise at the junction of the buccal segment with setiger i. Branchial
filaments from sedger i to near the posterior end and arise just above the notosetae
throughout (fig. 20. 3. f). Parapodial ridges well marked. Setae are slender flattened
capillaries throughout, hooks being entirely absent.
Type locality : W'estern Europe.
Records: South West Africa (23/14/d and 26/15/s) ; Cape (from 34/18/s to
35/20/d).
Distribution : Eastern Atlantic ; ? Sweden (d, vd), Scotland (s) to Morocco (s) ;
Senegal (s) ; ? tropical western Africa (s) ; ? South Georgia; Persian Gulf (s).
Cirratulus africanus Gravier, 1906
(fig. 20.3.g-k)
Cirratulus a/ricanus Gravier, 1906c; 152, pi i fig. 179, te.xt-figs. 315-317 ; Day, 1957: 102.
Body about 50 mm. long, brown in colour. Prostomium (fig. 20. 3. g) long,
pointed and without eyes. Three to four tentacular filaments arise above setigers
2-3. Branchiae from setiger 3 and extend over most of the body, arising close above
* Imajima and Hartman ( 1964:- have transferred this species to Cirri/ormia on the basis that the tentacular
cirri do not arise from setiger I. .As shown earlier, the cirri really originate from the peristomium.
51-
POLVc:iiAi.r.\ or southern afrk:a
I'l*.. jri.3. Cinattfliis (hrvwdernia. (a) Anterior end. (n) Entire worm (Tour times natural size).
iv.) Middle foot, (u) Clapillar\'seta. Cirratuhis filifomii^. (e) .Xnterior end. iv) Middle foot.
Chratulm africanus. (o) Anterior end. (h) Middle foot. (i) Capillary seta. (j) Abruptly
tapered seta, (k) Sigmoid hook. Cirratulus gilchrisd. (l) Anterior end. (m) Middle foot.
CiTTiilulus cirralus (after Fauvcl). (n) Middle foot. Cirralulus cominmis {after Elders).
(f)) Lateral view of anterior end.
CIRRATULIDAE AND HETEROSPIONIDAE 513
the notosetae (fig. 20.3.11). Capillary setae (fig. 20.3.!) in both rami of the parapodia
throughout. From the tenth foot setae appear which taper abruptly (fig. 20. 3.]')
and are thus intermediate between capillaries and acicular hooks. They change
to normal sigmoid hooks (fig. 20. 3. k) about the middle of the body.
Type locality: Djibouti, Gulf of Aden.
Records: Mocambique (26/32/i).
Distribution: Gulf of Aden.
C-rratulus gilchristi Day, 196 1
(fig. 20.3.1-m)
Cirraliilits gilchrisli Day, 1961 : 500, fig. 6 j-m.
Body thread-like, only 12 mm. long and markedly tapered. Prostomium (fig.
20. 3. 1) bluntly rounded with four pairs of eyes (or only one to two in juveniles).
Three to five pairs of stout tentacular filaments arise from the anterior margin of
setiger i in a transverse row. Long branchiae from setigcr i to the posterior end.
In the middle of the body (fig. 20. 3. m) each branchial filament arises as far above
the notosetae as the distance between notosetae and neurosetae. Unidentate acicular
hooks in both rami of the parapodia from setiger 3-6 onwards. Capillaries present
in all notopodia but absent from the neuropodia from setiger 6 onwards.
Type locality : Saldanha Bay, South Africa.
Records: Cape (33/17/s, 34/18/i, s).
Distribution : Three records only.
Cirratulus cirratus (MUller, 1776)
(fig. 20. 3. n)
Lumbridis ciT'atus 'WuMer^ 1776: 215.
Cirratuhis cirratus: Fauvel, 1927 : 94, fig. 33 a-g.
Body elongate and cylindrical, up to 50 mm. long and 1-2 mm. in diameter.
Prostomium bluntly conical with two to four pairs of eyes in a row. Peristome as
long as the first three to four segments. Numerous tentacular filaments form a row-
above setiger i. Segments two or three times as broad as long. Stout branchial
filaments from setiger i to the end of the body ; on middle segments (fig. 20.3. n)
they arise higher above the notosetae than the distance between notosetae and
neurosetae. Capillary setae in both rami of the parapodia throughout. One to two
sigmoid acicular setae in the notopodia from setiger 20 onwards and two to four in
the neuropodia from about setiger 12 onwards.
Type locality : Danish seas.
Records : No certain southern African record.
Distribution : Arctic (s) ; North Atlantic from the North Sea to the English
Channel ; subantarctic from Magellan area and the Falkland Islands to Kerguelen.
514 POI,VC:iIAI/rA OF SOUTHERN AFRICA
Cirratulus concinnus Ehlcis, 1908
(fig. 20.3.0)
Cirratulus tmuinniis. VM\eis, 1908: 128, pi. 17 figs. 13-14.
A small tliicad-like worm about 25 mm. long for 80 segments. Prostomium (fig.
20.3.0) an elongate cone without eyes. Buccal segment elongated with one to two
pairs of grooved tentacular filaments arising at the junction of the [leristome with
setiger i. Branchial filaments from the first setiger onwards. In the middle of the
body individual branchiae arise ( ?) close above the notosctac. Long capillaries in
both rami of the parapodia throughout. Acicular hooks absent from the notopodia
but present in the ncuropodia from setiger 32 onwards.
Type locality : Agulhas Bank, South Africa.
Records: Clape (35/'23/d).
Distribution : Only one record.
CIRRIFORMIA Hartman, 1936
Prostomium conical and usually without eyes. Buccal segment triannulate.
Numerous grooved tentacular filaments arise in a group from an extension of the
peristome above sctigcrs 2-7. Branchial filaments start on setiger i and are present
as single filaments on all subsequent segments over the greater part of the body.
Rami of parapodia well separated and bear both capillary setae and simple
acicular hooks.
Type sPEtTES : Teiehella tciitaculnta Mont:ii^u, 1808
Key to Species
1 Brancliiac of middle scgnicnls arise closer above the notoselae than distance between
notosetae and ncurosetae (fig. 20.4, b) ......... 2
- Branchiae of middle segments arise farther above the notosetae than distance l)etween
notosetae and ncurosetae (fig. 20. 4. j) ......... 5
2 Tentacular filaments arise above setiger 5-G. Branchiae slender and thread-like (fig. 20.
4-a) .:•.•■•• 3
- Tentacular filaments arise above setigers 2-4. Branchiae fairly stout (fig. 20.4. c) . . 4
3 Capillary setae in both rami throughout the body . . . . C. tentaculata (p. ■,!",)
- No capillaries in the ncuropodia after the 45th setiger .... C. tyiaryae*
4 Hooks sigmoid and present fniin ihe eighth neuropodium and from the 40th notopodium
C. saxatilis (p. ',15)
- Hooks straight ; ihey are absent from anterior segments .... C a/t'r (p. 515)
5 Body a uniform browii ; gills and tentacles uniform yellow or orange .... 6
- Bfjdy speckled with black; gills and tentacles barred (fig. 20.4. k) . C. punctata (p. -^ly)
6 Tentacular filaments arise above setiger 3 ..... C. capensis {p. -,17)
- Tentacular filaments arise above setiger 4-5 ..... C. Jiligera {p. -,i8)
CIRRATULIDAE AND HETEROSPIONIDAE 515
Cirriformia tentaculata (Montagu, 1808)
(fig. 20.4.a-d)
Terebella tentaculala Montagu, 1808 : 110.
Audouinia tentaculata : Fauvel, 1927 : 91, fig. 32 a-g.
Length up to 200 mm. in European forms but never more than 80 mm. in South
Africa. Prostomium pointed (fig. 20.4.3). Numerous tcntacuhir filaments arise in
two clusters above setigers 5-7. Slender branchial filaments from setiger i to the
end of the body. They arise close above the notosetae (fig. 20.4. b). Segments
narrow and crowded. Capillary setae (fig. 20.4.d) in both rami of the parapodia
throughout. Four to five unidentate acicular hooks (fig. 20. 4. c) per ramus after
about the first third of the body (i.e. about segment 50 onwards). Body uniformly
brown, gills red in life.
Type locality : South Devon, England.
Records: South West Africa (22/14/1, 26/15/i, s and 29/16/1); Cape (from
20/17/i to 34/23/e, i, s and 33/27/i) ; Natal (from 31/29/1, s to 27/32/i) ; Mocam-
bique (from 26/32/i to 23/35/e, s).
Distribution : Eastern Atlantic from the North Sea (i) to the English Channel
(i, e) and south to Morocco (i, s) and tropical western Africa (s) ; Persian Gulf (s) ;
tropical Indian Ocean (i, s) ; Pacific from Japan and New Caledonia to New Zealand
(d) and Campbell Is. (i).
Cirriformia saxatilis (Gravier, 1906)
(fig. 20.4.e-g)
Audouinia saxatilis Gravier, 1906c : 154, pi. i figs. 180-182 text-fig. 318.
Body about 50 mm. long, yellowish brown with the anterior segments darker
ventrally. Prostomium (fig. 20. 4. e) rounded. Tentacular filaments arise above
setigers 2-4. Branchiae from setiger i to the posterior half of the body ; they arise
immediately above the notosetae (fig. 20. 4. f). Saw-edged capillaries are present in
both rami of all parapodia. Sigmoid hooks (fig. 20.4.g) first appear in the neuro-
podium of setiger 8 and in the notopodium of setiger 42.
Type locality: Djibouti, Gulf of Aden.
Records: Mocambique (23/35/s).
Distribution: Gulf of Aden (s).
Cirriformia afer (Ehlers, 1908)
(fig. 20.4.h-i)
Cirratulus afer Ehlers, 1908 : 127, pi. 17 figs. 10-12.
[Non] Cirratulus afer Monro, 1930 : 156, fig. 60 (= Timarete sp).
Body up to 20 mm. long, fairly stout with the parapodia forming ridges at the
sides. Prostomium (fig. 20. 4. h) broadly rounded and without eyes. Tentacular
filaments arise above setiger 2 and number five to six on each side. Branchiae from
setiger i and extend to the posterior half of the body ; they arise close above the
SI 6
P()i.vt:piAr,TA OF s(u riir.RX airica
Fic;. 21). 4. Cirriformia lenlaculntn. (a) Anterior end. (b) Fiiot from mid-region, (c) Hook.
'd) C^apillary sela. Cmiformiu saxatiln. (il .Anterior end. (r) Foot from mid-region.
(i,i Hook. Cirriformia afer (after Ehlers). (11) Head end. (i) Foot from mid-region.
Cirriformia jmmlata. (j) Foot from mid-region, (k) Head end. (l) Hook, (m) t^ajiillary
seta. Cirriformia capmsis. (n) Head enfl. (o) Font from mid-region. Ctrrijinniia Jiligcra.
(p) Head end. (q) Foot from mid-region.
CIRRATULIDAE AND HETEROSPIO.MDAE 517
notosetae (fig. 20.4.!). Capillaries in both rami of all parapodia. No hooks anteriorly
but in some of the posterior segments a single spine may accompany the capillaries.
Type locality : Great Fish Bay, Angola.
Records : Not recorded from Sou th Africa.
Distribution: Angola (s).
Cirriformia pujtctata (Grube, 1859)
(fig. 2o.4.j-m)
Cirratuhis punctaltii Grxihe, 1859: 107.
Audouiniti piDKtnla: Day, 1951 : 47.
Audoidnia stmkincta Ehlers, 19053; 292; Fauvel, 1953: 330, non fig. 1 74CI.
Body about 40 mm. long and fairly broad. It is brown flecked ^vith black pigment
and the tentacles and branchiae are barred. Prostomium (fig. 20. 4. k) rounded in
front without obvious eyes. Numerous tentacular filaments arose about setiger 4.
Branchiae from setiger i to near the posterior end. In the middle of the body
(fig. 20.4.J) they arise further above the notosetae than the distance between noto-
setae and neurosetae. Capillaries (fig. 20. 4. m) in both rami of all parapodia.
Slender sigmoid hooks (fig. 20.4.I) appear about setiger 12.
Type locality : West Indies.
Records: Natal (31/29/i to 27/32/1) ; Mocambique (26/32/i).
Distribution: Circumtropical (i, s).
Cirriformia capensis (Schmarda, 1861)
(fig. 20.4.n-o)
Cinaiulm capensis Schmarda 1861 : 56, pi. 27 fig. 213.
Cirratulm mislratis Stimpson, 1856: 391. [Preoccupied by Cinalulus auslralis Blanchard, 1849 =
Cirriformia jiligera (Delle Chiaje, 1828).]
Audouinia australis : Day, 1955: 418.
Body large, reaching 150 mm. by 6 mm. with numerous segments. It is uniformly
brown with orange gills and tentacles. Prostomium (fig. 20. 4. n) broadly rounded
in front without obvious eyes. Numerous tentacular cirri arise above setigers 3-4.
Branchiae from setiger i to the posterior end. Individual filaments are stout and
in the middle of the body they arise further above the notosetae than the distance
between notosetae and neurosetae (fig. 20.4.0). Capillaries present in both rami
of all parapodia. Sigmoid hooks appear about setiger 12.
Type locality : False Bay, South Africa.
Records: South West Africa (22/14/i, s and 26/15'i, s) ; Cape (from 30/17/i
and 34/18, i, s to 34/23/e and 32/28/i).
Distribution : Angola (s)
5i8 ?OI.VC:H.\F.TA OF SOITHERN AFRICA
Cirriformia filigera (Dclle C'hiaje, 1825)
(tig. 2n.4.p-q)
l.umhrku^ fdi^frt/s Dcllc Clhiajo i8'.J3 : 178.
Audouinin filigera: Fauvpl, i'j'-!7 : 92, tis;. 3J li-in.
Body uniformly brownish in colour \vith reddish gills. Length up to 250 mm.
Prostomium (fig. 20. 4. p) bluntly conical. Eyes not visible in adults. Numerous
tentacular cirri arise abo\e setigers 4-6. Branchiae from setigcr i to near the posterior
end and in the middle of the body (fig. 20. 4. q) they arise further above the notosctae
than the distance between notosctae and neurosetae. Capillary setae in both rami
ol' all parapodia. Sigmoid hooks appear about setigcr 12.
Type locality : Naples.
Records: Natal (29/31 i, s) ; Mocaniljiquc (26/32/1 and 23/35/s).
DisTRiBi TioN : Warm and tropical Atlantic (Morocco (i, d), San Thome ; North
Clarolina (i) ; Gulf of Mexico (s) ; Mediterranean (i, s) ; Persian Gidf ; tropical
Indian Ocean (i) ; ? Pacific.
Familv HETEROSPIONIDAE Hartman, 1065
(including LONGOSOMIDAE Hartman, 1944)
Body long and slender, rounded in section and divided into two regions. Pros-
stomium a blunt depressed cone without appendages. Pharynx soft, eversible and
miarmed. Peristomium \vcll developed with a pair of grooved palps but no para-
podia or setae. An anterior region of nine short segments with poorly dcxeloped
biramous parapodia and a pair of cirriform branchiae above the notosctae. Simple
capillaries in both rami and sometimes acicular spines in the first ncuropodium.
Posterior region with greatly elongated segments without parapodia! projections
or branchiae. Posterior setae are simple capillaries and/or acicular spines.
Remarks: Hartman (1944b and 1965) gives a general account of the family and
the few known species. None has been recorded from southern Africa.
TROCHOCHAETIDAE AND APISTOBRANCHIDAE 519
Family TROCHOCHAETIDAE Pettibone, 1963
( = DISOMIDAE Mesnil)
Prostomium rounded ; peristome reduced, with or without posterior tentaculiform
appendages. Dorsal lip with or without an anterior digitiform projection. Two long
gi-ooved palps. Body divided into distinct regions. Parapodia well developed and
usually biramous with prominent postsetal lobes but some of the middle segments
lack notopodia in Trochochaetus. Setae of several types but always simple ; they include
acicular ncurosetae on setigers 2, 3 and 4, also slender capillaries and barbed and
spiniform forms on posterior parapodia. No hooded hooks.
Key to Genera
I Parapodia all biramous. One or more tentaculiform lobes behind the prostomium
POECILOCHAETUS
— Middle region uniramous (without notopodia). No free tentaculiform lobes behind the
prostomium though a keel may be present ..... TROCHOCHAETA*
(= Disoma auctorum)
Records from southern Africa
Poecilochatus serpens AWen .... 5iCs, 55Ca
REMARKS
The lar\ae of Poecilochaetus, which are easily recognised, were known long before
the adult. They may be found even in oceanic plankton samples which suggests that
the larval stage is prolonged. Allen (1904) gave a very careful account of the mor-
phology of tlie adult and some interesting notes on its habits. Apparently near
Plymouth they make burrows in mud-filled crevices and stretch out their very long,
grooved tentacles in search of detritus. In other areas they may be dredged from
deep water on muddy bottoms and appear to live in friable mucus tubes covered
with foraminiferan shells.
POECILOCHAETUS Ehlcrs, 1874
Prostomium rounded with two pairs of eyes. Peristome reduced, with 1-3
tentaculiform lobes posteriorly. Dorsal lip widi a digitiform process. Two long
grooved palps. First setiger with elongated notosetae and neurosctae. Postsetal lobes
elongated but vary in shape. Filiform branchiae beliind middle and posterior para-
podia. Setae are mainly capillaries which are smooth anteriorly and plumose
posteriorly. Barbed setae from middle segments onward. Acicular hooks in the
neuropodia of setigers 2, 3 and sometimes 4 and in the notopodia of posterior segments.
Type species: Poecilochaetus fulgoris Ehlers, 1874
I'ol,^■c;IIAl•, r.\ oi- southern airica
Ki V 111 Si'Ei us
I Nciiropifdial linoks in setigcrs ■:;-4. Anterior dorsiini wilh tiibcrtles . . P. fulgoris'^
- Ntnu"0|:)odial huoks in sctigers 2-3. Anterior dorsum smooth ..... 2
'2 Three tentacnliforin lobes behind the prostominm. Filamentous gills on pcisterior face
of posteritir parapodia .......... P, serpens
— One tentaculiform lobe behind prostoniium. No gills on middle or posterior parapodia
P. johnsoni"^
Poecilochaetus serpens .MIcu, 1904.
(fig. -i.i.a-j)
Poecilochiiiiiis s,rpeii\ .Mien, 190.1: 79, pis. 7-12 text-fig. i ; Fativel, 1927: 67, fig. 23 a-m ; Day,
19IJI : 497.
Body (fig. 21. 1. b) slcndt-r, up to 55 mm. long witli iio scgnieiUs. Prostomitim
(fi.g. 21. 1. a) roundfd with Imir (.yes. Palps long, groii\c:d and caducous. A digiti-
form process arising from the dorsal lip. Three long nuchal processes arising from
the occipital region. First sctiger often with a vestigial notopodial jiostsetal lobe
and a long cirriform ]30stsetal neuropudial lobe. From setiger 2 to sctiger 6 both
i ':' if
Mi
„ ^^Lm
Fn.. Ji.i. PfMcilochai-tics serfiens. (a) Head, iv.) Entire worm (twice natural size), fc) Fourth
fool, fiji Anterior iiook. (i-) Posterior view (jf posterior foot showing branchiae, (f) Middle
foot ( loth). (g) Smooth capillary, (iij Feathered capillary, (i) Posterior spine (after
Fauvel;. {]) Posterior serrated spine (after Fauvelj.
TROCHOCHAETIDAE AND APISTOBRANCHIDAE 521
notopodia and ncuropodial lobes are tapered (fig. ai.i.c), from setiger 7-13 they
are club-shaped with swollen bases, slender shafts and knobbed tips (fig. ai.i.f)
but from setiger 14 onwards they are again tapered (fig. ai.i.e) with two to four
filamentous gills behind middle and posterior parapodia. Long capillaries of setiger
I directed forwards to form a cephalic cage. Two to four curved spines (fig. 2i.i.d)
in the neuropodia of setigers 2 and 3. The rest of the anterior setae are all smooth
capillaries (fig. 2i.i.g) plus a few feathered setae (fig. 2i.i.h) in middle segments;
these persist in posterior parapodia. On the last 16 or 17 setigers the notosetae are
replaced by simple or serrate spines (fig. 2i.i.i and j).
Type locality : Plymouth, England.
Records: Cape (from 32/i7/d,34/i7/vd and 34/18/s to 34/26/d) ; Natal (29/31 /s,d).
Distribution : N. Atlantic from Scotland (s), Ireland and the EngHsh Channel (i)
to Morocco (s) ; Mediterranean.
Family APISTOBRANCHIDAE Mcsnil and Caullery, 1898
Small elongate worms with numerous segments. Prostomium without appendages
but with well developed nuchal organs. Pharyn.x soft and unarmed. Peristome with
parapodia and a pair of long grooved palps. Notopodium reduced to a postsetal
cirriform lobe with an internal aciculum. An inter-ramal cirrus sometimes present.
No branchiae. Neuropodium with a lamellar postsetal lobe and a fascicle of simple
capillary setae. Pygidium with anal cirri.
Note: No representative has been recorded from South Africa. A single genus
Apistobranchus is know from northern seas. A review will be found in Orrhage (1962).
522 POI.VCHAKTA Of SOUTHF.RN AFRICA
Family CHAETOPTERIDAE Malnisrm, 1867
Very soft mucilaginous worms always protected in a tube. Body elongate and
composed of numerous segments always divided into three regions. Prostomitun
inconspicuous and enfolded by a broad, collar-like buccal segment. Head append-
ages include a pair of long groo\ed palps and sometimes a pair of inconspicuous
tentacular cirri. Anterior region of nine to fifteen uniramous, dorsally flattened
segments with stout acicular setae in setigcr 4 and elongate simple setae elsewhere.
Middle region of biramous segments with fine internal notosetae and minute ser-
puliform uncini in the neuropodia ; the noto|3odia are often bilobed and one or
more pairs produce mucus bags which are caught by cup-shajsed organs and carried
lorward to the mouth. Posterior region of biramous segments always with simple
notopodia which do not produce mucus bags.
Records front southern Africa
C/taiio/ilciiis vaiicnpedalui (Renier)
as C.'hactopterus ctipiiisii .Stimpson
as Chaetopterus hamalus Schmarda .
Mesochaetopterus capensis (Mcintosh) .
as Ranzania capensis Mcintosh
as Ranzanides capensis (Mcintosh) .
Mesochaetopterus minutus Potts .
Phyllochaetopterus elioti Crossland
Phrllochaetopterus herdmani Crossland .
PhyUnchaeloptcrus sncialis Claparcde .
as Phyllochaetopterus pictus Crossland
Spiochaetopterus costarurn (Claparcde) .
as Telepsanis costarurn Claparcde .
Spiochaetopterus trpicus Sars
Spiochaetojitcrus vilrarius (Ehlcrs)
as Tehpumts vitratiiis Ehlers
THE MAIX niAC;NOSTI(; CI lARACI I'.RS
Useful accounts of the family Cihaetopteridae will he finn<l in Potts (1914),
FauN'el (1927) and Barnes (1964).
All gcner.i arc tubicolous with an extremely fragile 1 pithc Hum which produces
abundant mucus. When the worm is preserved in its tube, tiie mucus hardens iiround
it and it is \ery dillicult to remo\e the worm from its tube without damaging the
I'ragile parapodia upon which identification depends. The most important charac-
ters include the nature of the head appendages, the number and shape of the enlarged
setae of the fourth setiger, the shape of the notopodia of the middle region and the
nature of the tul)i'.
i lie h( ad is Iruni ate with a small ovoid proslomiuni eiiliildcd by a broad glandular
i^Ci,
26Ais
, 27Mi
, 32PS,
:33As,
36C:i,
4oPi,
44Ci, 45PiNi
, 5iC:s
2Ci
4Ci
loCs
t") I Cs
3f!Xi,
4oXi,
4,-,Pi.\
i, 48C;
sWs
451^1,
51C1
45 Pi
32Cd
33AS
, 4oNi,
44Ci,
45Pi,
5 1 Cs
15CS,
2lCi
2 7 Mi
5.Cs
— ?C
— Ns
i5Ai
CHAETOPTERIDAE 523
buccal segment. The latter is always provided with a large funnel-shaped lower lip
and a pair of grooved palps similar to those of the family Spionidae. These palps
often show a characteristic pigment pattern which is worth noting. In Phyllo-
chaetopterus alone there is a pair of minute tentacular cirri with fine internal acicula.
They arise from the dorsal surface of the first setiger and lie on either side of the
prostomium. In well preserved specimens they are easy to sec but when the specimen
has been fixed inside the tube the head is usually covered with congealed mucus
and the tentacular cirri arc difficult to distinguish. Since the possession of these
tentacular cirri is the main character which distinguishes Phyllochaetopterus from
Spiochaelopterus the two are often confused.
The regions of the body. The number of uniramous segments which forms the
anterior region is probably constant but allowance must be made for autonomy and
subsequent regeneration. The fourth setiger however is always recognisable by its
enlarged and modified setae which are used to cut the partitions between the annuli
of the tube or even the tube wall itself when a side branch is formed. The number
and shape of these speciahsed setae is a useful character.
All the segments of the middle and posterior regions arc biramous but the foH-
aceous, bilobed notopodia are limited to the middle region since Barnes (in press)
has shown that even Telepsavus has a short posterior region of simple conical noto-
podia ; it thus becomes a synonym of Spiochaelopterus. In Spiochaelopterus and Phyllo-
chaetopterus the change from the middle region with its bilobed notopodia to the
posterior region with its simple knobbed notopodia is well marked ; in Chaetopterus
the last three segments of the middle region have fused notopodia but in Meso-
chaetopterus where the notopodia of the middle region are simple, the change from
middle to the posterior region is gradual and depends on the relative length of
the segments.
The nature of the tube. In Chaetopterus the tube is broad, papery and opens on the
the surface at both ends. In Mesochaetopterus it is fragile and covered with sand or
foraminifera shells and one end is buried. In Phyllochaetopterus and Spiochaetoptnus
the tube is horny and ringed with one end buried or attached to the substrate.
Barnes has shown that the rings correspond with internal partitions pierced by
minute pores.
Feeding mechanisms. It has long been known that Chaetopterus is a cihary feeder and
Barnes (1964) has shown that all the other genera feed in essentially the same way.
They produce a feeding current either by the pumping action of the fused notopodia
(in Chaetopterus) or by the beating of cilia on the smaller non-fused notopodia of the
middle region in other genera. The food current passes down the dorsal surface
of the body and the enlarged notopodia of the second segment of the middle region
secrete a mucus bag in which food particles are trapped. In some species more
than one mucus bag is formed. Posterior to the mucus bags there are cup-shaped
organs or cupules which roll up the food bags and these are then passed forwards
along a ciliated mid-dorsal groove to the mouth. Faecal pellets are carried forwards
in the same way but on the head these are transferred to the palps and voided from
the tube. The palps eliminate not only egested material but also large inedible
j:^ I'OI.VCIHAETA OI' SOU Ili KRN AFRICIA
particles which enter the tube and their primitive function of detritus c;atherini^
organs appears to be of ininor importance in some cases.
Kf.v to Genera
1 Middle region with Ijilobed notopodia. lube often horny and ringed .... 2
- Middle region with the notopodia never bilobed. Tube not horny or ringed ... 3
2 A pair of minute tentacular cirri as well as large grooved palps PHYLLOCHAETOPTERUS
(P- 524)
- Xo tentacular cirri, only a pair of large grooved palps . SPIOCHAETOPTERUS (p. 527)
3 Bodv large. Palps short. Some notopodia of middle region fused to ionn pariflles
CHAETOPTERUS ip. -,29)
- Bod\' iisualis" small. Palps large. Notopodia fif mitldle legiim never fused
MESOCHAETOPTERUS [p. :,3i)
PHYLLOCHAETOPTERUS Griibe, 1863
Body with three well delincd body regions. A pair of long groo\ed palps and a
pair of minute tentacular cirri arising froiri the dorsal surface of setiger i on cither
side of the prostomium. Anterior region uniramous with enlarged setae on setiger 4
and paddle-shaped setae elsewhere. Middle region biramous with bilobed foli-
aceous notopodia, lateral branchial lobes and bilobed neuropodia. Posterior region
biramous without branchial lobes but \vith cxlindiical notopodia and ncinopodia
as uncigcrous ridges.
Type speciies : Phvllnchaclnpterus niacilis Grube, 1863
Key to Si'Ecirs
1 Middle region of two segtnents. A srjiitary species with a sand-encrusted tube P. herdinani
- Middle region with seven or more segments ........ 2
2 A large solitary species with a fragile, sand-encrusted tube. Two or mote spines on setiger 4
P. elioti
- A small gregarious species with a tough, horny ringed tube. .\ single spine on setiger .\
P. socialis
Phyllochaetopteriis herdtuani Willey, 1905
(hg. 22.i.a-c)
Plivlliiiliaiiii/'Iniis lurdmani Willey, 1905: 292, p. 5 fig. 127-132; Fauvel, i9-,3 : 342, fig. 177 i-m ;
Day, I9",7 : 100, fig. b m.
A large species reaching 80 mm. Prostomium without eyes. Tentacular cirri
(fig. .12. 1. a) long, tapered and supjiorted by fine internal setae. Buccal segment
large, truncate and cur\ing upwards. Palps barred. Anterior region of nine to ten
segments. Segment 4 with six to eight stout setae obhquely truncate at the ends
(fig. 22. 1. c). Middle region with two segments with elongate bilobed notopodia
(fig. 22. 1. b), flattened branchiae and large expanded wing-like neuropodia with
uncini along the whole external margin. Posterior region of numerous segments
bearing digitiform notopodia with knobbed tips. Neuropodia are inferior scjuare
CHAETOPTERIDAE 525
projecting pinnules. Uncini (fig. 22.i.d) witii twelve to fourteen teeth. Tube
(fig. 22. 1. e) fragile and thickly encrutscd with sand-grains and shell fragments.
Type locality : Galle, Ceylon.
Records: Natal (30/30/i) ; Mocambique (26/3 i/i).
Distribution : Ceylon.
Phyllochaeopterus elioti Crossland, 1 903
(fig. 22.1.f-g)
PhyUochaetoptena elioti Crossland, 1903 : 172, pi. 16 figs. 1-3, 8, pi. 17 figs. 10-13 ; Day, 1957 : 100.
A large species 50-100 mm. long by 2-3 mm. broad. Buccal segment truncate,
palps barred with brown. Tentacular cirri which are well developed and digiti-
form, arise at the sides of the prostomium which bears one pair of eyes. Anterior
region of fifteen segments. Fourth segment with two to three enlarged and truncate
setae (fig. 22.i.g). Middle region of more than twenty short segments with bilobed
notopodia (fig. 22.i.f) containing six slender setae. Triangular lateral branchiae
are condnuous with the ventro-lateral neuropodia which bear numerous minute
uncini with about thirty teeth. Posterior region with slender knobbed notopodia
each supported by a single spear-headed seta. Tube horny, not ringed and densely
encrusted with sand.
Type locality : Zanzibar.
Records: Cape (34/22/s) ; Mocambique (26/32/i).
Distribution : Zanzibar (i) ; Ceylon and the Gulf of Manaar.
Phyllochatopterus socialis Claparede, 1870
(fig. 22.i.h-r)
Phyllochaetopterus socialis Claparede 1870 : 345, p. 21 fig. i ; Fauvel, 1927 : 84, fig. 30 a-I.
A small worm (fig. 22.i.i) seldom exceeding 30 mm. in length and dark green
posteriorly. Prostomium (fig. 22.1.J) ovoid with two lateral eyes often hidden by
the small tentacular cirri. Buccal segment truncate in front but curving up laterally
on the sides of the prostomium. Two long stout grooved palps. Anterior region of
ten to eighteen segments. Fourth setiger with one stout brown specialised seta
(fig. 22. 1. n) having an irregularly truncate end. Middle region of seven to twenty-
four segments bearing bilobed notopodia dorsally and triangular gills laterally
(fig. 22.1.1). Uncigerous portion of ncuropodium ventro-lateral. Uncini (fig. 22.1. r)
with about twenty-four teeth. Posterior region of numerous segments with digiuform
52G
l'OL^■c:HAI■:^A oi' .souiulr.x atrica
Fi'.. jj.i. Phyllocltiieloph'ru\ hnihnanl. (A) Anlcrior end. (b) I'ool from middle region, (c)
I^iilnigetl seta from fourih Icjot. (i>; L'ncinus. (i:) Part of sandy tuljc. Phvllocli(uto/'leriii
elioli. (I-) Foot from mid-region, (c;) Enlarged srla Irom fourth foot. Phyllochnelnplertn
iocialis. (h) Part of branching group of tubes, (i) Entire worm (four times natural size).
{ConUl. tit foot of fHi'^f 7-'7]
CHAETOPTERIDAE 527
notopodia bearing one to two spear-headed setae (fig. Q2.l.q). Tube (fig. 22. 1. h)
horny and ringed distally, often aggregated into branching colonies.
Type locality : Italy.
Re(;ords ; South West Africa (22/14/s and 26/15/i, s) ; Cape (from 2g/i6/i and
34/i8,i, s, d to 33/28/s) ; Natal (29/31/i, s) ; Mocambique (26/32/i, 24/34/s, 24/35/d).
Distribution : Atlantic from the U.S.A. and the Enghsh Channel south to the
Falkland Islands and South Africa ; Mediterranean ; tropical Indian Ocean
(i, s, d, vd) ; Australia.
SPIOCHAETOPTERUS Sars, 1853
(including TELEPSAV US CosU, 1861)
Fairly large but very soft worms in ringed horny tubes. Three distinct body regions.
Prostomium small and enfolded by the truncate buccal segment which bears long
grooved palps but no tentacular cirri. Fourth setiger with one to two stout spines.
Middle region with a variable number of segments bearing bilobed notopodia. A
short posterior region bearing conical notopodia.
Type SPECIES : Spiochaetopterus tjpicus Sars, 1853.
Key to Species
I Middle region of two to ten segments ........ S. typicus
- Middle region of about 20 segments ....... .S. vitrarius
— Middle region of over 30 segments ........ S^ costarum
Spiochaetopterus typicus Sars, 1853
(fig. 22.I.S-t)
Spiochnetopltrus typicus Sars, 1853 : 390 ; Fauvel, 1927 : 82, fig. 29 a-i.
A fairly large species measuring 50-60 mm. with a horny, ringed tube. Anterior
region (fig. 22.1.S) of nine to ten segments. Fourth setiger with a single giant seta
obliquely truncate at the end. A brown patch on the vcntrum of setigers 5-6.
Middle region of two to ten segments bearing large biolobcd notopodia with fine setae
embedded in the larger inner lobe. A triangular lateral gill on the side of each
segment. Neuropodia ventro-lateral. Uncini with about thirty teeth. Posterior
(j) .Anterior end (palps omitted), (k) Foot from anterior region, (l) Foot from mid-region.
(m) Foot from posterior region, (n) Enlarged seta from fourth foot, (o and p) Xotosetae
from anterior region, ((j) Notosetae from posterior region, (r) Uncinus. Spiochaetopterus
typicus. (s) Anterior end. (t) Notopdoium from posterior foot. Spiochaetopterus vitrarius
(after Ehlers, 1908). (u) Lateral view of anterior region, (v) End of enlarged seta of fourth
foot.
528 I'Ol.VtiHAK TA OK SOriHERX AFRICA
regicni with liiigcr-likc iidtopodi.i sujiportccl by two spcar-tippccl setae (fig. jj.i.t).
No lateral gills. Uiicini with minute teeth.
Type locality: Northern Norway.
Records: ? Cape (34/1 8 s).
DisTRiBiTiON : .\rctic (s) ; North Si-a ; North Carolina (d).
Spiochaetopterus vitrarius (Ehlers, 1908)
(fig. 22.1.U-V)
Tekpmvus vilrarius Ehlers, 1908 : 1 14, pi 15 figs. i-8.
A slender species over 35 mm. long, consisting of an anterior region of nine
uniramous segments, a middle region of about twenty segments of decreasing length
and a posterior region of numerous short segments. Prostomium ovoid with a pair
of lateral eye-spots. Buccal segment broad, truncate and curving upwards laterally.
Palps long and barred distally. No tentacular cirri. Segment 7 with a sharply
defined arc of black pigmenton the ventruin (fig. 22.1.U) preceding a w'hite glandular
patch on segments 8 and 9. Segment 4 with a fan of hastate setae and a single
enlarged seta with a swollen and conical tip (fig. 22.1.V). Notopodia of middle region
large, flattened and bilobrd. Lateral branchiae present. Posterior region with
short blunt notopodia supportetl by one to two long hastate setae. No posterior
branchiae. Neuropodia of middle and posterior regions bear undivided rows of
serpuhform uncini with ( ?j teetli. Tube c le.ir, horny and closely ringed.
Type locality : Great Fish Bay, Angola.
Records: N.ital (3o/30;s, 29/31,^).
Distribution: Tropical western .Africa from Senegal (s) to Angola (s).
Spiochaetopterus costariim (CHaparcde, 1870)
TdepuiiUi (uitmum Clapari'de, 1870: ^40, pi. 20 fig. 1 ; Fauvel, 1927 : 82, fig. 28 a-h.
Spiochaetofttnus costdTwn: Barnes (in press .
Body 50-60 mm. long, slender and fragile. Tube horny, ringed and transparent.
Palps long and speckled with brown. .Vnterior region ot nine to ten srtigers with
a large white glandular patch on the ventrum of setiger 7. One enlarged seta in
the fourth setiger w'ith the end swollen and truncate and the cutting edge serrate.
Middle region with \ery numerous segments, each bearint; bilobed notopodia
lateral triangular branchiae and neuropodia with mmn-rous untini with 25-30
sm.dl teeth. ,\ < upule on middle segment 2. Pusterior region short with crowded
segments bearing digitiloini notrjpddia .md reducerl neuropodia.
CHAETOPTERIDAE 529
Type locality : Naples.
Records: Cape (34/18/s).
Distribution- : Atlantic coast of France (i, s) ; Mediterranean (s) ; Madagascar
(i) ; Japan and \V. Canada to Southern California.
CHAETOPTERUS CuMCT, 1827
A large flabby worm inhabiting a broad parchment-like tube opening onto the
surface at both ends. Body divided into three distinct regions, an anterior region of
uniramous segments, a middle region which has the posterior notopodia fused
across the dorsum to form paddles and a posterior region with digitiform notopodia.
Buccal segment with short tapered palps but no tentacular cirri. Fourth setiger
with several stout spines.
Type species : Tricoelia varieopedatus Renicr, 1 804.
Chaetopterus varieopedatus Renier, 1804
(fig. 22.2.a-g)
Yricotlia varieopedatus Werner 1804: xviii.
Chaeloplenis varieopidaltis : Fauvel, 1927: 77, fig. 26 a-n.
Body about 100-150 mm. long by 10 mm. broad with 70 segments (fig. 22. 2. a).
Head truncate with a very small prostomium and a very large buccal segment
which forms the glandular lower lip. A pair of short tapered palps and a pair of
small eyes. Anterior region with eight to twelve segments bearing triangular
notopodia. Notosetae with flattened knife-Uke blades except in setiger 4 where
there are several stout, obliquely truncate setae (fig. 22. 2. g) in addition to a few
capillaries. Middle region with five biramous segments, the first short with small
notopodia, the second long with large wing-like notopodia anteriorly and a cupulc
posteriorly and the third, fourth and fifth with large notopodia fused across the
dorsum to form paddles. Neuropodial uncini (fig. 22. 2. c) with six to eight teeth.
Posterior region with over 20 segments bearing digitiform notopodia with knobbed
ends and bilobed neuropodia (fig. 22. 2. b).
Type locality : Mediterranean Sea.
Records: Cape (from 33, i8,i, s and 34,'23/e, i to 32/28/i) ; Natal (29'31,'i, s) ;
Mocambique (26/32/i, s).
Distribution : Cosmopolitan apart from the Arctic and Antarctic.
53°
l'()I.Vt:HAi:rA OF SOUTHERN AFRICA
I'lG. 22.2. Chat-!nptfrii\ iiu iei'/irfldfit^. ( \i Entire worm [natural size), (iii Foot from tlie poster-
ior region, (r.l L'nrinus. (Oi Lateral view of fourth and fifth segments of mid-region.
'1 I .Anterior capillary seta, (f) .■Anterior oar-shaped seta, (o) Enlarged seta of setiger 4.
Mfsoclmfloplrmt minultis. (11) Part of sandy tube, (i) Uncinus. (j) Second foot of mid-
region. (Kj Foot from tail region, (l) Enlarged seta of setiger 4. (m) Head end. (n)
Entire worm {four times life size). Mewchaetoptmis cajicmii. (o) .'\nterior end. (p) Second
uncigerfjus foot, fq 1 Enlarged seta of setiger 4. (r) Uncinus. Spinchaftopti-riis costariun.
(Sj Part (jf tube, (ij Lateral view of head end. (v) Posterior foot. (\'} L'ncinus.
CHAETOPTERIDAE 531
MESOCHAETOPTERUS Potts, 1914
(including RAAJ^AAIA Claparcde, 1870 (preoccupied) and
RANZ-'iMDES Chamberlin, 1919)
Body with three regions. Buccal segment with long grooved palps but no tenta-
cular cirri. Anterior region with uniramous segments. Fourth setiger with several
specialised setae. Middle region with a few long segments bearing simple notopodia
and neuropodia with divided rows of uncini. Posterior region not sharply marked
off from the middle region but the segments are short, the notopodia digitiform and
there are no cupulcs. Tube fragile and encrusted with sand or shell fragments.
Type species : Mesochaetopterus minulus Potts, 19 14.
Key to Species
1 Middle region of three segments of which the second and third bear cupules. .\ large
solitary' species measuring 100 mm. or more ...... M, taylori*
- Middle region of two segments of which the second bears a cupiile. .Small species
measuring less than 35 mm. .......... 2
2 .\nterior region of 10-13 segments with four to se\-en modified setae in setiger 4. Posterior
notopodia each with a single seta. Animals gregarious . . . Af. minutus
- .interior region of 12 segments with "several" modified setae in setiger 4. Posterior
notopodia each with ( ?) notosetae. .\nimals not gregarious . M. sagittaria*
- .\nterior region of nine segments with eight modified setae in setiger 4. .Vnimals not
gregarious ... ........ M. capensis
Mesochaetopterus minutus Potts, 19 14
(fig. 22.2.h-n)
Mesochaetoplerus mlmilus Potts, 1914: 963, pi. 2 fig. 4, pi. 3 figs. 7-8; text-figs. 4 and 5.
A small species about 15 mm. long living gregariously in dense masses of fragile
sandy tubes (fig. 22. 2. h). Body (fig. 22. 2. n) not pigmented. Prostomium relatively
large and ovoid with a pair of eyes. Buccal segment broad and truncate with a
pair of long grooved palps. Anterior region of ten to thirteen segments. Setiger 4
with four to seven modified setae (fig. 22.J). Middle region of two long segments
with simple flattened notopodia supported by two to three setae. One cupule in
the middle of the second segment. Branchiae absent. Neuropodia notched and
uncigerous row divided. Uncini with seven to nine teeth (fig. 22. 2. 1). Posterior
region of numerous segments decreasing in length each bearing short conical noto-
podia supported by a single seta and finally a few segments with knobbed notopodia
(fig. 22. 2. k).
Type locality : Torres Strait.
Records: Natal (30/30/1 and 29/31/i) ; Mocambique (26;32/i).
Distribution: Tropical Indo-west-Pacific (i) to Japan.
53J roi.YCIlAE TA OT SOUTHERN AFRICA
Mesochaetopterus capensis (Mcintosh, 1885)
(fiy-. 22.2.0-1)
Rdii-iiniii iii/hnsis .\Ii Inlnsh. iHfi", : ^jyfi, pi. 2J\\ figs. i()-2u.
Riinzdiiides ail»'nsis : O.iy, ii|(ii : 411!!.
Body about 15 mm. long with more than 20 segments. Anierior region ol nine
uniramous segments (fig. 22.2.0). Prostomiuni ovoid. I'.ilps long, Liinlly ijarred
distally and with a pair of eyes at llieir bases. Fourth setiger with eight modified
setae having obliquely truncate and serrated tips (fig. 22. 2. q). Middle region ot
two long segments ofwhicii the first has a pair of short notopodia close to the last
anterior segment. Tiie second has a pair of longer strap-like notopodia (fig. 22.2. p)
and a cupule posteriorly. Neuropodia as bilobcd ridges with a di\ided series of
uncini. Individual uncini (fig. 22. 2. r) with six to nine teeth. Posterior region with
short segments bearing simple conical notopodia supported by two setae with
flattened, tapering blades. Tube fragile and enenisted with sand or foraminifera
shells. Animal solitary.
Type locality : Dredged off the Cl.qic of Good Hope.
I\.Ei:oRDs: tlape (from 34/18/s, d and 36 2i,d to 34/23/s) ; Xatal (30/30/s).
Distribution: Endemic.
ORBINIIDAE
533
Family ORBINIIDAE Hartman, 1942
Body vermiform and divided into an anterior thoracic region of rather flattened
segments and a posterior abdominal region of numerous rounded segments. Pro-
stomium conical without sensory appendages or palps. Proboscis unarmed. Buccal
segment achactous and sometimes also the next. Cirriform branchiae are inserted
on the dorsal surface between the notopodia and usually extend over most of the
body. Notopodia consist of finger-like postsctal lobes each with a bundle of crenu-
late capillaries and often a few forked setae. An interramal cirrus may be present
on abdominal segments. Neuropodia arc in the form of vertical ridges on the sides
of the thorax and may be accompanied by a postsetal row of foot-papillae which
extend onto the ventral surface as stomach papillae. Thoracic neurosetae include
crenulate capillaries and usually acicular hooks as well. Abdominal neuropodia are
dorso-lateral projections usually bilobed. A ventral cirrus is usually present.
Abdominal neurosetae are crenulate capillaries.
Records front southern Africa
Haploscoloplos cLfragilis Verrill
Haploscoloplos kerguelensis (Mcintosh)
as ? Theosloma oersetedi Kirkegaard (non
Claparede) .....
Naineris laevigata (Grube)
as Anthostoma hexaphjUum Schmarda
as Theodisca hexaphyllum (Schmarda)
asScoloplos {Naidonereis) hexaphyllum (Schmarda
as Nainereis hexaphyllum (Schmarda)
as Scoloplos armiger Monro (non Muller)
Orbinia angrapequensis (Augener)
as Alicia angrapequensis Augener
Orbinia bioreti (Fauvel) .
as Aricia bioreti Fauvel
Orbinia cuvieri (Aud. & M. Edw.)
Phylo foetida australis (Fauvel) .
as Aricia foetida var. australis Fauvel
Phylo foetidia Ugustica (Orlandi)
as Orbinia foetida (Claparede) var. Ugustica
(Orlandi)
Phylo capensis Day .
Orbinia monroi Day
Proscoloplos cyngochaetus Day
Schroederella pauliani Laubier
Scolaricia capensis Day
Scolaricia dubia (Day)
as Orbinia dubia Day .
56CS
5iCsd
48CS
36Ci, 4oNi, 4461,
Ni, 51CS
4Ci
isCi
2 6 Wis
33Ci
33Ci
44Ci, 51CS
26\Vs
45Ni, ?5iCis
2 7 Mi
— Ns
27Mi, 45Pi
44Ci, 5iCsd
5iCsd
44Ci, 45Pi
5iCi
54^Vi
51CS
51CS
44Ci
534 l'01.\t:HAKIA OI" SorillKKX AFRICA
Scololilelhi cajunsis Day ..... 56Cd
Scoloplds anniger Miillcr .....? ^iCsd
Sioloploi [Leodamas) jonstonei Day . . . StJCi, 44C'i,
45PiNi, 5iC:s
Scoloplos [Leodainai) madai:,aicariensh Faiucl . 27Mi, 4oPi, 4')Pi,
48CS, — s
Scnlojdns marsujiialis Sdiithcrn .... 45?'
Scoloplos sp. ....... 44Ci
Scoloplos {Lcodcmcis) iiniramus Day . . . 5iCsd, — Xs
RKM.VRKS
.\ rccrnt review of the family Orbiniidae will be found in Hartmaii (1057). ^lost
of the orbiiiiids burrow in banks of. sandy mud and do not make permanent tube.s.
They ingest jiartieulate organic matter but do not appear to be selective as tite gut
is often packed with large sand grains. The pointed prostomiimi and the flattened,
muscular anlcrinr region appear to do most of die burrowing work while the posterior
region, where the whole parapodium is dorsal to protect the gills, is respiratory and
digestive. Forms with more rounded prostimiia such as Aalneiis and Pioscoloplos are
found on rocky shores among tin- holdfasts of algae.
The important taxonomic characters include the .shape of the prostomiiim, the
numbi;r of segments whit h liirm the anterior region, tiie position of the first gill, the
shape of both the anlerior and posterior neuropodia and (inally the character of the
setae.
The head. As noted, the prostoniiiun m.iy be either a sharply pointed cone or a
rounded lobe above the mouth. The eversible proboscis is soft and often lobed but
it is usually retracted and not important. The buccal segment is always achaetous
and in tin- subfamily Protoariciiiiae it is followed by a second achaetous segment.
The anterior rai'ioti or thorax. This pai t of the body is flattened to varying degrees
and the neinripi>dia are mere lateral ridges bearing vertical rows of neurcisetae.
Behind the neurosetac however, there are one or luore conical postsetal lobes ov foot-
papillae whose position (superior, median or inferior) is important if there are only
one to three. In the abdomen the superior one becoines the postsetal lobe of the
neuropodium and the other one or two persist for several segments as ventral cirri.
In Orhitiia and Phxlo there are numerous foot pa]iillae and, on the transitional seg-
miMits between the tiiorax and .ibdomen, a xcntral row ct{ stomach papillae is developed.
The jw'scnce or absence of such papillae is important but their exact niunbcr and
segmental disposition is too variable to be of \alue.
The segment on which the first pair of branchiae arises is constant to within two
segments if branchiae start on setiger 5 -7 but varies up to four segments if the first
branchia a]i]jears between the 15th and -'")th setiger. This ability of the worm to
"cotnit more accurateh" inMrer its head is fairly conunon amimg polychaetes.
The ihor.icic notiisetae .uc .ill crenulate (laddered) capillaries, 'i'he neurosetae are
arranged in vertical rows. In the genus I/aploscoloplos they are all crenulate capillaries
but in otlier genera the anterior rows of neurosetae arc short, stout hooks (sometimes
ORBINIIDAE 535
called crocthels). These often have serrated ends protected by delicate bivalve guards.
Setae which are intermediate between hooks and capillaries are termed subuluncini
and have stout shafts which abruptly taper to delicate crenulated tips. Finally in the
genus Phjlo one or more stout spines much larger than any of the other setae arise
from the superior edges of the neuropodia of the last few thoracic setigers.
The posterior region or abdomen. The change from thorax to abdomen is marked
by the loss of neuropodial hooks and the movement of the whole neuropodium to the
dorsal surface of the segment. The postsetal lobe enlarges and between it and the
neuropodium a short interramal cirrus may develop. The neuropodium itself elongates
and becomes bilobed. The relative size or the presetal and postsetal lobe is of specific
importance and also the character of the whole lateral margin of the neuropodium.
Usually the basal part is flanged and may even form a broad lamella which divides
into two lamellae on posterior segments. In other cases a small ventral cirrus persists
over the first half of the abdomen.
Abdominal notosetae are mainly crenulate capillaries but these are often accom-
panied by a few forked setae. The latter are short and difficult to see unless the whole
notopodial bundle is frayed out on a slide. The neuropodium is strengthened by one
or two acicula which usually have tapering tips but in some species of Scoloplos the
ends are stout and project through the surface. The neurosetae are usually long
crenulate capillaries but in Scolaricia the blades taper very abruptly to whip-like tips
and arc aptly known as, flail setae.
As will be seen the characters change along the length of the abdomen and the
segmental position of a parapodium should be noted before it is removed for micro-
scopic examination.
Key to Genera
1 Both the buccal segment and the next is achaetous (fig. 23.1.3) (subfamily Protoariciinae) 2
- Only the buccal segment is achactous (fig. 23. 2. g) (subfamily Orbiniinae) ... 6
2 Branchiae present on abdominal segments ........ 3
- Branchiae completely absent ........ ORBINIELLA*
3 Only crenulate capillaries present ..... SCOLOPLELLA (p. 336)
- More than one type of seta present ......... 4
4 .'\bdominal neurosetae include swan-shaped hooks (fig. 23.i.g). Piostomiimt rounded
PROSCOLOPLOS (p. 53G)
- Swan-shaped hooks absent. Prostomium rounded or pointed ..... 5
5 .Abdominal neurosetae are all crenulate capillaries. Prostomium rounded PROTOARICIA*
- .\bdominal neurosetae include acicular hooks (fig. 23.i.k) as well as crenulate capillaries.
Prostomium pointed SCHROEDERELLA (p. 538)
6 Prostomium rounded or square. Thoracic neurosetae include subuluncini
NAINERIS (p. 539)
- Prostomium pointed. No subuluncini ......... 7
7 Thoracic neuropodia with numerous foot papillae (fig. 23.2.11). .Stomach papillae often
numerous ............. 8
- Thoracic neuropodia without foot papillae or only one to three. Few if any stomach
papillae .............. g
8 One or more enlarged spines in the superior part of last few thoracic neuropodia (fig.
23.2.m) . PHYLO (p. 540)
- No enlarged spines in thoracic neuropodia ..... ORBINIA (p. 543)
536 POIACllIAl.lA OK SOU rill.RX AiRUlA
9 Thoracic neuropodia with (Tcinilatc cajiillaries only ; no lioolcs IIAPLOSCOLOPLOS (p. 546)
- Thoracic neuropodia witti both liooks and crenulatc capillaries ..... 10
10 Flail-tipped ncurosctae (fig. 2.'3-4-l| in abdominal segments. Abdominal neuropodia with
an inferior lamella ........ SCOLARICIA (p. 347)
— No llail-tipped neurosetae. .Abdominal neuropodia without a distinct iiiTerior lamella
SCOLOPLOS (p. 549)
Sub-fnniilv PROTOARICIINAE Hartman, 1957
Small Oibiniidae with thr fust iwn ,sct;nicnts apodmis and achactous. Branchiae
sometimes absent.
SCOLOPLELLA Day, 1963
Prostomium conical. Both the buccal segment and the next achactous. Thorax
of few segments and not obviously flattened. Parapodia biramous but reduced to
small papillae, each ramus bearing crenulate capillaries only. No specialised setae.
Branchiae absent from the thora.x. No intermediate c irrus ; no \ entral cirrus nor
stomach papillae.
Type SPECIES : Scolophlla capcmis T>:\\\ 1963.
Scoloplella capensis Day, 1963
Mig. 23.i.a-d)
ScoloptiUa capensis Day, 1963 : 413, fig. 8 m-c).
Body small and thread-like, an incomplete specimen measuring 6 mm. for 25
segments. Prostomium (fig. 23.1. a) bluntly conical and without eyes. Peristome
and the next segment achactous. Thorax almost rounded, sharply marked off from
the abdomen and consists of seven short setigerous segments. Parapodial rami
(fig. 23. 1. b) close together and each reduced to a small papilla plus a bundle of
crenulate capillaries. Abdominal segments longer tlian broad, each with a short
blunt postsctal lobe in the ncjtopodium and a stout bilobed neuropodium ffig. 23.1.C).
No intermediate cirrus iror ventr.tl cirrus. Branchiae do not appear before setiger
24 and are thus absent from the thorax. Neurosetae are short broad-bladed crenu-
late capillaries (fig. 23.i.d) similar tu those of the thorax. No specialised setae.
Type Lor:.\i.nY : \\'. cjf Ciape Town in 183 metres.
Records: Cape (33/i7,d and 34/23;dj.
Distribution' : No other specimens known.
PROSCOLOPLOS D.\\, 1954
Minute worms with a rounded prostomium and the first two segments achactous.
No sharp di\ision between thorax and abdomen, the parapodia being poorly
developed and essentially similar. Setae include crenuhilc capill. tries plus swan-
shaped hooks in posterior neuropodia.
Type species : Proscoloplus cygnnchactus Day, 1954.
ORBINIIDAE
537
Fig. 23.1. Scoloplella capensis. (a) Lateral view of anterior end. (b) Thoracic segment,
(c) .Abdominal segment, (d, d') Crenulate capillary. Proscoloplos cygmchaetus. (e) Dorsal
view of anterior end. (f) Branchiate foot, (g) Swan-shaped hook. Schroederella pauliani
(after Laublier, 1962). (h) Lateral view of anterior end. (i) Thoracic foot, (j) Abdominal
foot, (k) Thoracic uncinus. (l) Abdominal neuroaciculum.
538 POl.VC:HAETA OF SOUTHERN AFRICA
Proscoloplos cygnochaetus Day, 1054
(fig. '-'S.i.e-g)
Proscoloplos n'gttocluitiiis Day, 1954 : 21, tig. 3 a-f.
Body 3-5 mm. long. Prostomium (fig. 23.1.0) rounded; first two segments
achaetous. Parapodia essentially similar throughout. Each notopodium consists
of a digitiform postsctal lobe with a few crenulate capillaries arising from its base.
Stout fusiform branchiae (fig. ag.i.f) from setigcr 8. Ncuropodia similar to noto-
podia throughout but lateral in position and smaller. Neurosetae include a few
crenulate capillaries and, in posterior segments, one to two swan-shaped hooks
(fig. 23. 1. g) with a row of four to fi\c denticles above the rostrum.
Type locality : Tristan da C'unha.
Records: Cape (34/18/i).
Distribution': Tristan da Cunha (i).
SCHROEDERELLA Laubier, 1962
Minute worms \vith a very pointed prostomium consisting of two annuli. Eyes
present. Buccal segment and the next both achaetous. Body divided into an
anterior thoracic region without branchiae and a posterior abdominal region with
branchiae. Parapodia poorly developed and foot papillae absent. Notosetac are
crenulate capillaries with the addition of a pointed aciculum in the abdomen.
Netirosetae include both cremdate capillaries and acicular hooks.
Type species : Schroederella pauUani Laubier, 1962.
Schroederella pauliani Laubier, 1962
(fig. 23.1. h-1)
Schrot'dtTtlhi /itiuliarii hauh'ier, 1962: 231. figs. 1-2.
Body minute, 3-4 mm. long for 40 segments. Prostomium (fig. 23.1.11) a long
pointed cone divided into two annuli. One pair of eyes. Thorax of about eight to
twenty segments without branchiae. Postsctal lobe of notopodium small and appears
on setiger 7-9. Thoracic neuropodia (fig. 23.1.!) as slight lateral swellings without
foot papillae. A short intermediate region of four to five segments between thorax
and abdomen. Abdomen of 16-20 segments v\ith digitiform dorsal branchiae (fig.
23. 1. j) and small postsctal notopodial lobes. Neuropodia better marked than those
on the thorax but essentially similar and lack ventral cirri. Notosetac are crenulate
capillaries throughout with the addition of a sharp pointed aciculum in the abdomen.
Thoracic neurosetae include a few crenulate capillaries and several serrated acicular
ORISIMIDAE 539
uncini (fig. 23.i.k). Abdominal ncurosetac include one to two crenulate capillaries
and a single blunt "acicular seta" (or ? projecting aciculum fig. 23.1.I).
Type locality : Sandwich Bay, South West Africa.
Records: South West Africa (23/14/i).
Distribution : No further records.
Subfamily ORBINHNAE Hartman, 1957
Orbiniidae with only the first or buccal segment achaetous and apodous. Branch-
iae always present.
NAINERIS BlainviWe, 1828
Prostomium rounded to square in front, depressed. Two subdcrmal eyes. Pharynx
with a frilled or lobed margin. Only the buccal segment achaetous. Branchiae on
most of the anterior segments. Thoracic notosetae may include a few forked setae
as well as crenulate capillaries. Thoracic neuropodia with a posterior foot-papilla.
No stomach papillae. Thoracic neurosetae include rows of hooks (uncini), crenulate
capillaries and often intermediate forms (subuluncini). Abdominal notosetae may
include a few forked setae with the crenulate capillaries. Abdominal neuropodia
bilobed with three to five acicula and a sheaf of crenulate capillaries. No vental cirri.
Type species : J^ais quadricuspida Fabricius, 1 780.
Key to Species
I Thoracic neuropociia with subuluncini and a median foot-papilla . N. quadricuspida*
- Thoracic neuropodia with subuluncini and a superior foot-papilla . . N. laevigata
Naineris laevigata (Grubc, 1855)
(fig. 23.2.a-f)
Aricia laevigata Grube, 1855 : 1 12, pi. 4 figs. 6-8.
Naineris laevigata : Fauvel, 1927: 22, fig. 7 a-1 ; Hartman, 1957: 297, pi. 35 figs. 1-8.
Body about 40 mm. long by 2 mm. across the thorax. Prostomium depressed and
smoothly rounded in front. Subdermal eyes not obvious. Pharynx eversible with
numerous marginal lobes (fig. 23.2.3). Thorax flattened with 15-30 setigers. Taper-
ing branchiae start on the fifth to eighth foot. Thoracic notopodium (fig. 23. 2. b)
with a sheaf of crenulate capillaries and a stout postsetal lobe. Thoracic neuropodia
as stout lateral ridges each with a superior postsetal lobe or foot papilla. Abdominal
notopodia (fig. 23. 2. c) more slender and elongated than those on the thorax. No
intermediate cirrus. Abdominal neuropodia bilobed, each with a short blunt
presetal lobe and a slightly longer and more pointed postsetal lobe. No ventral
cirrus. Thoracic notosetae include many stout, well tapered crenulate capillaries
and one to two forked setae with distally serrated shafts and unequal forks spinulose
on their inner margins. Thoracic neurosetae in four to five vertical rows ; most are
540 pc">i.vc;iiAr,i.\ oi- sol thekx airk:a
subuluncini (fig. '23.2.0) \vith stout bciit shafts abruptly tapering to weakly serrated
tips; inferiorly there is also a group of hooks (uncini) (fig. 23. 2. d) which arc bent
and faintly barred with guards near the blunt ti]5s and numerous crenulatc capillaries
(fig. 23. 2. f) like those in the notopodia. Abdominal notosetae are fine crenulatc
capillaries and two to three forked setae similar to those of the thorax. Nemosctae
include three to five acicula with projecting tips which are cpiite smooth and faintly
cur\cd as well as numerous crenulatc capillaries.
Type; locauty : Mediterranean Sea.
Records : South West Africa (22/14,1 - 26/15/1, s) ; C^ape (30/17/i and 34/18/i, s
to 33 26 s) ; Natal (29/31/i) ; Mocambicjue (26/32/i).
DisTRiBL'TiON : Bay of Biscay, warm and tropical Atlantic from North Carolina
(i) to Gulf of Mexico (i) and Brazil ; Gold Coast (i) ; Mediterranean ; Persian Gulf;
Ceylon ; Japan.
PHYLO Kinbcrg, 1866
Prostomium conical, proboscis lobed. Only the buccal segment achactous.
Notosetae arc crenulatc capillaries sometimes accompanied by a few forked setae in
the abdomen. Branchiae start on setiger 6-9. Thoracic ncuropodia with numerous
foot-papillae. Stomach papillae usually present at the junction of the thorax and
abdomen. Thoracic neurosetac include two to four rows of hooks, a few crenulatc
capillaries and towards the end of the thorax, one or more giant superior spines
arising from a glandular organ. Abdominal ncuropodia bilobed. An interramal
cirrus may be present.
Type species : Phylofclix Kinbcrg, 1866.
Krv TO Species
1 Xumcrous stomacli papillae ........... 2
- No stomacli papillae or only two to three ...... P. capensis
2 Twenty to Iwcnty-lhree thoracic segments ..... P. foetida ligustica
- Thirty-six to tiiirty-nine thoracic segments ..... P. foetida aiistralis
Phylo capensis Day, 1961
(fig. 23.2.g-l)
Phvlo cafien^h Day, 1961 : 47*3, fig. i a-f.
Length up to 50 mm. Seventeen to twenty-four thoracic segments (fig. 23. 2. g)
with branchiae from the sixth. Postsctal lobe of the notopodia well developed from
the first foot onwards but not expanded and chopper-shaped in the abdomen.
Thoracic ncuropodia (fig. 23.2.11) with up to nine foot papillae. Stomach papillae
either absent or limited to two to three at the junction of thorax and abdomen.
Abdominal ncuropodia (fig. 23. 2. k) bilobed and the ventral cirri well developed.
Intermediate cirrus cither absent or very small. Notosetae include the usual
crenulatc capillaries plus a few forked setae (fig. 23. 2. f) in the abdomen. Thoracic
ORBINIIDAE
541
f,
d e
Fig. 23.2. Nahuris laei'igala. (a) Lateral view of anterior end. (b) Tenth thoracic foot,
(c) Abdontinal foot, (d) Thoracic hook, (e) Subuluncinus. (f) Crenulate capillary from
thoracic neuropodium. Phylo capensis. (g) Lateral %iew of anterior end. (h) Fifteenth
foot, (i) Superior spine, (j) Forked seta, (k) Abdominal foot, (l) Thoracic hook.
Phylo foelida ligustka. (m) Fifteenth foot, (n) Superior spine, (o) Abdominal foot,
(p) Thoracic hook.
542 POLVC:HAK lA OF SOUTHERN AFRICIA
neurosctac include three to four rows of blunt, lightly serrated hooks which lack
guards (fig. 23. 2.1), a few capillaries and, from setigcr 14 to the end of the thorax,
a single enlarged superior spine with a smooth shaft (fig. 23.2.1).
Type locality : Saldanha Bay, South Africa.
Records: Cape (33/17/s, 34,t8's, 34/22/d).
Distribution- : Endemic.
Phylo foetida foetida (Claparede, 1870)
(fig. 23.2. m-p)
Arkia foetida C\?ip:\v^f\Gy 1870: 306.
Aricia foetida : Fauvcl. 1927: 14, fig. 4 a-e.
A large reddish brown species reaching 105 mm. and with a characteristic foetid
odour. Twenty to thirty-nine thoracic segments with branchiae from setiger 6-9
onwards. Postsetal lobe of notopodium well developed tlirougliout. Thoracic
ncuropodia (fig. 23. 2. m) with ten to fifteen foot papillae and, at the junction of thorax
and abdomen, the segments have continuous rows of stomach papillae. Thoracic
neurosetae include three to four rows of flanged and serrated hooks (fig. 23. 2. p), a
row of crenulate capillaries and, from setiger 12 onwards, a large, dark, superior
hastate spine (fig. 23. 2. n). Abdominal notopodia with cultriform postsetal lobes
and a few forked setae among the crenulate capillaries. A well developed inter-
ramal cirrus. Ncuropodia with luiequal lobes and a well marked \entral cirrus.
Type locality : Naples.
Phylo foetida ligustica (Orlandi, i8g6)
Aricia foetida var. ligustica Orlandi, 1896 : 12, pi. 2 figs. 3-12 ; Fauvel, 1927 : 14, fig. 4 a-1.
Length up to 80 mm. A total of 20-23 thoracic segments with branchiae fnjm the
sixth or seventh.
Type locality : Mediterranean.
Records: C^ape (from 32/18 s to 34:'22/d) ; Mocambique (26/32/i and 23/35/e).
Distribution: Mediterranean (i) ; tropical western Africa (s).
Phylo foetida australis (Fau\el, 1919)
Aricia foetida var. auitralii Fauvcl, 1919: 429.
Length about too mm. A total of 36-39 thoracic segments with branchiae from
the sixth or seventh.
Type locality : Madagascar.
Records: Mocambique (23/35/e).
Distribution: ^Ladagascar (i).
ORBI.MIDAE 543
ORBINIA Quatrefages, 1865
Prostomium conical and pointed. Proboscis lobed. Buccal segment achaetous.
Notosetae are crenulate capillaries on the thorax but a few forked setae may also be
present on the abdomen. Branchiae start between segments 4 and 10. Thoracic
neuropodia with numerous foot papillae. Segmental rows of stomach papillae at
the junction of the thorax and abdomen. Thoracic neurosetae include two to four
rows of hooks plus a variable number of crenulate capillaries but no superior enlarged
spines. Abdominal neuropodia bilobed. An interramal cirrus may be present or
merely a ciliated button. Ventral cirrus present or absent.
Type species : Aricia cuvieri Audouin and Milne Edwards, 1833.
Key to species
1 Interramal cirrus between abdominal notopodia and neuropodia absent. (18-24 thoracic
setigers) .......... O. angrapequensis
- Interramal cirrus present ........... 2
2 22-26 thoracic setigers ........... O. cuvieri
- 32-39 thoracic setigers ........... O. bioreti
- .^bout 50 thoracic setigers .......... O. monroi
Orbinia angrapequensis (Augener, 19 18)
(fig. 23.3.a-d)
Aricia angrapequensis Augener, 1918 : 413, pi. 6 fig. 146, pi. 7 fig. 225, text-fig. 56.
Orbinia angrapequensis: Day, 1955 : 408.
Length up to 30 mm. for go segments. Eighteen to twenty-four thoracic setigers
with branchiae from the sixth (fig. 23. 3. a). Postsetal lobe of notopodium ob\ious
from the first foot. About 12 foot papillae per neuropodium and continuous rows
of stomach papillae on segments near the junction of thorax and abdomen (fig. 23. 3. b).
Thoracic neurosetae include three to four rows of flanged, bent, serrated hooks (fig.
23. 3. d) and a row of crenulate capillaries. Abdominal notopodia with only a few
forked setae among the capillaries. Abdominal neuropodia (fig. 23. 3. c) bilobed with
the inner ramus longer than the outer. No interramal cirrus but a ciliated cushion is
present. \'entral cirri present on all abdominal neuropodia.
Type locality : Luderitz, South ^Vest Africa.
Records; South \Vest Africa (22/14/s and 26/15/i, s) ; Cape (from 32'18,'s to
34/26/d).
Distribution : Endemic.
Orbinia cuvieri (Audouin & Milne-Edwards, 1834)
jlriWa Cuotfri Audouin and Milne-Edwards, 1 834: 258, pi. 7 figs. 5-13; Fauvel, 1927: lafig. 3e-l;
Fauvel, 1953: 301, figs. 155, 156.
Body up to 300 mm. long with 400 segments. Twenty-two to tv\enty-six thoracic
setigers with branchiae from setiger 5. Postsetal lobe of notopodium obvious from
the first foot. Thoracic neuropodia with 10-15 foot papillae and continuous rows
544
POLVCHAETA OF SOUTHERN AFRICA
^^}^;r^riifmfk^
-sjvkttv^^
nr~ h
Fig. 23.3. Orbinia angrapequemis. (a) Lateral view of anterior end. (b) Eighteenth thoracic
fr>ot. {<:.) Abdominal foot, (d) Thoracic neuropodial hook. Orbinia biorcti. (e) Twenty-
fifth thoracic foot, (f) Thoracic neuropodial hook, (g) Anterior abdominal fool. Orbinia
riionroi. (hJ Forty-fifth thoracic foot, (i) Abdominal foot, (j) Forked seta.
ORBIMIDAE 545
of Stomach papillae on segments at the junction of thorax and abdomen. Thoracic
neurosetae include three to five rows of flanged, bent and serrated hooks and a few
crenulate capillaries, but no enlarged superior spines. Abdominal notopodia with
chopper-shaped postsctal lobes and larger tapered gills. A well developed inter-
ramal cirrus. Abdominal neuropodia bilobed and with a small conical ventral
cirrus. Abdominal notosetae are mainly crenulate capillaries plus a few forked
setae with unequal prongs ; neurosetae are a few short crenulate capillaries.
Type locality : France.
Records: Natal (29/31 /s).
Distribution: North Atlantic from Greenland (s), Sweden (s), North Sea (s),
English Channel (s) ; Mediterranean (s).
Orbinia bioreti (Fauvel, 191 9)
(fig. 23.3.e-g)
Aricia bioreti Fauvel, 1919 : 430, pi. 16 figs. 52-56 ; Fauvel, 1927 : 13, fig. 3 a-d.
Length up to 130 mm. with more than 120 segments. Thirty-one to thirty-nine
thoracic setigers with branchiae from the fifth or sixth. Postsetal lobe of notopodium
obvious from the first foot. About 12 foot papillae per neuropodium (fig. 23. 3. e)
and continuous rows of stomach papillae on segments at the junction of thorax and
abdomen. Thoracic neurosetae include four to five rows of flanged and bent but
smooth hooks (fig. 23. 3. f) and a very few crenulate capillaries. Abdominal noto-
podia without forked setae. Intcrramal cirrus rudimentary in anterior abdominal
segments but well marked posteriorly. Abdominal neuropodia (fig. 23.3.g) with
small unequal lobes. Abdominal neurosetae (fig. 23. 3. g) with small unequal lobes.
Abdominal neurosetae include two acicula with blunt, bent tips and two to three
crenulate capillaries some of which have abruptly tapered dps.
Type locality : Madagascar.
Records: Natal (27/32/i).
Distribution : Bay of Biscay (i) ; Madagascar (i).
Orbinia ntowroi Day, 1955
(fig. 23.3.h-j)
Orbinia monroi Day, 1955 : 409, fig. i e-h.
Length up to 100 mm. for 200 segments. About 50 thoracic setigers with branchiae
from the fifth. A row of about 1 1 foot papillae on posterior thoracic neuropodia
and about 30 stomach papillae over setigers 40-45 (fig. 23.3.h). Thoracic neurosetae
include four to five rows of flanged, bent but smooth hooks and a few crenulate
546 POLVCHAF.IA OF SOUTHERN .\FRU:\
capillaries. IntcM'iamal cinus (fig. ^3.3.1) well developed. Postsctal lobe of abdom-
inal notopodia with a chopper-shaped blade. Forked setae (fig. 23.3.J) with weakly
serrated shafts.
Type lociality : Knysna Lagoon, South .Africa.
Records: Cutpe (■24/23/e) ; Mocambique (26/32/i).
DisTRiBiTiON : Endemic.
HAPLOSCOLOPLOS Monro, 1933
Prostomium conical and pointed. Proboscis lobed. Buccal segment achaetous.
Branchiae on all except a few anterior segments. Thoracic seginents not markedly
flattened. Notopodia with a cirriform postsctal lobe bearing a bundle of crenulate
capillaries at its base. Thoracic neiiropodia with not more than two foot papillae.
No stomach papillae. Only crenulate capillaries in the thoracic neuropodia, there
being no hooks. Abdominal neuropodia bilobed and bear a bundle of crenulate
capillaries. An interramal cirrus may be present.
Type species : Scoloplos cylindrifer Ehlers, 1905.
Key to Species
1 An interramal cirrus ]:>rescnt on anterior abdominal segments (tig. 23.4.1') ■ H. fragilis
- No interramal cirrus .......... H. kerguelensis
Haploscoloplos cC. fragilis (Verrill, ifiys)
(fig. 23.4.e-f)
? ,l«//iOito;n(i/r(7i;(7c Wrrill, 1873: 59!!.
? Haploicolofilos fragiUs: Hartman, 1937: 271, ])1. Jj figs. 1-3.
Haploscoluplos c{.fragilii : l^ay, 1963: 416.
Length up to 20 mm. Thorax with 15-16 setigers. Setae are crenulate capillaries
in both rami throughout the body. Thoracic notopodia with a well marked post-
sctal lobe from the first foot onwards ; abdominal notopodia slender. Branchiae
from setiger 14-15 ; they are larger than the notopodia. An interramal cirrus
appears at the junction of the thorax and abdomen but decreases in size posteriorly.
Anterior thoracic ni-uropodia with a single postsctal lobe ; the last two thoracic
neuropodia (fig. 23. 4. e) with two to three postsctal lobes. \n the abdomen the
neuropodia are bilobed (fig. 23. 4. f) with at first two and later one ventral cirrus.
Records: C^ape (34/26/d).
Distribution : Doubtful - H. fragilis is recorded from the Atlantic coast of LI. S. A.
between Massachusetts and the Gulf of Mexico on sandy shores and from drcdgings,
but the South African specimens may be distinct.
ORBINIIDAE 547
Haploscoloplos kerguelensis (Mcintosh, 1885)
(fig. 23.4.a-d)
Scoloplos kerguelensis Mcintosh, 1885 : 355, pi. 43 figs. 6-8, pi. 22A fig. 19.
Haploscoloplos kerguelensis : Monro, 1936: 160; Hartman, 1957: 275, pi. 27 figs. 1-3; Day, 1961 :
477-
Length up to 20 mm. Thorax (fig. 23.4.3) with 10-14 slightly flattened segments
with branchiae from the ninth to fifteenth onwards, at first small but much larger
than the notopodia in the posterior abdomen. Notopodial postsetal lobes obvious
from the first foot. Thoracic neuropodia (fig. 23. 4. b) with a single long median
foot papilla. No stomach papillae. Abdominal notopodia (fig. 23.4.C) small and
tapered. No interramal cirrus. Abdominal neuropodia with unequal lobes. No
ventral cirrus. Thoracic and abdominal neurosetae are crenulatc capillaries (fig.
234-d).
Type locality : Kerguelen Island.
Records : Cape (from 32/16/d to 34/18/s and 34/22/d).
Distribution : Kerguelen (s, d) ; Magellan area (s) ; South Georgia (s) ;
Antarctica (s, d) ; Western Australia (e) ; ? ? India.
SCOLARICIA Eisig, 19 14
Prostomium conical and pointed. No eyes. Only the buccal segment achaetous.
Branchiae from the middle of the thorax onwards. Notopodia with a cirriform
postsetal lobe bearing a bundle of crenulate capillaries and sometimes a few forked
setae in the abdomen. Thoracic neuropodia as lateral ridges or lamellae which
may be notched and may bear a posterior foot papilla. Abdominal neuropodia
bilobed with a lamellar expansion at the base. Stomach papillae few or absent.
Thoracic neurosetae include rows of hooks and crenulatc capillaries. Abdominal
neurosetae are crenulate capillaries and flail setae (capillaries with abruptly tapered
tips).
Type SPECIES : Scolaricia typica Eisig, 1914.
Key to .Species
I Six to twelve stomach papillae. Each posterior abdominal neuropodium with a notched
inferior lamella ........... S. dubia
- Stomach papillae absent. Inferior lamellae of abdominal neuropodia not notched S. capensis
Scolaricia dubia (Day, 1955)
(fig- 234-g-l)
Orbinia dubia Day, 1955: 409, fig. i a-d.
Scolaricia dubia : Day, 1961 : 481.
Length up to 35 mm. for 100 segments. Nineteen thoracic setigers with branchiae
from the thirteenth (fig. 23. 4. g). A single median foot papilla increasing to three at
the end of the thorax (fig. 23. 4. h). Six to twelve stomach papillae at the junction
548
POI.VCHAETA OF SOUTHERN AFRICA
Fig. 2'5.4. HiiplnHnln/iliu l.irgiulcn\i\. (a) Lateral S'iew of anterior end. (b) Tenth foot.
(cj .\bdominal foot, (d) Crcnulatc capillary. Ha/ihuoloplo^ cf. fytigilis. (e) Fourteenth
foot, fr) F'irst abdominal foot. Scularicia ritibia. I g) Dorsal view of anterior end. (h) Last
thoracic foot, (i) .interior abdominal fi>ot. (j) Thoracic hook, (k) Posterior abdominal
foot, (l) Flail seta.
ORBINIIDAE 549
of thorax and abdomen. A large interramal cirrus (fig. 23. 4. i). Abdominal neuro-
podia with equal lobes but with no ventral cirrus. A blunt glandular lamella appears
below the ncuropodium at the beginning of the abdomen, later becomes notched
and may separate into two lamellae at the end (fig. 23. 4. k). Notosetae as crenulate
capillaries throughout, and forked setae absent. Thoracic neurosetae include three
rows of flanged, bent and serrated hooks (fig. 23.4.J) plus a posterior row of crenulate
capillaries. Abdominal neurosetae few, including crenulate capillaries and longer
flail setae with abruptly tapered tips and serrated shafts (fig. 23. 4. 1).
Type locality : Langebaan Lagoon, South .Africa.
Records : Cape (from 33/1 7/s to 34/23/s, d).
Distribution : Endemic.
Scolaricia capensis Day, 1961
(fig- 23.5.a-d)
Scolaricia capensis Day, 1961 : 480, fig. i j>-s.
Length up to 45 mm. for 1 1 2 segments. Seventeen thoracic sedgers with branchiae
from the fifteenth onwards. Thoracic neuropodia (fig. 23.5.3) faintly bilobed with
a median foot papilla. No stomach papillae. No interramal cirrus. Abdominal
neiu"opodia (fig. 23. 5. b) bilobed with a single inferior lamellar expansion. No
ventral cirrus. Thoracic neurosetae are blunt hooks (fig. 23. 5. c) having a few coarse
serrations and a posterior row of crenulate capillaries. Abdominal notosetae do not
include forked setae. Abdominal neurosetae include one to two crenulate capillaries
and one to two flail setae (fig. 23. 5. d).
Type locality : Agulhas Bank, South Africa.
Records: Cape (34/17/d, 34/18/s, 34/21/s, 34/25/s).
Distribution : Endemic.
SCOLOPLOS BlumviWt, 1828
Prostomium conical and pointed, with or without eyes. Proboscis lobed. Only
the buccal segment achaetous. Branchiae on all except a few anterior segments.
Notopodia with a cirriform postsetal lobe bearing a bundle of crenulate capillaries
and somedmcs a few forked setae in the abdomen. Thoracic neuropodia with
nought to three postsetal foot papillae. Stomach papillae usually absent. Thoracic
neurosetae include rows of hooks plus a few crenulate capillaries. Interramal cirrus
usually absent. Abdominal neuropodia bilobed. A ventral cirrus may be present.
Type species : Lumbrkus armiger }^i\x\\cT, 1776.
Key to Species
I Membraneous pockets between the abdominal neuropodia (fig. 23.5.!). 18 thoracic
setigers ............ S. marsupialis
- No membraneous pockets ........... 2
550 I'OI.VCIIAK TA OF SOITHERX Al-RItlA
2 Branchiae start on setigcr 6 .......... . 3
- Branchiae start on .setii;er 12 or later ......... 4
3 .\bdominal ncuropodia bilobeci ifig. 23.5.1111 ; abdominal neurosetae all with tapcretl tips
S.johnstonei
- .\bdoniinal ncuropodia unilobed (lig. 23. 6. c^ ; many abdominal nenrcjsctae with abrujith-
pointed tips ........... S. uniramus
4 Less than 22 thoracic sctigers. Thoracic neuropodial hooks serrated . . .5. arntiger
- Between 24 and 29 thoracic setigers. Neuropodial hooks not serrated . S. madagascariensis
Scoloplos niarsiipialis Southern, 1921
Scoloplos inarsupialis .Southern. 1921 : 632. pi. 27 fig. 19 a-g. text-fig. 18; Day, 1957: 96.
Lcn,G;th up tn 50 mm. with 210 segments. About 18 thoracic setigers (fig. 23.5.6)
with branchiae from the ninth to thirteenth onward. Thoracic ncuropodia with a
single foot papilla (fig. 23. 5. f) starting on setigcr 4 to 12. No stomach papillae.
Xeurosetae are mainly crenulate capillaries and the few hooks may be limited to
the first 10 feet or be entirely absent in juveniles. \Vhen present the hooks (fig.
23.5.J) are serrated, flanged and faintly recurxed. .\bdominal segments (fig. 23. 5. h)
have membraneous pockets between adjacent ncuropodia. Forked setae very few.
Interramal cirrus absent.
Type locality : C^hilka Lake, India.
Records: Mocambique (26'32/i).
Distribution: India (e, i) ; Gulf of Manaar.
Scoloplos (Leodamas) johnstonei D^.)', 1934
(fig- 23-5-k-o)
Scoloplos johmlotiei Day. 1934: 5B, fig. 11 a-e.
Length up to 35 mm. for 180 segments. Between 21 and 24 thoracic setigers with
branchiae from the si.xth (fig. 23. 5. 1). Thoracic ncuropodia (fig. 23.5.0) either lack
foot papillae entirely or ha\'e only one to two on the last two to three thoracic
segments. Abdominal ncuropodia (fig. 23. 5. m) bilobed with the outer ramus larger
than the inner ; inferior lamella absent. Interramal cirrus absent. Thoracic ncuro-
podia with three to five rows of hooks plus a few capillaries. Each hook (fig. 23. 5. n)
is slightly bent near the tip, lightly serrated and flanged, .\bdominal notosetac are
crenulate capillaries plus a few forked setae with equal prongs (fig. 23. 5. k). Abdom-
inal neurosetae are crenulate capillaries with normally tapered tips. The aciculum
has a blunt tip.
Type LOf:ALrrY : False Bay. South .\frica.
Records: Cape ifrom 32 18 i, s to 34 18 i) ; Natal (29/3 1 /i) ; Mocambique
(26(32 i).
DisTRiBLTio.v : Endemic.
ORBINIIDAE
55"
Fig. 23.5. Scolarkia capensis. (a) Fifieenth (thoracic) foot, (b) Posterior abdominal foot,
(c) Thoracic hook, (d) Flail seta. Scoloplos marsupialis (after Southern, 192 1). (e) Lateral
view of anterior end. (f) Fifth foot, (g) SLxteenth foot, (h) .Abdominal foot, (i) Three
abdominal segments showing membraneous pockets between neuropodia. (j) Thoracic hook.
Scoloplos johnstonei. (k) Forked seta, (l) Lateral view of anterior end. (m) .\bdominal foot.
(n) Thoracic hook, (o) Tenth (thoracic) foot.
552 POI.VCHAErA CM' SOUTHKRN A1'RIC:A
Scoloplos (Leodamas) unirainus Dav, 1961
(fig. 23.6.a-r)
Scolu/iloi {L.) tmiramus Day. i;|l'ii : 477, Hg. i g-o ; Day, 19G3 : (").}7,
Length up to 35 mm. for 100 segments. Twenty-one to twenty-two thoracic
sctigcrs with branchiae from the sixth. Postsctal lobe of the notopodium always
with a filamentous tip even on the abdomen. Thoracic neuropodia with a single
long median foot pajiilla and a long ventral cirrus from sctiger 18 onwards (fig.
23. 6. a). Xo interramal cirrus. Abdominal neuropodia each with a stout base and a
single (outer) ramus (fig. 23. 6. c). Juveniles with ventral cirri on first few abdominal
segments (fig. 23. 6. b). No expanded lamellae. Notosctac as crenulate capillaries
(fig. 23. 6. c) plus a few forked setae in the abdomen of juvenile s]Decimens but not
adults. Thoracic neurosetae include three to four rows ol bkuU, bent hooks lightly
serrated near the end (fig. 23. 6. d) plus a few fine capillaries on the last few segments.
Abdominal neurosetae include one to two fine crenulate capillaries, two to four long
stout setae witli poorly marked serrations and abruptly pointed tips (fig. 23. 6. f) and
one to two stout, projecting acicula.
Type locality : Agullias Bank, South Africa.
Reciords : Clape (from 34/18/s and 34/22/d to 34/25/s; Natal (29/31/s).
Distribution : Ende
nuc.
Scoloplos ntadagascariensis Fauvel, iQic)
(lig. 23.6.g-j)
Scolojilos tiioiiiiga^carunns I\aii\-cl, HMO : 433' pl- '7 ^'g^- 3i-86.
Length up to 120 mm. with o\er 250 segments. Between 24 and 30 thoracic
sctigcrs (fig. 23. 6. g) with branchiae from about the 22nd. Postsctal lobe of noto-
podium ininute on anterior segments but obvious from the tenth. Thoracic neuropodia
(fig. 23.6.11) without foot pa]iillae except sometimes on the last one to three. Neuro-
podial hooks in two rows accoinpanied by a very few capillaries ; they are quite
smooth and almost straight without serrations or a trace of a flange (fig. 23.6.J).
Forked setae present with unequal rami. No interramal cirrus. No \entral cirri on
abdominal neuropodia (fig. 23.6.1).
Type locality : Tulcar, Madagascar.
Records: Cape (33/1 8/s and 34/18/s) ; Natal (29/31/1,5) ; Mocambique (26/32/i
and 23/35/e).
Distribution: Madagascar (i) and tropical West Africa; ALituctania (s) Gulf;
of Guinea (s) ; Angola (s).
ORBINIIDAE
553
Fig. 23.6. Scoloplos uniramus. (a) Eighteenth (thoracic) foot, (b) Anterior abdominal foot,
(c) Posterior abdominal foot, (d) Thoracic hook, (e) Notopodial capillary, (f) Neuro-
podial capillary from the abdomen. Scoloplos madagnscarieruis. (g) Anterior end. (h)
Thoracic foot, (i) Abdominal foot, (j) Thoracic hook. Scoloplos armiger (modified from
Fauvel, 1927). (k) Thoracic foot, (l) Abdominal foot, (m) Thoracic hook, (n) Forked
seta.
554 I'Ol.VCHAETA OF SOUTHERN AI-RK:.\
Scoloplos armiger (MuUcr, 1776)
(fig. 23.6.k-n)
Lumbricwi armi ger '\{u\\er , 1776: 215.
Scoloplos armiger: Faiu'cl. 1927 ; 20, fig. 6 k-q.
Length up to 120 mm. for 200 segments. Prostomium pointed with subdermal
eyes. Sixteen to eighteen thoracic setigcrs with branchiae starling on the 12th-
15th. Thoracic neiiropodia (fig. 23. 6. k) with at least one foot papilla but two
on the last three to four segments. One to two stomach papillae at the junction
of the thorax and abdomen. Abdominal neuropodia (fig. 23. 6.1) bilobcd with the
inner lobe longer. No interramal cirrus or ventral cirrus. Notosctae as crcmdate
capillaries plus one to two forked setae (fig. 23. 6. n) on the abdomen. Thoracic
neurosetae include five rows of setae among which crcnulatc capillaries and short
blunt "hooks" arc mixed.
Type locality : Southern Norway.
Records: ? Cape (34/18/s and ? 34/22/d).
Distribution : Arctic (s, d, vd, abyssal) ; North and South Atlantic (i, s) ;
North Pacific from Japan to California (s) ; ? Indian Ocean.
PARAONIDAE
555
Family PARAONTOAE Ccrruti, 1909
Small thread-like worms up to 40 mm. long with numerous segments, somewhat
flattened anteriorly but rounded posteriorly. Prostomium conical with well devel-
oped nuchal shts and sometimes a median dorsal antenna and eye-spots. Palps
absent. Phaiynx soft and unarmed. Buccal segment reduced and fused to the
lower surface of the prostomium ; it is seldom visible dorsally. All subsequent
segments with small biramous parapodia reduced to a postsetal notopodial lobe
and a lateral ridge-shaped neuropodium. Cirriform dorsal branchiae start on
setigers 4 to 18 and continue over a variable number of anterior segments but are
absent posteriorly. Acicula absent. Setae are mainly smooth capillaries in both
rami but specialised setae may be present in either ramus posteriorly.
Records from southern Africa
Aedicira belgicae (Fauvel)
Aricidea capensis Day
Aricidea curviseta Day
Aricidea fauveti Hartman
Aricidea jejfreysi (Mcintosh) sensu Cerruti
Aricidea longobranchiata Day
Aricidea suecica simplex Day
Cirrophortis branchiatus Ehlcrs
Paraonis gracilis gracilis (Tauber)
as Aonides gracilis Tauber
Paraonis gracilis oculata Hartman
Paraonides lyra capensis Day
as Paraonis Ijra var. capensis Day .
Paraonides lyra lyra Southern .
? Paraonides sp. .
56Cd
51CS
56CS, — Xd
48Cd
56CS
51CS
55Ca
i5Cs,56Cd,— Nd
56Cd
2lCi
55Ca
44Ci
56Cd
56CS
REMARKS
Useful reviews of the family will be found in Cerruti (1909) and Hartman (1957).
The paraonids are tiny thread-like worms which burrow just below die surface of
sandy mud. They are non-selective deposit feeders and the gut is full of sand grains
ingested together with the detritus. For systematic purposes one of the most important
characters is the presence of a median antenna on the prostomium and it should be
noted that this is often broken off.
At the generic level the possession of a median antenna and the disposition of
speciahsed setae are the most important characters. At the specific level the shape
of the prostomium, the length of the antenna, the segmental position and number of
gills, the size of the postsetal lobe of the notopodium and the exact shapes of the
setae are useful.
Head structures. The median antenna varies in length but is easily broken off
leading to generic confusion. The eye-spots are never deeply pigmented and prob-
ably fade in alcohol. The shape of the head is reasonably constant but there has
556 POI.VCIHAKTA OF SOUTHERN AFRICA
been some uncertainty as to whether the head consists of a prostomiuin alone and
whether the buccal segment possesses setae or not. Accordintr to Hartman "the
first or second visible segment has biramons parajiodia provided with setae". After
careful examination of several species the view adopted here is that the first or buccal
segment is usually reduced and not visible dorsally. According to this view the
buccal segment lacks setae and is fused to the ventral surface of the prostomium so
that the head consists of a fused prostomium plus ]3eristomium. This appears to
be the case in Aricidea but in Ciirojihorus the posterior dorsal margin of the peristome
is \isible between the prostomium and the first setiger while the ventral part is
outlined by a shallow groove. In Parannis the mouth is further back with the \entral
surface of the first setiger forming the posterior lip.
Paxijioilia ami hranchiac. The notopodium is reduced to a bundle of setae and a
postsctal lobe. The latter is usually best developed in the anterior region and the
comparative length of the postsctal lobe and branchia is worth noting. In the
posterior region the postsctal lobe usually becomes a slender filament which may be
elongated on the last few segments. The neuropodium is seldom more than a lateral
ridge from which the setae arise but a minute postsctal papilla is occasionally present
in the anterior region.
The branchiae usually appear on setiger 4 or occasionally on setiger 5 but in
Paiaoriis gracilis they do not appear before setiger 7 and sometimes not before setiger
9. The branchiae are cylindo-conical projections which overlap in the mid-dorsal
line. The last few tend to have swollen bases and occasionally they have elongated
filiform tips. The number of branchiae varies from three to over 60 and this number
is somewhat variable within a single species, e.g. 3-14, 9-14, 18-24, 34^50-
Siiar. The majority of the setae in both rami are smooth capillaries. They are
curved and may ha\e llattened blades in the anterior region but are straight and
much finer in the posterior region. The specialised setae may be present in either
ramus, sometimes as early as the last few branchiae and sometimes not until the
last third of the body. They vary from setae with thick shafts and slender tips (and
thus very similar to the capillaries) to hooded hooks with a rounded guard or dehcate
tapering arista extending beyond the apex. Forked setae also occur.
Hartman (1957) recognises two genera Aricidea and Paraonis based on the presence
or absence of the median antenna. In each genus there are subgenera depending
on whether the specialised setae occur in the notopodia or neuro]5odia. These
distinctions arc clear cut and easy to observe and as some 30 species oi Aricidea have
been described it is suggested that the subgenera be raised to full generic rank.
The following generic key is adapted from Hartman (1957 p. 314)-
Key to Gener.\
1 Prostomium with a median dorsal antenna (fig. 24.1. j) ...... 2
- Prostomium without an antenna .......... 3
2 Specialised setae among the capillaries of posterior neuropodia . . ARICIDEA (p. 557)
- .Specialiiied setae among the capillaries of posterior notopodia . CIRROPHORUS (p. ^62)
- No specialised setae among the capillaries either in the notopodia or neuro]>odia
AEDICIRA (p. 5G3)
PARAONIDAE 557
3 Specialised setae among the capillaries of posterior notopodia . PARAONIDES (p. ^66)
- Specialised setae among the capillaries of posterior neuropodia . . PARAONIS {p. ^6^)
ARICIDEA Webster, 1879
Prostomium with a median antenna. Peristome reduced, achaetous and fused to
the ventral surface of the prostomium. All subsequent segments with two bundles
of setae. Branchiae from setiger 4 and number 10 to 60 or more pairs; they are
absent from posterior segments. Smooth capillaries present in both rami of all feet.
In addition the neuropodia of posterior segments have specialised setae which may
be pseudo-articulate, acicular or abruptly tapered, their distal ends often being
provided with a hood or deUcate tapered arista.
Type SPECIES : Aricideafragilis V^thstcr, 1879.
Key to Species
1 Specialised posterior neurosetae numerous, each with stout shaft abruptly tapered to a
slender blade ............. 2
- Specialised posterior neurosetae as a few (eight or less) sigmoid hooks often with a hood
or arista .............. 3
2 Specialised neurosetae with an incomplete joint at the junction of shaft and slender tip
A. fragilis*
- Specialised neurosetae with a kink but no joint at the junction of shaft and blade (fig. 24. i .e)
A. curviseta (p. 557)
3 Specialised neurosetae are sigmoid acicular hooks sometimes with a terminal filament but
no hood .............. 4
- Specialised neurosetae are hooks with a short or tapering hood or arista ... 5
4 Acicular hooks with a terminal filament in middle segments but plain posteriorly
A, suecica suecica*
- Acicular hooks without a terminal filament even in middle segments (fig. 24.1.1)
A. suecica simplex (p. 558)
5 Specialised neurosetae with a rounded spioniform hood covering the blunt ape.x (fig.
24. 1. m) A. Jeffreys! (p. 558)
- Specialised neurosetae with a delicate pointed hood or arista near the apex ... 6
6 Pointed hood or arista on the convex side of the apex (fig. 24.2. d). Posterior branchiae
longer and stoutei' than middle ones ...... A*fauveli (p. 560)
- Pointed hood or arista on the concave side of the apex (fig. 24.2. k) .... 7
7 Last few branchiae with swollen bases and greatly elongated tips (fig. 24.2.1) A. longobranchiata
(p. 560)
- Last few branchiae decrease in size ...... A, capensis (p. 562)
Aricidea curviseta Day, 1963
(fig. 24.i.a-e)
Aricidea cumsela Day, 1963a : 422, fig. 9 e-k.
Length up to 20 mm. for 100 segments. Prostomium (fig. 24.1.3) bluntly conical,
slightly longer than broad. Antenna short, not reaching tip of prostomium. .interior
segments six times broader than long, posterior ones about as broad as long. Thirty-
four to forty-four pairs of branchiae starting on setiger 4 ; anterior ones equal to
half the segmental width, posterior ones smaller. Postsetal lobes of notopodia (fig.
24.1.0) tapered and equal to one-third the length of the gill anteriorly but become
55ii ^()l,^■^;n.\K lA oi- souiherx Africa
\ei'y slender jjostciiorly. No visible postsetal lobes in the nenropodia. Anterior
notosetae and neurosetac are curved capillaries with stout shafts and tapering
blades. Posterior notosetae are a few slender capillaries (fig. 24.i.d). Posterior
neurosctae are all capillaries ; superior ones taper normally while inferior ones are
numercms rather short setae (fig. 24. i.e) having stout shafts and abruptly tapered
blades with a kink at the junction of the shaft with the blade. No articulation at
the junction.
Type locality : AguUias Bank, South Africa.
Records: Cape (34/23/s, d) ; Natal (29/31/d).
Distribution : ? Endemic.
Aricidea suecica simplex Day, 1963
(fig. 24.i.f-i)
Aricidea suecica simplex Day. 1963 : 364, fig. 3 a-b.
Body slender, about 12 mm. long for 80 segments. Prostoinium (fig. 24.i.f)
slightly longer than broad. Antenna short and club-shaped and does not reach the
tip of the prostomium. No eyes. Anterior segments six times as broad as long,
posterior ones a little longer. Branchiae from setiger 4 to about sctigcr 15 ; anterior
ones (fig. 24. 1. g) overlap their fellows in the mid-dorsal line but posterior ones are
smaller. Postsetal lobe of the notopodium slender and two-thirds the length of
the branchia in the anterior region ; posterior ones the same length. Anterior
notosetae and neurosctae are curved capillaries with tapered blades ; posterior
capillaries are straight and much finer. From setiger 24 tlie neurosctae include
six to ten stout, blunt, sigmoid, acicular setae (fig. 24.1.1) without any sign of a
terminal filament.
Type locality: West of Cape Town in 1240 metres.
Records: Cape (34/1 7/abyssal).
Distribution : No other records.
Aricidea jeffreysi Mcintosh 1879 {sensii Cerruli, 1909)*
(fig. 24.i.j-m)
Scokcolejns (?) jeffreysii Mcintosh, 1879: 506, pi. 45 figs. 13-14.
Aricidea jcffnysii : Ccrruti, 1909: 409, pi. 18 figs. 1-6, 9-18. 22-26, pi 19 figs. 28-42. Faiucl, 1927 :
75, fig. 25 a-e. Day, 19633 : 423.
Body about 15 mm. long by 0-4 mm. for too segments. Prostomium (fig. 24.1.J)
bluntly triangular or rounded anteriorly. Median antenna just projects beyond
anterior margin of prostomium. Eyes not seen. Anterior segments four times as
broad as long ; posterior segments a little longer. Branchiae from setiger 4 and
extend over about 14-18 segments; anterior ones (fig. 24.i.k) overlap their fellows
in the mid-dorsal line and are bliuitly cylindrical; posterior ones slightly shorter.
Postsetal lobes of notopodia well developed and tapered anteriorly, about half as
* Ih'-ic is some duiibl whether Cerniti's spcciinc-n from tlie Mediterranean is conspccific with Mi Intosh's
type.
PARAONIDAE
559
^^^i^
Fig. 24.1. Aricidea curviseta. (a) Anterior end. (b) Far posterior foot, (c) Branchiferoiis
foot, (d) Posterior notoseta. (e) Posterior neuroseta. Aricidea mecica simplex, (f) .interior
end. (g) Branchiferous foot, (h) Posterior foot, (i) .Specialised posterior neuroseta.
Aricidea jejfreysi. (j) Anterior end. (k) Branchiferous foot, (l) Posterior foot (m)
Specialised posterior neuroseta.
56o POLVCHAETA OF SOUTFIERX AFRICA
long as the branchiae ; posteriorly (fig. 24.1.I) they become very slender but remain
the same length. Minute postsctal papillae present on anterior ncuropodia. Anterior
setae are curved, tapering capillaries in both rami ; posteriorly they become straight
and slender and four to five hooks (fig. 24.i.m) appear in the neuropodia. Each
curves to a blunt apex which has a rounded spioniform guard.
Type locality : Greenland.
Records: Cape (34/18/s).
DiSTRiBUTio.N' : ? Da\is Straits (Greenland) ; ? North Carolina (s) ; ? Ireland ;
Mediterranean.*
Aricidea fauveli Hartman, 1957
(fig. .!4.2.a-d)
Aricidea fauveli Wiinman, 1957: 318.
Aricidea fragilis (non Webster) Fauvel, 1936: 65, figs. 6-7.
Body about 20 mm. long by 0-4 mm. for 100 segments. Prostomium (fig. 24. 2. a)
bluntly triangular, about as broad as long with indistinct eyes and a small median
antenna. Anterior region flattened with segments almost eight times as broad as
long ; posterior region rounded with slightly longer segments. Branchiae from
setiger 4 and extend over 20-24 segments. An average gill (fig. 24. 2. b) is equal to
two-thirds the segmental breadth but the posterior ones arc stouter basally and as
long as the segment is broad. Postsetal lobes of anterior notopodia are ^vcll developed
and half as long as the gill but they become slender posteriorly (fig. 24. 2. c). Anterior
setae are curved and tapered capillaries in both rami. Posterior setae are reduced
in number ; the notosetae consist of a few fine capillaries a little longer than the
notopodial lobes ; ncurosetae include a few capillaries similar to those in the
notopodia and about four curved unidentate hooks (fig. 24. 2. d) with a delicate
pointed hood or arista covering the convex side of the apex.
Type locality ; Morocco.
Records: Cape (34/18/d).
Distribution: Morocco (s, d) ; Tropical western Africa (s) ; Angola (s, d).
Aricidea longobranchiata Day, 1961
(fig. 24.2.e-k)
Aricidea longobranchiata Day, 1961 : 482, fig. 2 g-n.
Body (fig. 24. 2. e) vermiform and tapering. Length up to 20 mm. by o-8 mm. for
120 segments. Prostomium (fig. 24. 2. f) bluntly triangular, as broad as long. Xo
eyes. Median antenna very elongated, reaching setiger 5. Anterior region llattcncd
with segments six to eight times as broad as long ; posterior segments only slightly
PARAONIDAE
56-
Fig. 24.2. Africidea fauveli. (a) Anterior end. (b) Branchiferous foot, (c) Posterior foot.
(d) Specialised posterior neuroseta. Arkidea longobranchiata. (e) Entire worm (six times
natural size), (f) Anterior end. (g) Anterior branchiferous foot (tenth), (h) Notopodial
capillary-, (i) Posterior branchiferous foot (24th). (j) Far posterior foot, (k) Specialised
posterior neuroseta. Arkidea caperuis. (l) Anterior end. (m) Branchiferous foot, (n) Far
posterior foot, (o) Specialised posterior neuroseta.
jfij poi.vc:haeia of southkrx afric:a
broader than long. Br.inchiae from sctigcr 4 and extend over i8--:;2 segments.
Anterior ones (fig. 24. 2. g) arc uniformly tapered and just o\erlap in the mid-dorsal
line but the last three pairs (fig. 24.2.1) have swollen bases and long slender tips
longer than the width of the segment. Postsetal lobes of anterior segments well
developed but posterior ones (fig. 24.2.J) very slender. Anterior setae are curved
capillaries with rather broad blades in both rami. Posteriorly the capillaries (fig.
24.2.11) became fine and straight and four to five hooks (fig. 24. 2. k) appear in the
neuropodia. Each has a unidentate curved apex with a fine tapered guard or arista
arising from tlie conca\e side of the shaft.
Type locality : Off Saldanha Bay, southern Africa.
Records: South \Vest Africa (26/14/d and 26; 15/3); Clape (from 32/18/s and
33 17 s, d to 36 21 d and 34;25/s).
Distribution : Southern African endemic.
Aricidea capensis Day, 1961
(fig. 24.2.I-0)
Aricidea capensis Day. 1961 : 481, fig. 2 a-f.
A thrcad-Uke species measuring 10 mm. by 0-2 mm. for more than 75 segments.
Prostomium (fig. 24. 2. 1) conical, 1-5 times longer than broad. Xo eyes. Antenna
slightly longer than the prostomium and obscurely annulated near the tips. Anterior
segments three times as broad as long : posterior ones hardly broader than long.
Branchiae ifig. 24. 2. m) from setiger 4 and extend over 14 segments; most of them
are equal to two-thirds the segmental breadth but the last two to three pairs are
smaller. Postsetal lobes of anterior notopodia are one-third the length of the gills
but posterior ones (fig. 24.2.11) are smaller. Anterior setae are curved, broad-bladed
capillaries in both rami. Posterior capillaries are fine and straight, .^bout four
curved hooks (fig. 24.2.0) appear in the posterior neuropodia ; each is bidentatc
with a minute secondary tooth above the main terminal one and has a long tapering
arista arising from the conca\-e side of the shaft just below the apex.
Type loc.'vlit^' : Mossel Bay, South Africa.
Records: Clapc (34/22/s) ; Natal (30/30/s).
Distribution : Only two records.
CIRROPHORUS EhlcK, 190S
Prostomium with a median antenna. Peristome reduced, fused to the \entral
surface of the prostomium but occasionally visible dorsally as a fillet between the
prostomium and the first setiger. AU subsequent segments biramous. Xotopodiuni
reduced to a cirriform postsetal lobe. Neuropodium as a low lateral ridge. Branchiae
from setiger 4 or 5 and extend o\er 15-33 segments. Smooth capillaries present in
both rami of all feet ; in addition the middle and posterior notopodia bear acicular
or forked setae. Xo specialised neurosetae.
Type species : Cirrophorus branchiatus Ehlers, 1908.
PARAONIDAE 563
Cirrophorus branchiatus Ehlers, 1908
(fig. 24.3.a-e)
Cirrophorus branchiatus Ehler, 1908: 124, pi. 17 figs. 5-9; Day, 19633: 423, fig. 9 l-o.
Body about 25 mm. long by 0-4 mm. for 120 segments. Prostomium (fig. 24.3.3)
bluntly conical, slightly longer than broad. Eyes absent. Median antenna short
and stout, less than half the prostomial length. Peristome reduced ; it isjust visible
dorsally between the prostomium and the first setiger but the main part is fused to
the ventral surface of the prostomium. Anterior segments somewhat flattened,
each three to four times as broad as long. Posterior region with segments slightly
broader than long and separated by marked intersegmental constrictions (fig. 24. 3. c).
Branchiae from setiger 5 and extend over 20-25 segments ; each is a cylindro-conical
lobe (fig. 24. 3. b) equal to two-thirds the segmental breadth. The last two to three
pairs are shorter. Postsetal lobes of the notopodia well developed anteriorly, very
small in the middle of the body but longer posteriorly. Four pairs of cirriform
projections at the posterior end (fig. 24. 3. e) some of which may be the notopodial
lobes of rudimentary segments.
Anterior setae up to segment 12 are fine capillaries in both rami. From setiger
13, one to two heavy spines appear in the notopodia and the number of capillaries
is reduced. Each spine (fig. 24. 3. d) is straight and bluntly pointed with a very fine
curved filament arising some distance below the apex. No specialised setae appear
in posterior neuropodia but the capillaries are twice as long as those in the notopodia.
Type locality : Agulhas Bank, South Africa.
Records: Cape (32/17/d and 35/20/d) ; Natal (29/31/d).
Distribution: ? CaUfornia ; North Carolina (s).
AEDICIRA Hartman, 1957
Prostomium broadly rounded or flattened anteriorly with a median dorsal antenna
which is usually simple but may be branched. Peristome fused to the prostomium.
All subsequent segments biramous with two bundles of setae. Notopodia reduced
to small postsetal lobes ; neuropodia are mere lateral ridges. Cirriform branchiae
appear on setiger 4 and extend over 12 to 60 segments. The setae are simple capill-
aries in both rami throughout the body and specialised setae are absent.
Type species : Aricidea [Aedicira) pacijica Hartman, 1944.
Aedicira belgicae (Fauvel, 1936)
(fig. 24.3.f-j)
Paraonis belgicae Fauvel, 1963a: 29.
Aricidea belgicae: Monro, 1939: 127, fig. 16 a-b.
Aricidea (Aedicira) belgicae: Hartman, 1957: 327.
Aedicira belgicae : Day, 1936a: 424.
Body about 20 mm. long and i mm. broad for 100 segments. Prostomium (fig.
24. 3. f) broadly rounded anteriorly, about as broad as long with pale indistinct eyes,
well marked nuchal slits and a short stout median antenna which does not reach
5fH
?oLvt:ir.\E'rA or southern airkja
Fi'i. 1^4.3. Cirrojihiniis branchialw,. (a) Anterior end. (u) Brantliifenms foot, (c) Lateral
view of four posterior segments, (d) Xotopodial spine, (e) Posterior end and pygidium.
Afdicira belgkae. (f) Anterior end. (g) Branchiferous foot, (h) Posterior neuroseta.
fi) Posterior notoseta. (jj Posterior foot. Paraonis gracilis oculata. (k) Branchiferous foot.
(Lj Entire worm (seven times natural size), (m) Head, (n) Specialised posterior neuroseta,
(o) Posterior foot.
PARAOXIDAE 565
the prostomial margin. Peristome reduced and fused to the ventral surface of the
prostomium and not visible dorsally. Anterior region somewhat flattened with the
segments six times broader than long ; posterior region rounded with the segments
about twice as broad as long. Branchiae appear on setiger 4 and extend over 13-24
segments. An average gill (fig. 24. 3. g) is a stout, cylindro-conical organ as long as
two-thirds the segmental width but the last few are shorter with swollen bases and
short filamentous tips. In the anterior region the postsetal lobe of the notopodium.
is well developed, tapered and half the length of the gill. Posteriorly (fig. 24.3.J)
it remains the same length but becomes very slender and filamentous. Anterior
setae are numerous curved and tapered capillaries in both ranu. Posteriorly the
number of setae decreases but no specialised setae appear. The notopodial capillaries
are very fine ; the neuropodial ones are considerably longer, and most of them are
equal to the segmental breadth.
Type locality : Antarctic regions.
Records: Cape (32/17/d and 34/18/s).
Distribution: Subantarctic ; North Carolina (d).
PARAONIS Gruhe, 1878
Prostomium conical. No median antenna. Eyes present or absent. Peristome
usually reduced and fused to the ventral surface of the prostomium. All subsequent
segments biramous but the parapodia are poorly developed ; notopodia are repre-
sented by a small cirriform postsetal lobe, and neuropodia by mere Literal ridges
from which the setae arise. Dorsal branchiae start on the fourth to nineteenth
setiger and number three to 36 pairs but are absent posteriorly. Notosetae are
smooth capillaries throughout. Neurosctac include capillaries plus speciaUsed setae
in posterior segments.
Type species : Paraonis tenera Grube, 1B78 (? = Aonides gracilis Tauber, 1879).
Key to Species
1 Specialised neurosetae have a blunt, curved apex with a ventral guard. Branchiae
foliaceous ........... P.fulgens*
— Specialised neurosetae are unidentate acictilar hooks without hoods. Branchiae cirriform . 2
2 Prostomium with eyes (fig. 24.3.m) ...... P. gracilis oculata
— Prostomium without eyes ........ P. gracilis gracilis
Paraonis gracilis oculata Hartman, 1957
(fig. 24.3.k-o)
Paraonis gracilis oculata "HanTixan, 1957: 331, pi. 44 figs. 1-3; Day, 1963: 363.
Body (fig. 24.3.I) up to 17 mm. long by 0-3 mm. for more than 80 segments.
Prostomium (fig. 24. 3. m) broader than long with a pair of pale eyes. Anterior
segments about three to four times as broad as long ; posterior segments more
rounded, not more than three times as broad as long. Branchiae start on setiger 6-12
566 POI.VOHAEIA OF SOUTHKRN ArRIC;.\
and niiinbcr abciut lO to 20 pairs. An axnage gill (fig. 24. 3. k) is cirriform and just
overlaps its fellow in the mid-dorsal line ; the last few are slightly smaller. Post-
setal lobes of anterior notopodia inconspicuous, posterior ones (fig. 24.3.0) small.
Both rami of all parapodia have slender capillaries whose length is about half the
segmental breadth. In addition the posterior neuropodia bear about four sigmoid
acicular setae (fig. 24.3.0) w ith curved tips but lack a guard or arista.
Type locality : Dredged off California.
Records: Cape (34/1 7/abyssal).
Distribution: C^alilbrnia (d).
Paraonis gracilis gracilis (Tauber 1879)
(fig. 24.4.a-b)
Aonides gnuHii T-AuhcT. 1879: 115.
Paraoriii gracilis gracilis : Hartman, 1957 : 330. pi. 44 figs. 4-5 ; Day, 19633 : 425.
Body thread-like, about 20 mm. long by 0-3 mm. broad for 100 segments. Pro-
stomium (fig. 24. 4. a) twice as long as broad. No eyes. Anterior segments slightly
flattened and three times broader tJian long ; posterior region rounded with segments
as broad as long. Branchiae from setiger 6-1 1 onwards and number 3-14 pairs
(usually 1 1) Each is a cirriform organ equal to two-thirds the segmental breadth ;
the last few are a little smaller. Postsetal lobes of the notopodia are inconspicuous
and less than a quarter the lengdi of the gill. Both rami of all parapodia have
slender capillaries equal to half the segmental breadth. In addition the posterior
neuropodia bear two to four sigmoid acicular hooks (fig. 24. 4. b) with a narrow guard
(in South African specimens at least).
Type locality : North Sea.
Records: Cape (32/17/d, 35/20/d, 34/26/d).
Distribution; Closmopohtan (s, d).
PARAONIDES Cvmiu, 1909
(including FARADO.hElS Hartman, 1965)
Prostomium conical. No median antenna. Eyes present or absent. Peristome
greatly reduced and fused to the ventral surface of the prostomium. All subsequent
segments bear biramous but greatly reduced parapodia. Notopodium represented
by a small cirriform postsetal lobe. Neuropodium as an indistinct lateral ridge from
which the setae arise. Cirriform dorsal branchiae start on setiger 4 and extend
over about 10-20 segments. Notosetae are fine capillaries with the addition of a
few specialised setae from the branchial region onwards. Neurosetae are slender
capillaries and there arc no specialised neurosetae.
Type SPECIES : Paiaonis {Paraonidei) riedpnlilatui Cciruu, 1909.
PARAONIDAE
567
d 1 e
Fig. 24.4. Paraon'is gracilis gracilis, (a) Head, (b) Posterior neuropodial hook. Paraonides
lyra lyra. (c) Anterior end. (d) Notopodial forked seta, (e) Notopodial capillar^-.
(f) Branchiferous foot, (c.) Posterior foot. Paraonides lyra capensis. (h) Anterior end.
(i) Branchiferous foot, (j) Notopodial forked seta, (k) Posterior foot.
568 l'OLVC;ilAl.TA OF SOUIUKRN AFRICA
Key to Species
1 Modified iiotosctae arc short, stout, broatl-wiiigcd capillaries . . .P. neapolitana*
- Modified setae are forked setae .......... 2
2 Postsetal lobe of anterior notopodia one-tliird tlic length of the branchia. Proslomium as
broad as long ........... P. lyra lyra
— Postsetal lobe of anterior notopodia are minute, inconspicuous pajiillac. Prostoniium
longer tlian broad ......... P. lyra capensis
Paraonides lyra lyra (Southern, 1914)
(fig- ■^4-4-c-g)
Paraonis (Paraomdc^) lyra Southern, 1914: 94, fig. 22 a-g; Fauvcl. 1927 ; 72. fig. 24 a-f.
Paraonides lyra lyra : Day, 1963a ; 425.
Body thread-like, up to 20 mm. long by 0-2 mm. broad for luo segments. Pro-
stoniium (fig. 24. 4. c) broadly triangular, about as broad as long. No eyes. Anterior
segments twice as broad as long, posterior ones about as long as broad. Branchiae
from setiger 4 and number 8-14 pairs. Each is a digitiform lobe which overlaps its
fellow in the mid-dorsal line. Postsetal lobe of the nolopodium (fig. 2i.4.f) well
developed in the anterior region and about onc-quartir the length ol the gill ;
posteriorly (fig. 24. 4. g) they become inconspicuous. Both rami of all parapodia
contain fine capillary setae (fig. 24. 4. e). In addition the notopodia from the last
few branchiferous segments onward contain one to two short forked setae (fig. 24. 4. d)
with the longer limb spinulose on the inner margin. No specialised setae in the
neuropodia and the capillaries usually exceed the segmental width.
Type LOCALITY : \Vestern Ireland.
Records: Clapc (33/1 7/d).
Distribution; Sweden; Ireland (s).
Paraonides lyra capensis (Day, 195'))
(fig. 24.4.h-k)
Paraonh lyra var. ca/ionii Day, 1955 : 417.
Body thread-like, about 10 mm. long by o-i mm. broad for 80 segments. Pro-
stoniium (fig. 24. 4. h) conical, longer than broad. No eyes. Anterior segments
twice as broad as long, posterior ones as long as broad. Branchiae from setiger 4
and extend over 10-13 segments. Each is a digitiform lobe whose length equals
two-thirds the segmental breadth. Postsetal lobe of the notopodium (fig. 24.4.1)
minute and not visible before setiger 8 and sometimes only found in the posterior
region. Fine capillary setae present in both rami of the parapodia throughout the
body. In addition there are two to three short forked setae (fig. 24.4.J) with the
PARAONIDAE 569
longer limb spinulose on the inner margin. These 'appear in the notopodia of the
last few branchifcrous segments and persist in later segments (fig. 24.4.11). No
specialised setae in the neuropodia and the ncuropodial capillaries are shorter than
the segmental width.
Type locality : Knysna estuary, South Africa.
Records: Cape (36/2 i/d, 34/23/e, 34/26/d).
Distribution : South African endemic.
570 I'OI.VCHAETA OF SOUTHKRN A1RIC:A
Family OPHELIIDAE Malmgren, 1867
Body fusiform in shape with relatively few segments and often grooved \entrally
and laterally. Prostomium a tapered cone without appendages but with a pair of
evaginable nuchal organs and a pair of subdermal eyes. Proboscis unarmed.
Mouth a transverse sht at the level of setiger i. Parapodia biramous but not well
developed ; they have minute setigerous lobes and lack dorsal cirri and often
ventral ones as well. Cirriform branchiae may be present above the notopodia
and lateral eyes may be present between the parapodia. Setae are simple capillaries
throughout. Pygidium often prolonged and tubular and usually provided with
numerous anal cirri.
Records from southern Africa
Arinandia intermedia Fau\el
ArmatiJia leplocinis Grube
Armandia langicaudala (CauUery)
Ophelia africana Tebble .
Ophelia agulhana Day
Ophelia anomala Day
Ophelia capensis Kirkegaard
Ophelia peresi Bellan and Picard
Ophelia roscojfensis Augener
Ophelina acumitiata Oersted
as Ammotrypane aiilogaster Rathkc
Polyophthalmus picliis (Dujardin)
as Polyuphlhalmui papillatus Treadw
Travisia concinna (Kinbcrg)
as Dindjmene concinna Kinberg
Travisia forbesii Johnston
ell
26.^1, 45X1, fiiCls
4oNi, 44C;i, 5iC;s
4oNi
42CS, 51CS
5 ' C'S
5iCs
48CS, 51CS
5 7 Ms
? 56CS
?5iCsd, — Xsd
32NdCd
4oNi, — Ms
38Ai
3Cs
i5Csd, 51CS
REMARKS
The opheliids are fusiform worms which burrow head downwards in sand or
mud. The ventrum is grooved and die respiratory current is brought don by peristal-
tic action and escapes along the lateral grooves in which the gills arc situated. The
gut is often full of sand grains ingested along with the organic matter in the substrate.
Pulyophthalmus is found on muddy shores in all warm seas ; Ophelia and Armandia
are more common in dredging on fairly clean sand while Travisia lives on muddy
bottoms in deep water.
A brief review of the family will be found in Fauvel (1927). Slop-Bowitz (1945a)
gives a key to the several Norwegian species including five species o[ Ophelina. Tebble
(1953) gives a useful review of the known species of the genus Ophelia.
The whole family is well defined. The dilferent genera are easily recognised by
the development of the ventral groove, the nature of the pygidium and the presence
or absence of brancliiae and lateral eye-spots. Specific difTerences are based on
OPHELIIDAE 571
further details of the same structures plus the number of segments and the disposition
of the gills.
The ventral groove. In all genera except Travisia the longitudinal ventral muscles
are very strongly developed so that they form ventro-lateral ridges. In consequence
the ventrum between these ridges forms a ventral groove which may extend the whole
length of the body or be restricted to the posterior half. Often a pair of lateral
grooves develop just above the longitudinal muscles in which the parapodia and
branchiae are protected as the worm burrows through the sand.
Segmentation, parapodia and branchiae. The whole body is fusiform and interseg-
mental constrictions are poorly marked. Moreover each segment is annulated so
that the easiest way of counting the segments is by noting the number of bundles
of setae. It should be noted that the setae of the first segment are small and easily
overlooked and the several species possess a few achaetous posterior segments in
front of the pygidium. The total number of segments varies between 30 and 60
and is surprisingly constant for each species, at least in the genera Armandia and
Ophelia.
The parapodia though biramous, are small and the shapes of the parapodial lobes
seldom provide characters of importance. The setae too, are surprisingly uniform
though their development on the last few segments may be of systematic value.
The branchiae with few exceptions are cylindrical tapering projections which
arise postero-dorsal to the notosetae. They are absent in Polyophthalmus, Tachytrypane
and in a few species oi Ophelia and Ophelina but in most species they are well developed
and their segmental distribution is very constant. As shown by Tebble (1953) the
various species of Ophelia may be classified on the arrangement of the branchiae and
the total number of segments.
Lateral eye-spots occur in both Armandia and Polyophthalmus. They are restricted
to a certain number of segments in the middle of the body and occur at segmental
intervals between the parapodia.
Key to Genera
1 Ventral groove at least in the posterior part of the body. No segmental swellings above
and below the parapodia (fig. 25.1.C) ......... 2
- Ventral groove absent throughout. Body stout with segmental swellings above and below
the posterior parapodia (fig. 25.1.J) ...... TRAVISIA (p. 575)
2 A ventral groove along the whole body. Gills present from setiger 2 or entirely absent . 3
- A ventral groove posteriorly from setiger 7. Gills absent before setiger 8 OPHELIA (p. 571)
3 Gills present from setiger 2 onwards ......... 4
- Gills entirely absent ............ 5
4 Eye-spots present between the parapodia of middle segments . . ARMANDIA ip. ^•j&f
- Eye-spots absent from body ........ OPHELINA 1 p. 579)
5 Eye-spots present between the parapodia of middle segments. Surface normal
POLYOPHTHALMUS (p. 579)
- Eye-spots absent from body. Surface tough and cartilaginous . TACHYTRYPANE* (p. 580)
OPHELIA Savigny, 1818
Body fusiform with an anterior swollen region without grooves and a posterior
tapered one with a ventral groove. Prostomium a tapered cone. Segments with
572 roLVCiHAETA OT SOUIIIERN AI KItiA
superficial aniuilalions and often microscopic pits in the lateral walls of the branchial
region. Branchiae may be entirely absent or present over most of the segments
from setiger 8 or lo onwards. Both rami of the parapodia reduced to inconspicuous
lobes bearing a bundle of simple capillaries. A papilla between the parapodial
rami. P)gidium with two stout \enlral lobes and several small anal papillae
dorsally.
Type SPECIES : Ophelia bkoniis Svixigny, 1818.
Key to Species
Branchiae absent. Bndy with 26 setigers IfiE;. 25.1.3)
Branchiae present. Body with more than 26 setigers
Eight anterior abranchiate segments. A total of 32 setigers
Nine anterior abranchiate setigers. More than 32 setigers .
Ten anterior abranchiate setigers . . . . .
Thirty-si.x setigers .......
Thirty-nine setigers .......
A total of 29 setigers including 14-15 branchiferous .
A total of 32 setigers including 17-19 branciiiferous .
O. anojnala ip. 572)
2
O. roscoffensis (p. 572)
3
4
O. agulhana Ip. 573)
O. ajricana (p. 575)
O. peresi (p. 573)
O capensis (p. 573)
Ophelia anotnala Day, 1961
(fiff.
.i.a-b)
Opiklia anomnla Day, 1961 : 515, fig. lob.
Length up to 40 mm. Body (fig. 25.1. a) of 26 setigers with a doubtful achaetous
preanal. No branchiae at all. A constriction behind setiger 2. Notosetae short
and not obviously longer than ncurosetae. \'entral groove well marked from setiger 7.
Nephridiopores from setiger 11 to 14. No pits on lateral body \\all. Crimped
dorso-lateral ridges (fig. 25.1.!:)) from setiger 24 to pygidium which has two stout
ventral cirri and a dorsal arc of 1G-18 small ones.
Type locality : False Bay, South Africa.
Records: Cape (32/i8;s, 34,i8;s, ss/j^/s).
Distribution: Endemic.
Ophelia roscoffensis .Augener, 19 10
(fig. 23.1.ij
Ophelia roicnjfeiuis Augener, 1910: 237; Tebble. 1952: 565, fig. 5.
Body cigar-shaped, up to 51 mtn. long with a total of 32 setigers of which the first
eight are abranchiate, 23 bear slender branchiae and the last setiger is again
abranchiate. One doubtful achaetous preanal. Branchial fenestrations absent.
Setigers 12 to 20 with nephridiopores. Setigers 29 to 32 abruptly tapered with
crimped dorso-lateral ridges (fig. 25.1.1). Pygidium with an arc of 12 small anal
cirri and two stout ventral ones. Notosetae twice as long as ncurosetae or even
more on the last few setigers. (.South Afrit an specimens are doubtfiillv assigned to
OPHELIIDAE 573
this species as they have only 20-2 1 pairs of branchiae and three to four posterior
abranchiate setigers.)
Type locality : Roscoff, France.
Records; ? Cape (34.22.S).
Distribution : Enghsh Channel.
Ophelia peresi Bellan & Picard, 1 965
O/i/if /w /if r«i Bellan and Picard, 1965: 295, figs. 1-2.
Length 10-12 mm. Body of 29 setigers and 3 indistinct and doubtful achaetous
preanals. The first 10 or 11 are abranchiate, the next 14-15 bear short branchiae
and the last four setigers are again abranchiate. Branchial fenestrations absent.
Nephridiopores not seen. Prominent dorso-Iatcral ridges extend from setigcr 20 to
unite on the 27th setiger (2nd posterior abrancliiate setiger) and continue to the base
of the pygidium. Pygidium with a dorsal arc of 12 small anal papillae and two large
elongated ventral ones. Differs from 0. bicornis by the possession of the dorso-lateral
ridges on the last few segments and in having 4 not 7 posterior abranchiate setigers.
Type locality : Tulear, Madagascar.
Records : Madagascar (s).
Ophelia capensis Kirkegaard, 1959
(fig. 25.I.C-f)
Ophelia capensis Kirkegaard, 1959: 45, fig. 8; Day, 1961 : 514.
Length up to 50 mm. Body of 36-37 setigers including 9 (or occasionally 10)
abranchiate setigers, 17-19 branchiferous segments, three to four posterior abranch-
iate setigers and one to two achaetous preanals. Branchiae with a flattened sac-hke
basal projection (fig. 25.i.e). Nephridiopores on setigers 12-17. Notosetae (fig.
25. 1. f) in middle of body three times as long as neurosetae. Lateral swellings start
on setigcr 26 and condnue to the anus which has a dorsal arc of 12 small papillae
and two long stout ventral ones (fig. 25.i.d).
Type locality : In 50 metres off Table Bay, South Africa.
Records: Cape (33/18/s, 34/18:5, 34/23/s, and 33/27/s).
Distribution : Endemic.
Ophelia agulhana Day, 1961
(fig- 25-i-g)
Ophelia agulhana Day, 1961 : 513, fig. 10 a.
Length up to 50 mm. Body of 36-37 setigers including nine (or occasionally 10)
anterior abranchiate segments, 23-26 branchiferous segments, one to three posterior
abranchiate setigers and a doubtful achaetous preanal segment in front of the
574
I'Ol.VCHAETA OK SOUTHERN AFRICA
Fig. 25.1. Ophelia anomala. (a) Entire worm (twice life size), (b) Dorsal view of posterior
end. Ophelia capemis. (c) Entire worm (four times life size), (d) Posterior end. (e) Para-
podium, (f) Capillary seta. Ophelia agulhana. (g) Posterior end. Ophelia africana. (n)
Posterior end. Ophelia cf. roscoffensis. (i) Posterior end. Traiiiia forbesii. (j) Entire worm
(twice life size), (k) Anterior parapodium. (l) Posterior parapodium. (m) Capillary seta.
OPHELIIDAE 575
pygidium. Branchiferous segments with rows of pits in the lateral body wall. Branch-
iae irregularly annulated. Notosetae twice as long as neurosetae in the branchiferous
region. Nephridial pores on setigers 12-17. Crimped dorso-lateral ridges from
setiger 33 to pygidium which has two stout ventral cirri and a dorsal arc of 15 small
ones (fig. 25.i.g).
Type locality : False Bay, South Africa.
Records: Cape (32/17/d, 32/18/s, 33/17/d and 34/18/s).
Distribution : Endemic.
Ophelia africana Tebble, 1953
(fig. 25.1.h)
Ophelia africana Tebble, 1953 : 365, fig. i ; Day, 1961 : 513.
Length up to 70 mm. Body of 39-40 setigers including nine anterior abranchiate
setigers, 27-28 branchiferous segments, two to three posterior abranchiate setigers
and one to two achaetous preanals. Adults with rows of pits in body wall of branchi-
ferous segments. Branchiae irregularly annulated. Posterior setae short. Nephridio-
pores on setigers 11-16. Lateral swelhngs on setigers 36-39. Preanals and dorsal
part of pygidium crimped (fig. 25.i.h). Two short, stout ovoid papillae below the
anus and an arc of 10-12 small papillae above it.
Type locality : Table Bay, South Africa.
Records: Cape (33/1 8/s).
Distribution : A single record.
77?^ F/SM Johnston, 1840
Body fusiform without a ventral groove. Prostomium conical. Cirriform branchi-
ae from setiger 2 to the last few. Dorsal and ventral rami of parapodia both reduced
to a bundle of simple capillaries. Posterior setigers with cushion-Uke sweUings above
and below the setigerous bundles. No ventral cirri. A lateral organ (pit) between
the parapodial rami. Anus encircled by blunt lobes.
Type species : Travisiajorbesii ]ohns\.on, 1840.
Key to Species
I Twenty-three to twenty-nine setigers. Surface reticulate and sandy . . T. forbesii
— Thirty-five setigers. Surface granular with papillae ..... T, concinna
Travisia forbesii Johnston, 1840
(fig. 25.i.j-m)
TVauisia Forifsii Johnston, 1840 : 273; Fauvel, 1927: 138, fig. 48 g-k.
Length up to 30 mm. Body (fig. 25.1.J) short and fusiform with 23-29 setigers,
the anterior ones rounded and triannulate, the more posterior ones more rectangular
and rather telescoped with the ventral surface flat or faintly grooved. Branchiae
57« POI.VC:IIAETA OF SOL IHERX AFRU:A
from sctigcr ■:; to tin- last two. Xcphridioporcs from scligcr 3 to 14. Anterior para-
podia (fig. 25.i.kl without projections. Posterior parapodia (fig. 25.1.I) with
swellings representing dorsal and ventral cirri. Setae are fine hispid capillaries
(fig. 25. 1. m) throughout. Anus encircled by about eight stout papillae.
Type locality : Firth of C;lyde, Scotland.
Rfx.ords : Clape (34/18 s and 33'25's).
Distribution : .Arctic ; northern Atlantic from Norway (i, s) and Greenland
(i, s, d, abyssal) to the English Channel ; Behring Sea ; N.W. Japan and Okhotsk
Sea.
Travisia concinna (Kinberg, 1866)
Dindyiucnf toncinna Kinberg, 1866; 66, pi. 25 fig. j.
Tniiisia cuiu'innti : Hartm.in, 194B : 113.
A total of 35 setigers. Surface of body granular with fine papillae. Branchiae
from sctiger 2 to the last (33 in all). Anus lobed. Fleshy lobes above the notopodia
from setiger 17 and below tiic neurosetae from sctiger 18. They increase in size
posteriorly.
Type loc;ality : Algoa Bay, South Africa.
Records: Cape (33'25/s).
Distribution' : Known only from the orie;inal record.
ARMANDIA Filippi, 1861
Body elongated, pointed anteriorly and not di\ided into regions. A deep ventral
groove and a pair of lateral grooves along the whole length. Prostomium conical
with two subdermal eyes and two evaginable nuchal organs. Proboscis more or less
lobed. Segments annulated and lateral eyes present from about sctiger 2 onwards.
Parapodia with a presctal lobe, a short \entral cirrus and two bundles of capillary
setae. Pygidium tubular with a long internal \cntral cirrus and a semi-circle of
short dorsal cirri.
Type species ; Armandia cinliosa V\\\\>\)\, 1861.
Kev to Species
1 Presctal lobe of parapodia very long (fig. 25.2. a). Thirly to thirty-two setigers with gills
to the last .......... A, longicaiidata
- Presetal lobe of ])arapodia always short (lig. 2j.2.e) .......
2 Twenty-seven to twenty-nine setigers. Two or more posterior abranchiate setigers . . 3
- Thirty-three to thirty-seven setigers. Gills extend to the last setiger . . A. leptocirris
3 'Fhrcc to four posterior abranchiate setigers. Pygidial funnel pale with 12-20 anal cirri
A. intermedia
- Two postciinr abranchiate setigers. Pygidial funnel brown with si.\ anal cirri A. tnelanitra*
OPHELIIDAE 577
Armandia longicaudata (Caullerx', 1944)
(fig. 25.2.a-c)
Ammotrypane longicaudata Caullery, 1944 : 44, fig. 35.
Armandia longicaudata : Day, 1 95 1 : 50.
Body up to 30 mm. long with 30-32 setigers which bear gills from the second to
the last. Presetal lobe of parapodium prolonged especially on the anterior 6-10
segments (fig. 25.2.3, b). Lateral eyes between setiger 6 and 23. Anal funnel
(fig. 25. 2. c) obliquely truncate so that the anus opens downward. It is provided
with a very long ventral cirrus and an arc of about 12 tapered dorsal papillae.
Type locality : East Indies.
Records: Natal (30/30/s) ; Mocambique (26/32/i and 23/35/e, s).
Distribution : Madagascar (i), East Indies.
Armandia intermedia Fauvel, 1902
(fig. 25.2.d-g)
Armandia intermedia Fauvel, 1902: 86, figs. 29, 30; Day, 1957: 104.
Body (fig. 25. 2. d) about 12 mm. long and rounded at both ends. Twenty-seven
to twenty-nine setigers with gills extending over 24-25 segments from setiger 2 to
the last two or three. Lateral eye-spots (fig. 25. 2. e) on setigers 7 to about 18. Pre-
setal Up of parapodium not prolonged. Setae (fig. 25.2.f ) of last few segments not
obviously longer than those on preceding ones. Anal funnel (fig. 25. 2. g) short
with a long internal ventral cirrus and 10-20 clavate dorsal papillae.
Type locality : Senegal.
Records: Cape (34/i8;'s and 34/22,'i, s) ; Natal (29/31/i).
Distribution: Senegal (s), Ghana (s), Angola; Indo-west-Pacific from the Red
Sea (i), Persian Gulf and Ceylon to N.W. AustraUa, Japan and New Caledonia.
Armandia leptocirrus Grubc, 1878
(fig. 25.2.h)
Armandia le/itocirrus Gruhe, 1878: 194; Fauvel, 1919: 435; Day, 1951 : 50.
Body about 20 mm. long; 31-35 setigers with gills from setiger 2 to the last or
last but one. Lateral eye-spots on setigers 7 to about 18. Presetal lip of parapodia
not prolonged. Setae of last few segments longer than those of preceding segments.
Anal funnel (fig. 25. 2. h) long, and obhquely truncate so that the anus opens upward.
It has a long ventral cirrus and 12-18 fine dorsal papillae.
Type locality : PhiUppine Islands.
Records: Cape (33/27/s) ; Mocambique (26/32/i and 23/35/e, s).
Distribution : Tropical Indo-wcst Pacific from the Red Sea and Persian Gulf (s)
to the Andaman Is. (i) and New Caledonia.
578
POLVCHAETA OF SOUTHERN AFRICA
Fig. 25.2. Armandia longkaudala. (a) Anterior foot, (b) Posterior foot, (c) Lateral view of
po.sterior end. Armandia intermedia, (d) Entire worm (four times life size), (e) Three
segments from middle of body, (f) Capillary seta, (g) Posterior end. Armandia leplocirrus.
(h/ Posterior end. Ofthelina acuminata, (i) Entire worm (three times life size), (j) Posterior
end. Polyophthalmus fiictus. (k) Entire worm (four times life size), (l) Lateral view of
head end. (m) Posterior end.
OPHELIIDAE 579
OPHELINA Oersted, 1843
{inclnAmg AMMOTRTPANE Rathke, 1843)
Body vermiform, not divided into distinct regions. A deep ventral groove and a
pair of lateral ones along the whole body. Prostomium conical with subdermal
eyes. Segments annulated, lateral eye-spots absent. Cirriform gills, if present,
start on setiger 2 and continue to near the end of the body. Each parapodium with
a conical setigerous lobe, two bundles of simple capillaries and a small ventral
cirrus. Anal funnel with a long internal ventral cirrus and an arc of dorsal papillae.
Type SPECIES : Ophelina acuminata Oersted, 1843.
Ophelina acuminata Oersted, 1843
(fig. 25.2.i-j)
Ophelina acuminata Oersted, 1843; 46.
Ammotrypane aulogasler Rathke, 1843 : 188, 10 figs. 1-3 ; Fauvel, 1927 : 133, fig. 47 a-e.
Body (fig. 25.2.i) vermiform with 48-50 segments and up to 50 mm. long. Pro-
stomium conical with a filiform tip. About 46 pairs of gills from setiger 2 to the last
two to three. Setigerous lobe elongate, conical. Ventral cirrus small. Anal funnel
(fig. 25.2.J) scoop-shaped and slit ventrally. The posterior end is fringed by many
small marginal papillae and two much stouter ones arise at the anterior end of the
sUt and flank a long median internal cirrus.
Type locality : Danish seas.
Records: Cape (34/18/d to 33/27/s) ; Natal (30/31/d, 29/31/s, d).
Distribution : Arctic ; N. Atlantic from Norway (s) and Greenland (s, d) to
the English Channel (s) ; N. Pacific from the Behring Sea to Japan ; tropical
Indian Ocean (i, s).
POLYOPHTHALMUS Quatrefages, 1850
Body short, without distinct regions. A ventral groove along the whole length.
Prostomium short, proboscis small. No branchiae. Lateral eyes present. Parapodia
biramous with simple setae but no ventral cirri. Anal tube short with small papillae.
Type species : J\fais picta Dujardm, 1839.
Polyophthalmus pictus (Dujardin, 1839)
(fig. 25.2. k-m)
Nais picta Vyxiyzrdm, 1839: 293.
Polyophthalmus piclus : Fauvel, 1927: 137, fig. 48 1-n.
Body (fig. 25. 2. k) slender, up to 25 mm. long and consists of 27-28 biannulate
segments with a pattern of brown marks dorsally. Prostomium (fig. 25. 2. 1) a
rounded hood with subdermal eye-spots. A single bundle of very small capillaries
580 roi.Yc:HAi; lA oi- southerx Africa
per foot except on the last few. Lateral eye-spots present on seti,e;crs 7-21 but
diflleult to see. .\iial tube (fig. 25. 2. m) rudimentary with small marginal papillae.
Type locality : France.
Records: Xatal (30 '30 i and 29 '31,1) ; Mocainbic[ue (26/32/1 and 23/35/e).
UiSTRlBl'Tio.N : All warm and tropical seas - Atlantic from the English Channel
(i) to the Gulf of Mexico (i) and tropical West Africa (i, s) ; Mediterranean (i) Red
Sea (s) and Indo Pacific to Japan ; western C^ananda and Southern California
(i, s) ; Madagascar (i).
TACHYTRYPANE Mcintosh, 1879
Body elongate and vermiform with a smooth, tough, cartilaginous surface. A
ventral groove along the whole length. No branchiae. No lateral eye-spots. Para-
podia biramous but without ventral cirri. Pygidium marked off from the posterior
segments and curved down so as to open ventrally.
Type SPECIES : Tachrtnliam- jeffrcyn Mcintosh, 1879.
Records: Not recorded from southern Africa.
COSSURIDAE 581
Family COSSURIDAE Day, 1963
Small thread-like worms with numerous similar segments. Prostomium conical
and without appendages or eyes. Pharynx eversible, soft and unarmed. Peristome
achaetous and without appendages. The second segment sometimes achaetous as
well. All subsequent segments without parapodial lobes but with one or two bundles
of simple pointed setae with rather flattened spinulose blades. A single very long
cylindrical dorsal gill (or ? tentacle) on sctiger 2 or 3. Pygidium with anal cirri.
Records from sotithern Africa
Cossiira coasta Kitamori ..... 56Cd
COSSURA Webster and Benedict, 1887
The only recorded genus has the characters of the family.
Type species : Cossura longocirrata Webster and Benedict, 1887.
REMARKS
Cossura is a small thread-like worm which lives in sandy mud in fairly deep water
and appears to be a deposit feeder. A key to the known species will be found in
Laubier (1963). It is suspected that several of the specific names that have been
erected are really synonyms for there are few useful taxonomic characters. The
shape of the anal cirri seems to be distinctive but unfortunately the posterior end is
often missing.
Cossura coasta Kitamori, i960
(fig. 26.i.a-d)
Cossura coasla Tsiiamori, igGo: 1082, fig. i a-f; Day, 1963: 427.
Body thread-like, rounded in section and up to 15 mm. long with 107 segments.
Prostomium (fig. 26.1. a) a blunt depressed cone without eyes or appendages but
with a pair of nuchal shts. Pharynx eversible and lobed. Peristome and segment
2 partly fused, apodous, and achaetous. Subsequent segments broader than long
with poorly marked intersegmental constrictions. Parapodial lobes absent and the
setae arise directly from the sides of the body. Segment 3 (the first setiger) has a
single bundle of setae but all subsequent segments have two bundles which arise
close together (fig. 26.i.b). The setae are all simple capillaries (fig. 26.1. d) with
longer and shorter blades which are somewhat flattened and finely spinulose on
one margin. A single very long cylindrical gill arises from the dorsal surface of
setiger 3. It is about three-quarters the length of the body. The last few segments
S82
I'OI.YCllAETA or SOUTHERN AFRICA
(fig. 26. 1. c) lack setae and the pygidium bears three long filiform anal cirri. In
Japanese specimens these arc forked at the ends bnt not in South African specimens.
Type locality: Scto Inland Sea, Japan.
Records: South-west .\frica (2G/i4/d) ; Clape (from 33/1 7/d to 36/21/d) ; Natal
(•^9/30/s, 29/31/d).
Distribution : Japan.
Fig. 26.1. Cossiira comla. (a) Anterior end. (b) Foot from mid-region.
(c) Posterior end. (d) Blade of rapillary seta.
SCALIBREGMIDAE 583
Family SCALIBREGMIDAE Malmgren, 1867
Body either short and stout or longer and arenicoliform but never with more
than 30-60 segments. Prostomium either bilobed or with two divergent frontal
horns. Eyes present or absent. Nuchal slits present. Proboscis eversible, but soft
and unarmed. Buccal segment achaetous. Parapodia biramous but poorly developed
with or without dorsal and ventral cirri on the posterior part of the body. Setae all
simple and consist mainly of capillaries accompanied by a few forked setae ; in
addition there may be acicular setae in the first one to four segments. Branchiae if
present are branched and restricted to the first few segments. Pygidium with or
without anal cirri.
Records from southern Africa
Asclerocheilus capensis Day .... 56Cd
Hyboscolex longiseta Schmarda . . . 4Ci, aGWi, 4oNi,
44Ci, 45Pi, 5iCsd
i2Ci
36Ci
5iCis, 56\Vs
5iCd
48Cd, 5iCsd
as Lipobraiichhis capensis \Villey .
as Lipobranchius longisetus (Schmarda)
Parasclerocheilus capensis Day .
Polyphysia crassa (Oersted)
Scalibregma infiatum Rathke .
REMARKS
Scalibregmids are rare worms and not well known. They are found in muddy
situations and in all of them the gut contains mostly mud. On the otJier hand, they
do not burrow deep in the mud and presumably live in the surface layer of silt
feeding on detritus. Hyboscolex Uves in muddy rock crevices and is often found in
old tubes formed by other animals. Parasclerocheilus lives among loose muddy stones
and Scalibregma is dredged from similar habitats. Polyphysia is a deep water species
living on mud or foraminifera.
THE MAIN DIAGNOSTIC CHARACTERS
Useful discussions of the family will be found in Ashworth (igoi), Chamberlin
(1919) and Furreg (1925). Both ChamberUn (1919) and Fauvel (1927) provide
keys.
The most important taxonomic characters are the shape of the whole body,
details of the prostonuum, the presence of branchiae and acicular setae in the first
few segments and the nature of the posterior parapodia.
The body. In most genera the body is either arenicoliform with a swollen
anterior region and a narrowed tail or short and maggot-shaped. In Parasclero-
cheilus however, the body is elongate and fusiform. In some species the segments in
the middle of the body are divided into three annuli, in others there are four.
The prostomium and eye-spots. The prostomium is bilobed to varying degrees. In
Polyphysia there are merely two stout lobes divided by a deep median \'-shaped
584 I'Ol.VCniAKTA OF SOUTHERN AFRICIA
ntitch; ill other genera sueh as Sialilirfi^mn these lateral lobes are better dexeloped
so tli.it the whole prostoinium is broadly T-shaped. The exact shape in any single
speeinieii is greatly alleeled by the degree of contraction and only major differences
are important. The presence or absence of eye-s|5ots is useful at the species level
but it should be noted that when the prustomium is partly retracted into the peri-
stomial collar the eyes may be hidden.
Ihdiuiiidi'. Br.mching gills ma)' arise lichind the notopodia of three to six segments
from setiger 2 onwards. It is important to note, however, that the number of gills
incrc.ises with the si/e of tlir worm and that small juveniles ni.iy e\en lack gills
entirely.
The iiiat\ The typical setae in both r.imi rif the parapodia are ca])illaries accom-
panied by a few forked setae but a few genera have acicular setae in the first one to
four feet. The capillaries may be either smooth and cylindrical or minutely spinu-
lose. The forked setae usually have unequ.il prongs with spinules on their inner
margins. The acicular setae are stouter than the other setae and typically have
abruptly pointed tips but they m.i)' have blunt or tapered tips ; in the latter case
they are not very different from the capillaries. When acicular setae occur they
replace the forked setae in a few anterior segments.
Paiajioiliti. The parapodia are ne\'er well de\eloped in anterior segments. The
parapodial jirojections may even be entirely .ibsent .iiid the two bundles of setae
then project directly from the sides of the body with a small ciliated button or
'^lateral organ" between them. In many genera however, the anterior setae arise
from small conical sctigcrous lobes. These tend to become longer in the posterior
part of the body and may be accompanied by both dorsal and ventral cirri or by
\entral cirri alone. When both cirri are well developed they tend to be conical and
as long as tlie sctigcrous lobes but when only the \'entral cirrus is present it tends to
be filamentous and may arise posterior to the neuropodium.
The synonymy of the genera is confused. Schmarda (1861) described two genera,
Hvbnscolex (type //. lonninia) and Oncnscnkx (type (). dicraiidchactin). As Augner
(19 1 8) has shown after an examination of Schmarda's material, the two genera are
synonynifius although Oncoscokx hipatiitus Schmarda is a fragment of Dasyhranclnis
sp. Oncoscultx (licranochactus thus becomes Hvboscolex (licranncitachis (Schmarda). As
shown by Day 1961 Eumcnin rclkidala Mcintosh 1885 must be transferred to Hvho-
scoli'x. It differs from //. !o/igiscla in lacking eyes and in having parapodial projections
in posterior segments luit like //. ln)iai\eta it lacks dorsal and ventral cirri. Eiimcnia
oculala Ehlers 1901 which was transferred to the genus Onancokx by Ehlcrs (19 13)
also becomes a species <>[ IIyliu\cali-x. Kiimciiiii glahra Ehlers 1887 was used by Cham-
berlin (19 19) as the type species of his new genus Kcbuita. I have not seen Ehlers'
original description but judged from the remarks of Chamberlin and Furreg it is
essentially similar to Hvboscolex ap.irt from .m .ibruptly u.iirowcd .iii.il tube.
Furreg has shown that the genus /''iininiii: is preoccupied in the Li-pidoptera so
that Oersted's E. cr/i.\\ti becomes l-'o/Y/ihv\i(i crmsn. Etimniia ir/lii-rsii Mcintosh 1869
\vas made the type of a new genus Li/i(ihiiiiichiiis by ClunninglKini ,ind Ramage (li
SCALIBREGMIDAE 585
It differs from Poljphysia in the absence of gills but according to Stop-Bowitz (1945)
it is no more than a juvenile P. crassa in which the gills are undeveloped.
Key to Genera
1 Acicular setae present in one or more of the first few parapodia ..... 2
- Acicular setae absent ............ 4
2 Branched gills present on the first few segments. No dorsal or ventral cirri
PARASCLEROCHEILUS (p. 585)
- Branched gills absent ............ 3
3 Ventral cirri present on posterior segments ..... SCLEROCHEILUS*
- Ventral cirri absent ASCLEROCHEILUS (p. 586)
4 Body short and maggot-shaped (fig. 27.1 k). Prostomiuni broadly V-shaped. Branched
gills present POLYPHYSIA (p. 586)
- Body arenicoliform (fig. 27.2.3). Prostomium broadly T-shaped with lateral projections.
Gills present or absent ........... 5
5 Posterior parapodia with both dorsal and ventral cirri ...... 6
- Posterior parapodia if developed lack dorsal and ventral cirri. (No gills)
HYBOSCOLEX (p. 588)
6 Branched gills present SCALIBREGMA (p. 589)
- Gills absent PSEUDOSCALIBREGMA
PARASCLEROCHEILUS Fauvel, 1928
Body elongate and fusiform. Prostomium T-shaped with a pair of frontal lobes.
Eyes present. Buccal segment achaetous. Branched gills on sctigers 2 to 6-7.
Segments with four annuli after the first few. Parapodia biramous but poorly
developed and dorsal cirri absent. A ventral cirrus behind the neurosetae of
posterior segments. Acicular notosetae present in the first one to four anterior
segments and fine capillaries plus a few forked setae in subsequent ones.
Type species : Parasderocheilus branchiatus Fauvel, 1928.
Parasclerocheilus capensis Day, 1961
(fig. 2 7.i.a-f)
Parasclerocheilus capensis Day, 1 96 1 : 517, fig. i o c-f.
Body (fig. 27. 1. a) arenicoliform, up to 30 mm. long with 60 segments and blood-
red when alive. Prostomium (fig. 27.i.b) T-shaped with divergent frontal lobes
and a pair of large reniform eyes. Buccal segment incomplete ventrally and the
lower lip formed by setiger i. The first four segments biannulatc and subsequent
ones quadriannulate. Parapodia biramous with swollen setigerous lobes and a
ciUated lateral organ between them. A small ventral cirrus (fig. 27.1.C) below the
neurosetae on the last third of the body. Bushy gills behind the notopodia of setigers
2-7. Setiger i with five to six acicular setae (fig. 27.i.d) in the notopodium. Sub-
sequent setae are mainly fine capillaries (fig. 27.i.f) plus a few short forked setae in
K
586 rC)I.V(.;H,\ElA OF SOUTHERN AFRICA
both rami. Forked setae (fig. ay.i.e) with unequal prongs both feathered on their
inner margins.
Typk locality : Langcbaan Lagoon, South Africa.
Records; South West Africa (28/16/s) ; Cape (33/18/i and 34/18/s to 33/25/s).
Distribution : Endemic.
ASCLEROCHEILUS Ashwonh, 1901
Body arenicohform. Prostomium T-shaped with divergent frontal horns. No
eyes. Buccal segment achactous. No gills. Segments with four annuli after the
first few. Parapodia biramous but poorly developed anteriorly and never have
dorsal or ventral cirri. Curved acicular setae accompany the capillaries in the first
one to three setigers. Subsequent segments with fine capillaries and forked setae.
Anal cirri digitiform.
Type species: Lipobranchius intermedins Saint-Joseph, 1894.
Asclerocheilus capensis Day, 1963
(fig. 27.1.g-j)
Aiclerocheihts capemis Day, 1963a: 428, fig. 10 a-f.
Body arcnicoliform, about 10 mm. long with about 30 segments. Prostomium
(fig. ay.i.h) with stout, divergent frontal lobes but no eyes. Peristome short and
achaetous. The first seven segments vaguely biannulatc, later ones clearly quadri-
annulate and tessellated. No gills. Parapodia (fig. 2 7.i.g) biramous but poorly
developed. Setigers i and 2 with acicular setae plus long capillaries in both rami.
Acicular setae (fig. 27.1.1) with stout shafts and curved, tapered tips. Setiger 3 and
all subsequent segments have fine capillaries of varying lengths plus a few forked
setae (fig. 27.1.J) with unequal prongs.
Type locality : Agulhas Bank, South Africa.
Records: Cape (34.23. d).
Distribution : Only one record.
POLYPHYSIA Qualrcfages, 1865
( [= EVMENIA Oersted- preoccupied] indudmgUPOBRAM.HIUS Cunningham
and Ramage, 1888.)
Body short and swollen with about 30 segments. Prostomium bilobed. No eyes.
Buccal segment achaetous. Sctigerous segments with three annuli and the surface
tessellated. Parapodia biramous but poorly developed, consisting of two cushion-
shaped setigerous lobes with a lateral organ between them. Branching gills present
on anterior segments. Setae include slender capillaries and forked setae but no
acicular setae. No anal cirri.
Type species : Eumenia crassa Oersted, 1843.
SCALIBREGMIDAE
587
a
'5^
¥
Vr
IM^
(^ i)
G (i\)
,j7 ■■ •
- h
'M: r ' m
Fig. 27.1. Parasclerocheilus capmsis. (a) Entire worm in dorsal view (four times life size).
(b) Head, (c) Posterior foot, (d) .^cicular seta, (e) Forked seta, (f) Capillary seta.
Asclerocheilus capensis. (g) Twelfth foot, (h) Dorsal view of anterior end. (i) .Vicular seta.
(j) Forked seta. Polyphysia aassa. (k) Lateral view of entire worm (three times life size),
(i.) Head, (m) Foot from mid-region, (n) Capillar^' seta, (o) Forked seta.
588 POI.VCIIAETA OF SOUTHKRX .\1RIC:A
Polyphysia crassa (Oersted, 1843)
(fig. 27.i.k-o)
Eumenia crassa Oersted, 1843 : 47 ; Fauvel, 1927 : 127, fig. 45 i-k.
Polyjihysia crassa : Stop-Bowitz, 194',: 75, figs. 4-6 ; Day, 1961 : 516.
Body (fig. 27. 1. k) maggot-like, up to 30 mm. long with about 33 segmcnt.s. Seg-
ments triannulatc and surface tessellated. Prostomium (fig. 27.1.I) deeply bilobed
and without eyes. Buccal segment achactous. Parapodia (fig. 2 7.i.m) biramous
with two indistinct setigcrous lobes each bearing long capillaries plus a few forked
setae. Dorsal and \cntral cirri absent. About five pairs of branched gills behind
the notopodia starting from setigers 2 but not de\'elopcd in juveniles. Capillary
setae (fig. 27.i.n) very long and smooth; forked setae (fig. 27.1.0) with very long
prongs feathered on their iimer margins.
Type loc.\lity : Denmark.
Records: Clape (31/16'd).
Distribution' : Atlantic from Greenland (s, a) and Norway (s, d) to the North
Sea ; ? Mediterranean.
HYBOSCOLEX ^chmaxAn, 1861
(including ().\C(>SCVLEX Schmarda, 1861, and ? KEBUITA Chamberlin, 19 19.)
Body small, swollen antcriorh', arenicoliforni. Prost(jmium T-shaped. Eyes
present or absent. Buccal segment achactous and fused to the first setigcr ventrally.
Anterior segments with three annuli. Parapodia biramous but may lack setigcrous
lobes and never possess dorsal or \entral cirri. Numerous simple capillaries and a
few forked setae but no acicular setae.
Type species: Hrboscnlex lonziseta Schmarda, 1861.
Hyboscolex lotigiseta Schmarda, 1861
(fig. 27.2.a-d)
Hvhoicolcx longisela Schmarda, 1861 : ^4, pi. 27 fig. 211.
Upobramhitii capcnsii Willey, 1904: 266. pi. 14 fig. 14, pi. 15 figs. 23, 24.
Body (fig. 27.2.3) about 15 mm. long with 50 segments and often dark brown in
colour. Prostomium (fig. 27. 2. b) with a pair of stout lateral projections so that the
whole is T-shaped. Two fused pairs of large eyes often hidden by the peristomial
fold. Peristome short, achactous, ob\ious dorsally and laterally but not visible
ventrally so that the posterior lip is formed by setigcr i. Anterior segments inflated
and triannulatc, but posterior ones not annulated. No branchiae or parapodial
projections of any sort in any part of the body, the setae arising directly from the
body wall (fig. 27.2. c). Both notopodial and neuropodi.il bundles of setae contain
SCALIBREGMIDAE
589
numerous smooth capillaries and a few forked setae (fig. 27. 2. d) with unequal
limbs. Five small anal cirri.
Type locality : Cape of Good Hope.
Records: South-west Africa (22/14/i) ; Cape (from 29/16/i to 35/20/s and
33/25/s) ; Natal (30/30/i) ; Mocambique (26/32/i).
Distribution: Auckland Is. (New Zealand) (i).
SCALIBREGMA Rathke, 1843
Body arenicoliform. Prostomium T-shaped with a pair of latero-frontal lobes.
Eyes absent. Buccal segment achaetous. Segments with four annuli after the first
few. Branched gills present on some anterior segments. Parapodia biramous with
Fig. 27.2. HyboscoUx longiseta. (a) Dorsal view of entire worm (four times life size), (b) Head.
(c) Foot from mid-region, (d) Forked seta. Scalibregma inflatum. (e) Branchiferous foot.
(f) Head end. (g) Posterior foot, (h) Forked seta, (i) Capillary seta.
590 POLYCHAETA Of SOUTHERN AFRICA
lateral organs between the short sctigerous lobes. Dorsal and ventral cirri present on
posterior segments. The setae are mainly capillaries plus a few forked setae but large
acicular setae are absent. Anal cirri digitiform.
Type species: Scalibregma inflatum Rathkc, 1843.
Scalibregma inflatum Rathke, 1843
(fig. 27.2.e-i)
Scalibregma inflalum Rathkc. 1843 : 184, pi. 9 figs. 15-21 ; Fauvel, 1927 : 123, fig. 44 a-f.
Body rusty brown, swollen anteriorly but narrowed posteriorly. Length up to
60 mm. with 60 segments. Skin tessellated. The first few segments triannulate, the
remainder quadriannulate. Prostomium (fig. 27.2.f) pale with divergent processes
forming a stout T. No eyes. Buccal segment achaetous. Four pairs of bushy gills
(fig. 27. 2. e) behind the notopodia of setigers 2-5. In anterior segments both the
notopodia and the neuropodia are truncate cones with a lateral organ in the form
of an evaginablc pit between them. Conical dorsal and ventral cirri from setiger
16 or 18. In the posterior region the setigerous lobes (fig. 27. 2. g) are more prominent
and compressed. The setae are mainly capillaries (fig. 27.2.1) but a few forked
setae (fig. 27. 2. h) are also present with the inner sides of the unequal prongs minutely
feathered. Five slender anal cirri.
Type locality : Moldc, Norway.
Records : Cape (from 32/1 7, d and 33/17/5, d to 36/2 i/d and 34/26/d).
Distribution : Cosmopolitan from the Arctic to Antarctic ; in depths of loo-iooo
metres in the tropics but at all depths from a few metres down in cold seas. Habitat
muddy sands.
CAPITELLIDAE
59'
Family CAPITELLmAE Grube, 1862
Body elongated, reddish, rounded in section and without obvious parapodia so
as to appear like oligochaetcs. Prostomium voluminous, evaginable but unarmed.
Body divided into an anterior, rather short and swollen thoracic region and a longer
abdominal region which often bears inconspicuous gills. Lateral sense organs
commonly present on many segents as small eversible, ciliated pits between the
notopodia and the neuropodia. Parapodia biramous but poorly marked, being
reduced to two bundles of setae on the thorax and two setigerous ridges on the
abdomen. Genital pores on some segments become visible in sexually mature
adults. Branchiae if present, are either branched and retractile organs on some
abdominal segments or are represented by vascular enlargements of the notopodia
and'or neuropodia. Setae include capillaries and hooded hooks. Special genital
setae may be present.
Records from southern Africa
Capitella capitata (Fabricius)
Dasybranchus bipartitus (Schmarda)
as Oncoscolex bipartitus Schmarda
as Branchoscolex craspidochaetus Schmarda
as Branchoscolex sphaerochaetus Schmarda
as Branchoscolex oUgobranchus Schmarda
as Dasybranchus caducus (non Grube)
Dasybranchus caducus (Grube) .
Heteromastus filiformis (Claparede)
Leiochrides africanus Augener
Mediomastus caperuis Day
Notomastus aberans Day .
Notomastus fauveli Day
Notomastus latericeus Sars
Parheteromastus tenuis Monro
Pulliella armata Fau\'el .
Scyphoproctus djiboutiensis Gravier
1 5 As, 2iCi, 26\Vi,
44Ci, 45Pi, 51CS
44Ci, 51CS
4Ci
4Ci
4Ci
4Ci
i3Ci, 32Ci, 3601, 48CS
27Mi, 4oPi, Ni, 45Pi
4oNi, 44Ci, 48Cd
26Ai, 35Ci, 44Ci, Ns
5iCsd, Nsd
45PiNi, 3iCi
44Ci
32Cd, 38Csd, siCsd, — Nd
45Pi
45Pi
— Ps
BIOLOGICAL .\OTES
Capitellids have a superificial resemblance to eardiworms and seem to have
similar habits. They burrow in various grades of sandy mud, their guts are filled
widi mud suggesting that diey ingest a good deal of inorganic matter along with the
organic particles on which they subsist. The only burrowing organ is the eversible
proboscis which is used in the same way as that oi Arenicola. The thorax is stout and
muscular and is highly extensible. The abdominal segments with their long rows of
hooks grip the substrate as the thorax is extended.
Capitella lives in black mud and is often foimd in estuaries. It is capable of toler-
ating very low oxygen tensions for enormous numbers may occur in polluted areas
592 roi.VOlIAElA OK .SOLTHERX AIRICA
such .IS harbour basins. .Xnlomastiis latericetis and Htieromaslus filijoiinis H\c hi clc.incr
sandbanks and arc common in shchercd bays and in drcdgings. Notomastus Jauvelii
which has well developed gills makes deep burrows in sandy mud and Dasybranchus
caducus has similar habits. Tiic latter is widespread in many parts of the world and
harbours a number of commensals in its tube, in particular the scale worms Ilarmolhoe
lunulala and Lepidailhenia maculala.
THi; MAI.\ DI.AG.NOSTIC CHARACTERS
The most important works are those of Eisig (1887) who gave a detailed account
of the anatomy, Fauvcl (1927) who gave a key to the European species and Hartman
(1947) whose review covers all genera erected up to 1944. Her chart showing the
distribution of the various types of setae is particularly valuable.
The most useful taxonomic characters are the number of dioracic segments, the
distribution of the setae, the detailed structure of the hooded hooks and the presence
or absence of branchiae. Several other specialised features are useful in individual
genera.
The head consists of a conical prostomium followed by a well de\'eloped pcristo-
niium. The prostomium always possesses a pair of evaginable nuchal sUts and
often a pair of eyes though the latter are rather indefinite pigmented areas. The
proboscis is soft and eversible, often voluminous. The peristomium is biannulated
and may be either achaetous or bear a few capillary setae in both rami. It may be
noted in passing that these setae are not always easy to find.
The Ihnriix and i/s selae. The anterior region of the body or thorax includes the
peristomium and 8-18 setigerous segments. Although these segments are often
rather shorter and stouter than the abdominal segments which follow, the thorax
is not always well defined. Changes in the nature of the parapodia and setae are
the best criteria for determining the junction of the thorax and abdomen. The
parapodia are never well developed and in the thorax they are mere grooves from
which the short series of notopodial and neuropodial setae arise. In the abdomen
the parapodia form raised ridges and the setal rows are considerably longer than
those of the thorax. Moreover capillary setae are usually (though not always)
restricted to the thorax and the thoracic hooks, when such occur, project further
from the surface than do those of the abdomen.
The presence of setae on the peristomium, the number of thoracic segments and
the distribution of different types of setae, e.g. winged capillaries, hooded hooks and
genital setae, are all cliaracters of generic importance.
The abdomen and its selae. The abdominal segments are much more numerous than
those of the thorax and the segments are longer and usually better marked, occasion-
ally becoming moniliform or campanulate towards the posterior end. The neuro-
podial rows of setae are longer than those of the notopodium though the proportions
change from the anterior to the posterior segments. The two notopodial rows may
fuse in certain species and the distance across the ventrum between the two neuro-
podial rows seem to be of specific importance so long as similar parts of the abdomen
are compared.
CAPITELLIDAE 593
The last few segments of the abdomen develop stout acicular setae in both
Pulliella and Scyphoproctus and in the latter genus the pygidium is flattened and saucer-
like. In some genera there is one short ventral anal cirrus and in others there may
be two but as the body fragments readily, these are often missing.
Branchiae. A closed circulatory system is absent and the ventral part of the
coelome acts as a hacmocoele. The blood is red and when the worm is alive it is
easy to see that certain thin walled projections or branchiae connected with the
abdominal parapodia have a respiratory function. \Vhen the worm is preserved
however, the branchiae are seldom obvious. In many genera (e.g. Capitella) branchi-
ae are entirely absent. In others (e.g. Notomastus) the branchiae are rather indefinite
projections of the notopodia and the superior edges of the neuropodia. In Branchio-
capitella the branchiae arc digitiform projections from the medial margins of the
notopodia and in Dasjbra?!ckus they are branched filaments which may be retracted
into a small pocket immediately above the superior end of the neuropodium. In
brief the branchiae may be absent, indefinite or retractile and in preserved material
they are seldom easy characters to use for identification.
Lateral sense organs, nephridiopores and genital pores. The lateral sense organs are small
ciliated papillae between the notopodia and neuropodia. They arc seldom easy to
see and may be limited to certain parts of the body or be entirely absent.
Nephridiopores are also difiicult to see in sexually immature specimens but in
ripe males and females where the nephridiopores act as genital pores they may be
very conspicuous with prominent glandular lips. They are situated on the same
horizontal level as the lateral sense organs but slightly posterior to them. They are
not found on all segments but usually occur on segments near the junction of the
thorax and abdomen. Their exact segmental arrangement is said to be of specific
importance.
Structure of the hooded hooks. The detailed structure of these setae is surprisingly
constant along the length of the body and Hartman (1947) has shown that they are
of great systematic importance at the species level. Unfortunately they are very
small and it is necessary to examine them under oil immersion both in profile and
in frontal view to see all the details. According to Hartman's terminology (p. 396)
each hook has the following parts :
"The long embedded stem is the shaft with the node near its middle length. It
extends distally often forming a thickened shoulder and a slender neck and terminates
distally in a hooked beak provided with a larger yan^ (= main fang) surmounted by
a crescent composed of smaller teeth or denticles. The beak is more or less loosely vested
with a hyaline hood or sheath (rarely absent in Protocapitella*) ; this is capeUke,
with an oval aperture distally to allow for protrusion of the teeth ; the aperture is
continued basally as a cleft that extends nearly to the basal end of the hood."
The most important characters are the proportions of the hood, the number of
teeth abov-e the main fang, their arrangements and relative size. These latter
characters can only be seen under oil immersion in frontal view.
* Protocapitella b now known to be a synonym of Brancliiomaldane a genus of the family Arenicolidae.
594 I'OI.VCHAKTA OF SOUTHERN AFRICA
Key to Gener.\
1 Setae present on the peristomial segment. Only nine thoracic setigers ... 2
- Setae absent from the peristomial segment. Nine or more thoracic setigers . . 3
2 l*oslerior abdominal segments with cirriform dorsal branchiae . BRANCHIOCAPITELLA*
- Abdominal segments without branchiae. (Genital setae in the male, iig. 2fi.2.i)
CAPITELLA (p. 594)
3 Nine thoracic setigers. Genital setae may be present ...... 4
- Ten or more thoracic setigers. Genital setae absent ...... 5
4 No stout spines on the last few abdominal segments. Genital setae present in the noto-
podia of setigers 7 and 8 CAPITOMASTUS*
~- Stout spines present on the last few abdominal segments {fig. iS.t.c). Genital setae absent
PULLIELLA (p. 595)
5 Ten thoracic setigers (with capillaries in setigers 1-4 and hooks in setigers 5-10)
MEDIOMASTUS (p. 600)
- More than ten thoracic setigers .......... 6
6 Eleven thoracic setigers ........... 7
- More than 1 1 thoracic setigers .......... 9
7 Capillary setae in both rami of thoracic setigers . NOTOMASTUS (p. ^()y)
- Capillary setae restricted to anterior thoracic setigers and hi toks in posterior ones . . 8
8 Hooks start on setiger 5. Branchiae absent . PARHETEROMASTUS (p. 605)
- Hooks start on setiger 6. Neuropodial branchiae present . . HETEROMASTUS (p. 6oi)
- Hooks start on setiger 7. Xotopodial branchiae present . . . BARANTOLLA*
9 Twelve or thirteen thoracic setigers ......... 10
- Fourteen or more thoracic setigers . . . . . . . . . 13
10 Capillary setae in both rami of all thoracic setigers . . . . . . . it
— Both capillary setae and hooks on the thorax. (Posterior end expanded to form an anal
plate) HETEROMASTIDES*
11 Twelve thoracic setigers . . . . . . . . . . . 12
— Thirteen thoracic setigers. (Retractile branchiae abo\e the neuropodia of posterior
abdominal segments (fig. 28.3.k) DASYBRANCHUS (p. G03)
12 .Xbdomen ends in an anal plate armed with spines. Branchiae absent
SCYPHOPROCTUS (p. 604)
— .\bdomen ends with normal segments. Retractile branchiae somclinics present
LEIOCHRIDES (p. 600)
13 Branchiae absent. Fourteen to seventeen setigers . PSEUDOCAPITELLA*
— Branchiae compound, notopodial. Sixteen to seventeen thoracic setigers
ANOTOMASTUS*
— Branchiae compound, neuropodial. Eighteen to twenty tiioracic setigers
CAPITOBRANCHUS*
CAPITELLA Blainville, 1828
Thorax of nine segments. Peristomitim and the next six segments with winged
capillaries. Hooded hooks present in posterior thoracic segments, starting from
segment 5 so that segments 5, 6 and 7 may have both hooks and capillaries. Seg-
ments 8 and 9 with copulatory setae in the notopodia of the male but hooks only
in the female. Lateral organs absent and genital apertures limited to the last few
thoracic segments. Abdominal segments with hooded hooks in both rami. No
branchiae.
Type species: Liimhriciis cajnlaliis Fabricius, 1780.
CAPITELLIDAE 595
Capitella capitata (Fabricius, 1780)
(fig. 28.2.i-m)
Lumbricus capitatus Vahricius, 1780: 279.
Capitella capitata : Fauvel, 1927 : 154, fig. 55 a-h ; Hartman, 1947 : 404, pi. 43 figs. 1-2.
Body (fig. 28. 2. i) thread-like, 30-40 mm. long by i -o mm. and dark red when
alive. Prostomium conical with a pair of ventro-lateral eyes. Thoracic segments
all biannulatc with capillaries in both rami from the first or peristomial segment to
the sixth. Seventh segment variable and may have capillaries only or hooks only
or both. In the female the eighth and ninth segments have hooks in both rami but
in the male (fig. 28.2.J) there are enlarged genital hooks dorsally slanting inwards
towards the genital opening and normal hooks ventrally. Abdominal segments
(fig. 28. 2. k) smooth in section with long-shafted hooks in both rami. Each hook
(fig. 28.2.1, m) with an arc of three to four teeth above the main fang.
Type locality : Greenland.
Records: South West Africa (22/14/i) ; Cape (from 32/18/ea nd 34/18/s lo
32/28/e).
Distribution : Arctic ; North Atlantic from Greenland (i, s) and Scotland (i) to
North Carohna (i) and .Senegal (s) ; Mediterranean ; Black Sea ; cold North Pacific
from Behring Sea to N.W. Japan ; Southern California (s) ; subantarctic (i, s, d).
PULLIELLA Fauvel, 1929
Thorax with an achaetous peristomium followed by nine setigerous segments
bearing winged capillaries in both rami. No specialised copulatory setae. Anterior
abdominal segments with hooded hooks in both rami, but the last 8-1 1 with acicular
spines in the notopodia and hooded hooks in the neuropodia. Branchiae absent.
Pygidium with a pair of anal cirri.
Type species : PuUiella armata Fauvel, 1929.
Pulliella armata Fauvel, 1929
(fig. 28.i.a-f)
Pulliella armata Fauvel, 1929 : 184, fig. 3 ; Fauvel, 1930 : 48, fig. 13 a-h.
Body about 20 mm. long for 70 segments. Prostomium (fig. 28.i.b) broad and
blunt with indistinct eyes. Peristomial segment biannulatc, achaetous. Nine
thoracic setigers with bundles of winged capillaries in both rami (fig. 28.1. a).
Anterior abdominal segments (fig. 28.i.d) with rows of hooded hooks in both rami,
but no parapodial ridges ; later segments have better marked tori. Most of the
abdominal segments are short and close to one another but the last 8-1 1 (fig. 28.1.C)
are enlarged, have deep intersegmental grooves and bear stout, straight, blunt,
acicular spines in the notopodia (fig. 28.i.f) and normal hooded hooks (fig. 28.i.e)
59'J
I'OI.VCHAETA OF SOUIHERN AlKKIA
Fic. jB.i. PuUiella armala. (a) Lalcral view of anterior end. (n) Dorsal view of head.
(c) Lateral view of posterior end. (ni DiaRranimatit T/S of anterior abdominal segment.
(e) Hooded hook, (f) Posterior notopodial spine. .Notommlus faiiveli. (g) T/S anterior
abdominal segment, (h) T/.S branchiferous abdominal segment, (i) Lateral view of anterior
end. (j, KJ Profile and face view of hooded hook, (l) Capillary seta. Kolomaslus abcram.
(M) Entire worm (twice natural size), (n) .Xnterior end. (o) T/S anterior abdominal
segment, (p, (j) Profile and face view of hooded hook.
CAPITELLIDAE 597
with two arcs of denticles in the neuropodia. Pygidium (fig. 28.1.C) with two stout
diverging ventral cirri. No branchiae.
Type locality : Pulli Island, Gulf of Manaar.
Records: Mocambique (26/32/i).
Distribution : Tropical Indo-wcst-Pacific (Ceylon, Indo-China, New Cale-
donia).
NO TO MASTUS Sars, 1851
Thorax with an achactous peristomium followed by 1 1 setigerous segments. The
first setiger has notopodial capillaries and usually neuropodial capillaries as well,
and the remaining 10 always bear capillaries in both notopodia and neuropodia.
Abdomen with hooded hooks in both notopodia and neuropodia. Branchiae are
triangular vascular projections from both parapodial ridges or occasionally as
branched filaments from the notopodia.
Type species : J^'otomastus lalericeus Sars, 1 85 1 .
Key to Species
1 Gills as branched filaments arising from the abdominal notopodia (fig. 28.i.h) . N.fauveli
- Gills as simple projections from the abdominal parapodia (fig. 28. 2. d) ... 2
2 Neurosetae absent from first thoracic setiger. Abdominal gills minute, often indistin-
guishable ........... N. aberans
— Neuropodial capillaries present on first thoracic setiger. Gills well developed N. latericeus
Notomastus fauveli Day, 1955
(fig. 28.1.g-l)
Notomastus fauvelii Day, 1955 : 422, fig. 2 h-1.
Notomastus giganteus (non Moore) Fauvel, 1932: 194.
Body up to go mm. long by 3 mm. with 80 segments. Prostomium (fig. 28.i.i)
small and conical. Peristome achaetous. First thoracic setiger with both notopodial
and neuropodial capillaries. Anterior abdominal notopodia (fig. 28.i.g) united to
form a continuous dorsal ridge with short rows of hooks on either side of the mid-
dorsal hne. From the 15 th abdominal segment they separate to become well marked
papillae in front of branching gills (fig. 28.i.h). Posteriorly the gills develop up to
20 filaments. Ncphridial pores not conspicuous but present on the first six abdominal
segments. Abdominal neuropodia with a small superior branchial projecdon
anteriorly, but shorter and without projecdons posteriorly. Hooded hooks (fig.
28. 1. j, k) with two transverse rows of denticles above the main fang, the first row
with 8-10, the second with three to four denudes. Capillar)- setae (fig. 28.1.I) with
fairly narrow blades.
Type locality : Knysna Estuary, South Africa.
Records : Cape (from 34/18,1 to 33/26/c).
Distribution : Ceyon (d, vd), India (s).
598
POLVCHAETA OF SOUTHERN AFRICA
Fin. ■j8.2. M)lomii\liii Inlerkeus. (a) Lateral view nf anterior end. (is) and (c) Face view and
profile of hooded hook. I'd) T/.S anterior abdominal segment (diagrammatic). Lciochrides
ii/ncanm. (E) .Xnterior end. (f) Lateral view of three branchiferous segments, (g, ii)Face
view and profile of hooded hook. Cupitella capitala. (i) Entire worm ( '^ four times life
size), (j) Lateral view of setigers 7-9 of ,^. (k) T/.S abdominal segment (diagrammatic).
(l, M) Profile and face view of hooded hook. Mediomasttis capensiu (n) Lateral view of anter-
ior end. (o, p) Profile and face view of hooded hook.
CAPITELLIDAE 599
Notomastus aberans Day, 1957*
(fig. 28.l.m-q)
Motomaslm aberans Day, 1957 : 105, fig. 7 a-b.
Body about 60 mm. long with 1 20 segments (fig. 28. i .m) . Prostomium (fig. 28. i .n)
an elongate cone with indefinite eyes. Peristome achaetous. First thoracic setiger
with notopodial capillaries but without neurosetae. Abdominal segments (fig.
28.1.0) with short rows of hooks in both notopodia and neuropodia. Gills as minute
papillae (often indistinguishable) on the outer edges of the abdominal notopodia
and the superior edges of the neuropodia. Abdominal neuropodia with short rows
of hooks well separated ventrally. Hooks (fig. 28. 1. p, q) with hoods twice as long
as broad and with at least two rows of denticles above the main fang, the first row
with four to five equal teeth, the second with five to seven minute teeth. Posterior
abdominal segments often campanulate when contracted.
Type locality : Kosi Bay, South Africa.
Records : Cape (34/18/i) ; Natal (27/32/e) ; Mocambique (26/32/i and 23/35/e).
Distribution: Madagascar (i).
Notomastus latericeus Sars, 1851
(fig. 28.2.a-d)
J^otomastus latericeus Sars, 1851 : 199 ; Fauvel, 1927 : 143, fig. 49 a-h.
Length up to 300 mm. with more than 150 segments. Prostomium a blunt,
biannular cone with ocular specks. Peristomium achaetous (fig. 28. 2. a). The
succeeding 1 1 thoracic setigers with at first three and later five annuli and capillaries
in both rami from the first onwards. Gills rudimentary and represented by small
swellings of the abdominal notopodia and sUghtly larger triangular projections of
the superior edge of the neuropodia (fig. 28. 2. d.) A lateral organ between notopodia
and neuropodia. Genital pores present behind the lateral organs in mature in-
dividuals on 7-20 segments starting on the second abdominal. Abdominal neuro-
podia have long rows of hooks which almost meet in the mid-ventral line. Hooks
(fig. 28. 2. b, e) with short hoods less than twice as long as broad and arcs of about
five denticles above the main fang.
Type locality : Norway.
Records: South West Africa (28/16/s) ; Cape (from 32/1 7,'d, 33'17/vd to
34/20,6 and 33,25/s; Natal (2g/3i;d) ; Mocambique (2434,5).
Distribution : CosmopoUtan from the Arctic to the Antarctic. .\t all depths
(intcrtidal to abyssal) in cold seas but usually below 50 metres in tropical seas.
Habitat sand, mud.
' Differs from A', hemipodm Hartman 1945 in the structure of the hooks.
6oo rOLYClHAr.TA OF SOUTHERN' AFRICA
LEIOCHRIDES AuRcmr, i()\.\
Thorax willi an achactous pcristomium followed by 12 setigcrs with capillary
setae in both rami. Abdomen with long-shafted hooded hooks in both rami. Re-
tractile branchiae with a few filaments arise from the superior ends of the posterior
neuropodia.
Type species: Leiochridcs luutralis Augencr, 191 4.
Leiochrides africanus Augener, iqiS
(fig. 28.2.C-h)
Leiochrides iifricantis Aus^cncr. I')i8: 472, pi. 7 figs. 199-200; text-fig. 74; Day, 1955: 423.
Body about 60 mm. long for 200 segments. Prostomium (fig. 28. 2. e) conical,
two-ringed. Abdominal hooks (fig. 28. 2. g, h) with long hoods and two arcs of one
to two large and then three to four smaller denticles above the rostrum. Retractile
gills arise between the notopodia and neuropodia of posterior abdominal segments
and have two to four lobes (fig. 28. 2. f). In posterior abdomin.il segments the rows
of notosctae are broadly separated dorsally.
Type locality : Sctte Gama, Brazzaville Congo.
Records: Cape (34/18/i) ; Natal (29/31/5).
Distribution : Tropical western Africa from Ghana (s) to Angola (i).
MEDIOMASTUS Hartman, 1944
Body thread-like. Thorax with an achaetous pcristomium followed by 10 sctiger-
ou.s segments. The first four setigcrs bear winged capillaries in both rami and the
last six bear long-shafted hooded hooks. No genital setae. Abdominal segments
with hooded hooks in both rami. Gills entirely absent.
Type species : Mediuiuastus calijoniiensi.s Hartm.ui, 1944.
Mediotnastiis capensis Day, 196 1
(fig. 28.2. n-p)
Mediomasliii ea/icnui Day. 19G1 : 318, fig. 11 a-d.
A thread-like worm about 14 mm. hmg with 60 segments. Prostomiimi (fig.
28. 2. n) biannulate, conical, witlmut eyes. Buccal segment achaetous. Segments
two to five biannulate with lajjillaries in both rami. Segments 6-1 1 narrowed,
biannulate to multiannulatc with long hooded hooks in both rami. First abdotninal
segment stouter than last thoracic. Abdominal segments flattened vimtrally and
thin-walled dorsally so that the large brown faecal pellets are visible giving a
beaded effect. No gills. Parapodia poorly marked with short notopodial and
ncuropodial rows of hooks equally spaced around the segments. Hooks (fig. 28.2.0, p)
CAPITELLIDAE 6oi
with two arcs of equal teeth above the main fang, the first with eight and the second
with three to four teeth.
Type locality : Saldanha Bay, South Africa.
Records: South West Africa (26/15/s and 28/16/s) ; Cape (from 32/17/s, d to
35/20,'d and 33/2 7 /s) ; Natal (30/30/s, 29/3 i/s, d).
Distribution : Endemic.
HETERO MASTUS Eisig, 1887
Body filiform. Thorax with an achaetous peristomium followed by 1 1 setigerous
segments. The second to sixth bear capillaries in both rami and the seventh to
twelfth bear long-shafted hooded hooks in both rami. Abdomen with hooded hooks
in both rami throughout. In the abdomen the gills are indistinct extensions of the
superior ends of the neuropodia and the posterior segments are campanulate or
strobiliform. Pygidium with a median ventral cirrus.
Type species: Capitella filiformis Claparede, 1864.
Heteromastus filiformis (Claparede, 1864)
(fig. 28.3.a-d)
Capitella filiformis ClaparMe, 1864: 509, pi. 4, fig. 10.
Heteromastus filiformis : Fauvel, 1927: 150, fig. 53 a-1 ; Hartman, 1947: 427, pi. 52 figs. 1-4.
Body filiform, up to 100 mm. long but only i mm. in diameter. No marked
distinction between thorax and abdomen (fig. 28.3.3). Prostomium conical. Thorax
biannulate and tessclate. Anterior abdominal segments long and cylindrical, later
ones shorter and posterior ones campanulate or strobiliform with short uncigerous
rows on projecting lobes (fig. 28. 3. b). Branchiae as indistinct superior projections
of the neuropodia from the 8oth segment onwards. Genital pores from the ninth to
twelfth thoracic segment. Thoracic hooks (fig. 28. 3. c and d) with long hoods and an
arc of about six denticles above the main fang; abdominal hooks with short hoods
and only three to four denticles above the main fang.
Type locality : Mediterranean Sea.
Records: Cape (34/23/e) ; Natal (29/3 1 /i) ; Mocambique (23/35/s).
Distribution : North Atlantic from Greenland (s) and Sweden (d, vd) south to
New England, North CaroUna (i) and the Gulf of Mexico (i) ; Morocco (s, d) ;
Mediterranean (s) ; Persian Gulf (s) ; North Pacific from the Bchring Sea to N.W.
Japan and Southern California.
L
603
I'OLVCHAKTA OF SOUTHKRX AFRICA
mnnn
Fig. 28.;^. Utiaouiaslus filifnrmis. (a) Lateral view of tliorax and anterior abdomen, [is) T/S
posterior abdominal segment, (c) Face view of thoracic hook, (d) Profile of same.
Da\vhtimhus t(uhicu\. (i) T/S anterior abdominal segment, (f) T/S posterior abdominal
segment. (*;) and (11 j Profile and face view of hooded hook. Daivbraiichin bifuitltfnu
(i) Entire worm (natural si/ri. fj) T'S anterior abdominal segment, (k) Postcrii.tr abdo-
minal segments bearing gills, (r.) and (m) Face and profile views of hooded hooks. Pm-
helrromaslu'i tenuis, (n) Prolile of thoracic hook. {<>) Profile of abdominal lumk (from
Monro), (p) Lateral view cjf anterior end. (q) T/S of abdominal segment.
CAPITELLIDAK 603
D ASYBR ANCHUS Gruhe, 1850
Thorax with an achaetous peristomium followed by 13 setigers bearing winged
capillaries in both rami. Abdominal segments with hooks only in both rami.
Retractile branchiae in the form of branched filaments arising from the superior
edges of the neuropodia of posterior abdominal segments.
Type species: Dasjmallus caducus Grube, 1846.
Key to Species
I Hoods of hooks less than twice as long as broad. Denticles above the main fang small and
equal (fig. 28.3. g, h). Branchiae from 20th abdominal segment . . D. caducus
- Hoods of hooks about three times as long as broad. Denticles above main fang unequal,
the first row consisting of one to four large ones (fig. 28.3. 1, m). Branchiae from
abdominal segment 40 ......... /). bipartitus
Dasybranchus caducus (Grube, 1846)
(fig. 28.3.e-h)
Dasymaltm caducus Grube, 1846: 1 66.
Dasybranchus caducus : Eisig, 1887 : 823, pi. 16 figs. 1-6, 8-12, pi. 32 figs. 1-4.
A large worm reaching 300 mm. in length and 10 mm. in diameter. Prostomium
short and conical. Thoracic segments biannulate and tesselate. Genital pores start
in the intersegmental constriction of setigers 12 and 13 (i.e. before last thoracic
segment) of sexually mature specimens. Abdominal parapodia as low ridges bearing
rows of hooks. A narrow mid-dorsal gap between anterior abdominal notopodial
rows of hooks (fig. 28.3.6). Retractile gills (fig. 28.3.f) above the neuropodia from
abdominal segment 20 onwards and reach a maximum of 20 branches. Hooks
(fig. 28.3. g, h) with hoods not more than twice as long as broad and with three to
four rows each with about eight equal denticles above the main fang.
Type locality : Mediterranean.
Records: Natal (29/31/i and 27/32/i) ; Mocambique (26/32/i and 23/35/e).
Distribution: Records confused - certainly present in Mediterranean (i), Red
Sea (i) and tropical Indian Ocean south to Madagascar (i, s).
Dasybranchus bipartitus (Schmarda, 1861)
(fig. 28.3.i-m)
Oncoscolex bipartitus Schmarda, 1861 : 55, pi. 26 fig. 205.
Dasybranchus bipartitus : Day, 1953 : 424, fig. 3 f and g.
Body (fig. 28.3.1) up to 300 mm. long by 6 mm. with more than 200 segments.
Prostomium short and conical. Thoracic segments biannulate and tesselate.
Nephridial pores from setiger 9-10. Abdominal parapodia as low ridges bearing
rows of hooks. A broad dorsal gap between right and left notopodial rows of hooks
and a small mid-ventral gap between the long rows of neuropodial hooks on anterior
abdominal segments (fig. 28.3.J). Nephridial pores continue to the 14th and 15th
Gii4 P0I.YC:MAKT.\ of SOL'THKRX AIRICV
abdominal segment. Posterior segments with short rows of hooks in both rami and
wide gaps between them. Retractile gills from abdominal segment 40, each \vith
a maximum of 13 branches in large specimens (fig. 28.3.!;). Hooks with hoods
three to five times as long as broad and three arcs of teeth decreasing in size above
the main fimg with one to four in the first row and four to eight in the second row
(fig. 28.3.1, m').
Type locality : Cape of Good Hope.
Re(:;ords : C^apc (from 29'i6 i to 34'i8''i, s and 33,'25/s).
Distribution : Records confused - certainly present in English Channel (s) and
probably Sweden (d).
SCYPHOPROCTUS C.i ivUi\ 1906 (emend. Day, 1962)
Thorax with 14-16 segments of which the peristome and the next segment arc
achaetous and the following 12-14 bear capillary setae in both rami though hooks may
replace capillaries in the neuropodium of the last thoracic setiger. Abdomen with
hooded hooks in both rami. No branchiae. Terminal segments of abdomen fused
to pvgidium to form a slanting plate on which the anus opens above a pair of anal
cirri. Notosetac of last few segments modified to form stout spines without hoods.
Type species; Scjphoproctus djihouticnsh Graxier, 1906.
Key to species
I Fourteen thoracic setigers. Anal plate with two rows of notopodi.Tl spines . S. steinitzi*
- Twelve thoracic setigers. .\nal plate with ten to twelve groups of notopodial spines
S. djiboutiensis
Scyphoproctus djiboutiensis Gnnkr, 1906
Sg/iltoproclus djiboulicmii Gravier, iyo6; i8i, pi. 3 figs. 200-204, text figs. 345-348; Fauvcl, 1933;
373. fig- 194 a-b.
Body slender, cylindrical, 23-30 mm. long. Prostomiiim a blunt cone without
eyes. Peristome and the next segment achaetous. TwcKe biaimulate thoracic
setigers with capillary setae in both rami, though the capillaries in the neuropodium
of the last thoracic segment may be partly or completely replaced by hooks. Abdo-
men of 70 or more thin walled segments without parapodial projections or any sign
of gills. The rows of hooks arc equal in length and excnly spaced in anterior seg-
ments, but the noto]3odial rows approach one another dorsally and almost fuse just
before the anal plate. At the same time the liooks, which are normally hooded
and denticulate over most of the abdomen, lose their hoods and denticles and are
enlarged to form stout spines. The anal plate is formed of a doubtful number of
segments fused to tlie pygidium. It slants posteriorly and is flattened dorsally with
the anus just above a pair of large terminal anal cirri. The margin of the plate is
beset with 10-12 groups of spines. The number of spines in each group decreases
posteriorly. The anterior margin has a continuous palisade of about 20 spines
CAPITELLIDAE 605
formed by the fusion of right and left groups but thereafter the groups decrease
from four or five to only one or two just before the origin of the anal cirri. There
is only one row of neuropodial hooks on the anal plate obviously corresponding to
the palisade of dorsal spines on its anterior margin. The capillary setae are quite
plain with narrow wings. Each hook has a short hood, almost as broad as long, a
well marked rostrum or main fang bent at right angles to the shaft and surmounted
by a crest of four rows of subequal denticles.
Type locality : Djibouti in 20 metres.
Records: Mocambique (24/34/s).
Distribution: Tropical Indian Ocean from the Gulf of Aden (s) to Ceylon (s).
PARHETEROMASTUS Monro, 1937
Thorax of 12 segments of which the pcristomium or first segment is achaetous,
the second to fifth segments bear capillaries only and the last seven have only long-
hooded hooks. Abdominal hooks with shorter hoods than those of the thorax.
Abdominal tori poorly developed and branchiae lacking. Pygidium with a single
short cirrus.
Type species : Parheteromastus tenuis Monro, 1937.
Parheteromastus tenuis Monro, 1937
(fig. 28.3.n-q)
Parheteromastus tenuis Monro, 1937a : 536, fig. 2 a-e.
Body slender, up to 50 mm. long by 0-5 mm. broad for 140 segments. Prostomium
conical and lacks eyes. Thorax (fig. 28. 3. p) slightly swollen with an achaetous
peristomial segment followed by 1 1 setigerous segments of which the first four bear
capillary setae in both rami and the last seven bear hooded hooks in both rami.
Change from thorax to abdomen not well marked, the main difference being the
hooks which have shorter hoods in the abdomen (fig. 28.3.0) than in the thorax
(fig. 28. 3. n). Parapodial ridges (fig. 28. 3. q) poorly developed. Branchiae absent.
Pygidium with a single short anal cirrus.
Type locality' : Maungmagan, Burma.
Records: Mocambique (26/32/i).
Distribution: Burma (i).
6oC POI.VCHAETA OF SOUTHF.R\ AFRICA
Family ARENICOLIDAE |.>hnst(in, 1835
Body elongate with numerous annulatcd segments. The head consists of a small
prostomium without appendages, a peristome with a large cversible proboscis which
is unarmed and then another achaetous segment. Behind this is a trunk of numerous
setigerous segments of which the first few lack branchiae and the remainder bear
branching gills behind the notopodia. A posterior tail region of achaetous segments
is sometimes present. Oesophagus with one or more pairs of glandular caeca. A
single pair of hearts. The first septum often bears a pair of septal pouches. Para-
podia small but biramous ; notosetae are simple capillaries and neurosetae are
acicular hooks without hoods.
Records from southern Africa
Abarenicola ajjuiis africana Wells
as Arenkola assimilis var. affinis Ashworth . iq\\'i
Abarenicola gilchristi Wells ....
as Arenicola assimilis war. affinis Ashworth
partim ...... iqCIi, 44Ci
as Arenicola marina (non Linnaeus) . . ii\Vi
as Arenicola claparedii (non Le\-insen)
Ashworth, 1910 . . . . . Wi
Arenicola loreni Kinberg .... 3Ni, i8C'i, igNiCi,
36Ci, 44Ci
Brancluomaldane vinccnti Langeriians
as Prolocapilella simplex Berkeley . . . 44CIi
BIOLOGICAL NOTES
The arenicolids are particle feeders and apart from the small and primitive form
Branchiomaldane which lives in muddy rock crevices, they live in muddy sand in the
intertidal zone. Their physiology has been described by Professor G. P. \V'ells in a
long series of papers. Arenicola and the related genus Abarenicola both construct
U-shaped burrows by means of an eversible proboscis. Power for pumping water
through the burrow is supplied by the contraction of the circular muscles of the body
and for burrowing it is transmitted to the proboscis cavity by the hydrostatic pressure
of the coelomic fluid. The intervening septa have pouches which bulge in consequence.
Three variations on this theme are illustrated in tlie accompanying diagrams.
The usual indication of an Arenicola burrow is a funnel-like depression which
marks the head end of the tube and earthworm type castings at the tail end. But
this is not always the case. Arenicola loreni which is one of the largest species and
may reach a length of one metre, burrows in estuarine sandbanks where the sub-
surface layers are semi-lic]uid. The head end of the burrow is marked by the usual
depression but the tail end lacks castings. Material \oided by the worm is in a semi-
liquid state and is squirted out a considerable distance. Abarenicola gilchristi which
is seldom more than 250 mm. long lives on more exposed shores. It ejects fairly
solid castings but these are carried avsay by wave action. Only the dwarf form
ARENICOLIDAE 607
Abarenkola affinis afrkana makes typical castings on South African shores. It hves
in the black mud of salt marsh pools high up on the shore where there are no waves.
THE MAIN DIAG.NOSTIC CHARACTERS
Important revisions of the family will be found in Ashworth {1912) and Wells
(1958). Wells recognised four genera, namely Abarenkola, Arenkola, Branchwmaldane
and Arenkolides, the last of which does not occur in southern African seas.
Head and internal organs. The prostomium varies in shape and statocysts may be
present or absent. If present they may be closed sacs or have ducts leading to the
exterior. The first double septum or "diaphragm" may be muscular and have a
pair of large, backwardly directed septal pouches ; alternatively the third septum
may have baggy projections extending forward. The number of glandular caeca
on the oesophagus is a most useful character ; there may be only one pair or as
many as 20. The number of nephridia varies from two to thirteen pairs and the first
may appear as early as setiger 4 or as late as setiger 15.
Body regions, number of segments and number of annuli. The presence or absence of
a posterior achaetous tail region is of generic importance. The number of segments
in the trunk region is very constant in Arenicola and Abarenicola but not in Branchio-
maldane and Arenicolides. Apart from Branchiomaldane which has two annuli per seg-
ment, all genera have five annuli per segment in the branchiferous region but a
reduced number immediately behind the head.
Parapodia, branchiae and setae. The notopodia are erect and uniform in structure
and the neuropodia are ventro-lateral ridges which vary in length. The short
neuropodia of Abarenicola provide an easy means of separating this genus from
Arenicola and Arenicolides which have long neuropodial ridges in the branchiferous
region. The branchiae are essentially similar in structure but the segmental position
of the first gill is important at the specific level. The notosetae are capillaries with
smooth blades in Branchiomaldane and spinulose blades in other genera. The neuro-
podial hooks have poorly marked denticles on the crest except in Branchiomaldane
where well marked teeth are present. The absence of a hood distinguishes the hooks
of the Arenicolidae from those of the Capilellidae.
Key to Gener.\
Branchiferous segments with two annuli (fig. 29. i. a), .\nimal small, not exceeding 40 mm.
BRANCHIOMALDANE (p. 608)
Branchiferous segments with five annuli (fig. 29.i.f). Animal large, exceeding 40 mm. . 2
No achaetous tail region. Gills never start before setier 12 (neuropodia long and approach
the mid-ventral line) ARENICOLIDES*
Achaetous tail region present. Gills start on setiger 7 or 8 . . . . . . 3
Neuropodia long and approach the mid-ventral line in the branchial region ifig. ag.i.i).
One pair of oesophageal caeca ....... ARENICOLA (p. 608)
Neuropodia short and are alwa>^ widely separated ventrally (fig. 29. LnV P'ive or more
pairs of oesophageal caeca ....... ABARENICOLA {p. f>io)
(k.8 Pt)I.VC:ll.\E TA of SOUTHI'.RX AFRICA
BRANCHIOMALDANE Lanscili.uis, 1881
Small thread-like worms 20-30 mm. long with numerous biannulatcd segments.
Prostomium a simjile cone with eves but without ntocysts. One pair of oesophageal
caeca. Body divided into an anterior region ofabout 14 -20 abranchiate segments and
a posterior region of numerous segments most of which bear a few branchial filaments
behind the notopodium. No achaetoirs tail region. Notosetae are narrow-bladed
capillaries throughout. Xcurosetae are long-shafted dentate hooks witliout hoods.
Type specifs : BruiHliinmaldane vincenii Langerhans, r88i.
Branchiomaldane vincenti Langerhans, 1881
(fig. 29.1 .a-cj
Branchiomaldane lincenli Langerhans, 1881 : 1 16. pi. 5 fig. 21 ; Fauvel, nj27 : 166, fig. -,8 m. p.
Prolocapilella iim/ilcx Berkeley & Berkeley, 1932 : 6tj9 ; Fiartman, 1947 : 440, pi. 57 figs. 3-4.
Body (fig. 29. 1. a) small and thread-like, about 10 mm. long with 30-50 biannulate
segments. Prostomium (fig. jq.i.b) an ovoid cone with two groups of ocelli. Xo
otocysts. Proboscis globular. One pair of oesophageal caeca. Peristome and thcne.xt
seginent achaetous. Anterior region of 14-20 abranchiate segments each with the
anterior setigerous annulus more prominent than the second one. Posterior region
with 15-30 biannul.ite segments (fig. 29.1.C) all of which bear parapodia and most
of them notopodial branchiae which arise as three to five simple filaments behind
the notosetae. Xotosetae ffig. 29.1.C) are capillaries throughout with flattened
blades and minutely spinulose tips. Xeurosctae are hooks (fig. 29.i.d) with three to
four teeth abo\'e the rostrum but witluau hoods. Two pairs of nephridia open on
setigers 5 and 6. Hermaphrodite.
Type locality : Clanary Islands.
Records: Cape (33/18 i and 34'18,'i).
DiSTRiBUTiox : North Atlantic from the English Cihannel (i) to the Clanary Is.;
N.E. Pacific to .S. CaUfornia.
ARENICOLA Lamarck, 1801
Body large ("100-400 mm.) and divided into a trunk region ofabout 20 setigeious
segments and a tail region of numeidus achaetous and apodous segments. Pro-
stomium small and retractile. Statocysts ]3rcsent. Proboscis large, papillose and
eversible. Diaphragm muscular with a pair of backwardly directed septal pouches.
Oesophagus with one pair rif glandular caeca. Fi\e to seven pairs of nephridia the
first openmg on setiger 4 or 5. Sexes separate. Setigerous segments after the first
two to three di\'ided into five annuli. Branched gills arise behind the notopodia
from setiger 7 to the end of the trimk. .Xotosetae erect and bear bundles of capillaries
with serrated and spinulose blades. Xeuropodia as low vertical ridges which are
elongated in the branchiferous region and almost meet on the mid-ventral line ; each
bears a row of long-shafted hooks with poorly marked denticles on the crest.
Type species : Lumhrkus marinus Linnaeus, 1 758.
AREMCOLIDAE
603
Fig. 29.1. Branchiomaldane vincenti. (a) Entire worm (four times natural size), (b) Dorsal view
of head region, (c) Lateral view of a branchiferous segment, (d) Neuropodial hook.
(e) Notopodial capillar\'. Arenkola loveni. (f) Entire worm (half natural size), (g) Dis-
section of anterior end. (h) Dorsal view of head and extruded proboscis, (i) T/S branchi-
ferous segment, (j) Neuropodial hook, (k) Notopodial capillary. Abarenicola qffinis africana.
(l) Entire worm (half natural size), (m) Dorsal view of head and proboscis, (n) T/S
branchiferous segment, (o) Dissection of anterior end. (p) Neuropodial hook, (q) Noto-
podial capillary.
6io l'OI,\C:ilAl. TA Ol' S(1L Ttll'.RX AIRK^A
Ki;v lo Species
I Seventeen setigerous segments and it pairs of gills .... A. bombayefisis*
- Nineteen setigerous segments and 13 pairs of gills ...... 2
■2 Six pairs of nepliridia on sctigers 4 to 9. Septal pouches small. .Statocyst open A. ty^arina*
- Five pairs of nepliridia on setigers 5 to 9. Septal pouches enormous. Statocyst closed 3
3 No prcN'cntricular dilatations of the dorsal vessel . . .A. loi'eni loz'eni
- .-\ pair of pre\enticular dilatations to the dorsal vessel . A, loi'eni sudaustraliensis*
Arenicola loveni loveni Kinbcig, 1867
(fig. 29.1. f-k)
Arrniciila liiient Kinherg. 1867: 3",5; Kinbcrg. 1857 igio: 73, pi. 29 fig. i; .\shv\orlh, 1910:
I. pi. I figs. I- 8 ; Wells, 1962 : 34!!, pis. 2 and 4.
A very large species (fig. 29. 1. f) with a long achaetous tail region reaching a
length of 800 mm. Prostomium (fig. 2g.i.h) trilobed with the incdian triangular
lobe broader than the \'-shaped pair of lateral lobes which support it. A pair of
closed statocysts each containing one secreted statolith. The tips of the enormous
septal pouches (fig. ag.i.g) reach back to the third diaphragm or further. Oeso-
phagus with one pair of glandular caeca. Five pairs of nepliridia opening on segments
5 to 9. Inner end of dorsal septal vessel moves from subintestinal to dorsal vessel
in setiger 12. Thirteen pairs of branchiae starting on setiger 7. Notosetae (fig.
29. 1. k) with lateral toothed-crests and a narrow, finely spinulosc lamina. Neuro-
podial hooks (fig. 29.1.J) with a faint indication of a denticle or none. Xeuropodial
ridges in the branchial region long and almost meet on the mid-ventral fine (fig.
29.1.1). The nominate form, ^-1. loveni loveni, lacks preventricular dilations of the
dorsal vessel.
Type LOCALrrY : Durban, .South .-\frica.
Records : Clape (from 33 18, i to 32 '26/e) ; Natal (29 31 'i, s) ; habitat : muddy
sand of protected bays and estuaries ; L.\V.N. to L.W.S.
Distribution; .1. /. loveni is endemic; .1. /. Mulamlraliensis is known from South
Australia.
ABARENICOLA Wells 1958
Body with a trunk region of 19-20 setigers and an achaetous tail region. Pro-
stomium nrin-retractile, in the form of a triangle with lateral extensions of its
(anterior) base. Statocysts either present with external openings or absent. Pro-
boscis large, eversible and papillose. First septum weak and lacks septal pouches ;
third septum with a pair of large, forwardly directed baggy projections. Five or
more pairs of oesophageal caeca of which the first is the largest. Five to six pairs
of nephridia starting on setigers 4 or 5. Sexes separate. Setigerous segments after
the first two to three divided into five annuli. Branched gills arise behind the
notopodia from setiger 7 or 8 to the end of the trunk. Xotosetae with spinulose
ARENICOLIDAE 6il
blades. Neuropodia as lateral ridges which are well separated vcntrally in the
branchiferous region. Neurosetae are long-shafted hooks with poorly marked
denticles near the apices.
Type species: Arenicola claparedii Levinsen, 1883.
Key to Species
I Thirteen pairs of gills starting on setiger 7. Eight to ten pairs of oesophageal caeca (fig.
29.1.0). (The first three setigerous annuli very prominent) . A. affinis africana
— Twelve pairs of gills starting on setiger 8. Fifteen to twenty-one pairs of oscophageal
caeca (fig. 29. 2. b) .......... A. gilchristi
Abarenicola affinis africana W ells, 1963
(fig. 29.1.1-q)
Abarenicola affinis africana Wells, 1963 : 142, fig. 6 a-b, pi. i and 5.
Arenicola assimilis var. affinis Ashworth, 191 1 : 18, text-figs. 4-5 (partira).
Body (fig. 29. 1. 1) small, less than 100 mm. long. Trunk region of 19 setigers with
13 pairs of well branched gills starting on setiger 7. Caudal region short with
prominent papillae. The first three setigerous annuli powerful and prominent,
intervening annuli small. Statocysts with ducts to the exterior and contain numerous
foreign statoliths as well as secreted ones. Posterior region of oesophagus (fig. 29.1.0)
with one pair of long delicate caeca followed by six to nine short ones in front of
the hearts. Five pairs of nephridia opening on setigers 5-9. Inner end of dorsal
septal vessel moves from subintestinal to dorsal vessel in setiger 13. Hooks (fig.
29. 1. p) with several poorly marked denticles. Notosetae (fig. 29.i.q) with finely
spinulose blades.
Type locality : Luderitzbucht, .South West Africa.
Records: South West Africa (26/1 5/i) ; Cape (33/18/i). Habitat: mud above
M.S.L. in protected bays.
Distribution : Endemic.
Abarenicola gilchristi Wells, 1 963
(fig. 29.2.a-c)
Abarenicola gilchristi Wells, 1963 : 147, fig. 6c, pi. 2 and 5.
Arenicola assimilis var. affinis .'Kshworth, 191 1 : 18, text-figs, 4-5 (partim) ; Day, 1955: 427.
Arenicola claparedii : (non Levinsen) .Ashworth, 1910; 351.
Body 150-250 mm. long. Trunk region with 19 setigerous segments and 12 pairs
of lightly branched gills on setigers 8-19. Caudal region fairly long with low papillae.
The first three setigerous annuli (fig. 29.2.3) prominent. Statocysts with ducts to
the exterior and contain a few (five to six) foreign and secreted statoliths. Oeso-
phagus (fig. 29. 2. b) with one large pair of caeca followed by 14-20 short pairs
arranged in double rows in front of the hearts. Five pairs of nephridia opening on
Gl2
roi,vc;iiAF, lA CM- southern ai ric:a
sctigcrs 5-q. IiiiKT end of dorsal septal \essrl ni(ncs fi(im suh-iiilislinal lo doisal
vessel in setiner 12. Hooks (fig. 29. 2. c) \\itli sexcral jioorly maikctl denticles.
Type lo(::alitv : Cape Peninsnla, South Aliie.i.
Records: South West Africa (26/15/i) ; Cape (32/18/i to 34/19/i) -habitat:
intertid.il sands lielow .M.S.L. on slK-ltered shores.
Distribution : Endemic.
Fif:. 20.2. Aliiimiiuilii i;il(li)i\n. (A) Lateral view of anterior end.
(li) Di»ettiijn ofanleriur end. (c) NcLiropodial liook.
MALDANIDAE
613
Family MALDANIDAE Malmgrcn, 1867
Body cylindrical with a relatively small number of greatly elongated segments.
Body regions not marked. Prostomium poorly defined and fused to the buccal
segment. Prostomial appendages entirely absent but nuchal slits well marked and
numerous eye-spots may be present. The prostomium may be followed by a
crest or surrounded by a flattened cephalic plate with a raised rim on the
dorsum of the buccal segment. Mouth ventral with an unarmed but papillose
proboscis. Parapodia biramous but poorly developed with a tuft of capillaries in
the notopodium and a vertical series of hooks in the neuropodium. Parapodia are
situated near the anterior margins of the first few setigers, in the middle of setiger 8
and near the posterior margins of all succeeding segments. Up to 10 achaetous
preanal segments may precede the pygidium which may be conical, truncate,
funnel-shaped or petaloid. The anus may be dorsal, terminal or sunk in the funnel.
In the first three setigers the neurosetae may be modified to form simple spines or
may be absent. Tube of cemented mud or sand grains.
Records from southern Africa
Asychis capemii Day
Axiothella jarli Kirkegaard
Axiothella quadrimaculala Augener
Clymenura tenuis (Day) .
as Leiochone tenuis Day
Euclymene glandularis (Day)
as Clymene glandularis Day .
as Praxillella praetennissa var. capensis
(non auctorum)
as Clymene praelermissa var. capensis
(non auctorum)
Euclymene lombricnides (Quatrefages)
as Clymene lombricoides Quatrefages
Euclymene luderitziana Augener
Euclymene lyrocephala (Schmarda)
as Clymene lyrocephala Schmarda
Euclymene mossamhica (Day)
as Clymene mossamhica Day
Euclymene natalensis (Day)
as Clymene natalensis Day
as Clymene sp. .
Euclymene oerstedi (Claparede)
Euclymene cf. quadrilobala (Sars)
Euclymene sp. .
Gravierella multiannulata Fauvel
as 1 Gravierella sp.
Johnstonia clymcnoides Quatrefages
5'Ci
48AS, PCs
45Pi, ?5iCis
45NiPi
51CS
44Ci
i3t;i, 35Ci
36Ci
51CS
35Ci,36Ci,44CiWi,45Pi
26WS, 44Ci, 51CS
lAVi, i6\Vi
4Ci
45Pi
5iCd
45PiNi
4oNi
48Cd
56Cd
56CS
27Mi, 45Pi
4oPi
48\Vd
(U4
'OI.YCHAKTA OF SOL TIIERN" AFRICA
Johnstonia knxsna Day .
l.umhrich'mene cvimdricauda Sars
Lumhriclymene minor Arwidsson
Macroclymene mnnilh Fauvcl .
Macroclymcne saldanha (Day) .
as Clymene iaidanha Day
Maldane stirsi Malmgren
Maldanelh capeiisis Day
Maldanella fibrillata Chamborlin
Mkromaldane (forma juvenis) .
Nicomache lumhricalis (Fabricius)
as Atcomache capensis Mcintosh .
as \icomache lumbrica/is \ar. capetnis
Mcintosh.
? as (dviiit'iic nucrn(ej)h(tld Sclimarda
Aicomaiiie inosianihicii Day
Petaloproctus terricola Qjaatrcfages
as .Mcomaclie niacintoshi Marenzellcr
as PelaloprncUis macintoshi (Marenzellcr
PraxUlella cf. affinis (Sars)
Piaxillella caperuis (Mcintosh)
as Praxilla capensis Mcintosh
Piaxillella praeiermissa Malmgrcn
Rhodirie gracilior Arwidsson
Inccrtac sedis
Maldanid gen. cl ^->.? .
44C:i, 5iC:s
56(;d
^4As
44C;i
32\dCs, 5iXd, — Nsd, Ps
5iCs
44C;i
33Ci, 36C:i, 44C;i, TjoCs
loCi
i3C:i, 35C'i
4C;i
40 Pi
44C1, 48\Vs
I iW'i, i3Ca
26\Vs
56C;d
4 1 Cs
loC^s
?32Cd
5iCsd, 56C:d
5iC;d
BIOLOtaCAL .NOTES
The maldanids are higlily specialised burrowcrs feeding on organic particles
buried in the mud. Those whose habits have been investigated appear to burrow
liead downward cementing the surrounding materials together to form a fairly com-
pact tube. The pygidium plugs the entrance of the tube and the anal cirri are
probably sensory.
.Xicnmache and P(i/iln/>rnctii'< burrow along the sides of boidders half binied in sand
or gra\rl so that the tube is attached to the stone. pAiclrmene, Clvmenella and Piaxillella
burrow in open sandlianks and their tubes are relatively fragile. Maldane and
Asychis are found in softer muds and their tubes are often bulky. Rlwdinr which is
dredged on the sanii- type of substrate has a thin horiu' tube.
Inigation of th<' burrow is brought about l:>y ]ieristallic action of the elongated
and highly coiUractile segments. According to Dr. C'harlolte Mangum's obser-
vations, the water i urrent is forced down Irom the tail towards the head and assists
in feeding. It is not surprising that in Johnstonia where branchial filaments occur,
they are found near the tail end.
The eggs are incubated in mucus cocoons .ittachcd to the entrance of the burrow
and till' larvae escape .U a fairly late stage of development.
MALDANIDAE 615
THE MAIN DIAGNOSTIC CH-ARACTERS
Useful reviews of the family will be found in Arwidsson (1906) and Fauvcl
(1927)-
In general the genera and species arc well defined but complete specimens are
essential for accurate diagnosis. Several very dubious species have been founded
on fragments.
The head. The prostomium is contractile and its shape is of limited value ; the
presence of ocelli is however, worth noting. The prostomium may extend back as
a median ridge or cephalic keel which is well marked and arched in genera such as
Maldane or undeveloped in Eucljmene and its allies. On either side of the keel there
is a pair of nuchal slits which may be straight and parallel or curved and divergent
anteriorly particularly if the prostomium is broad. The whole dorsal surface of the
head may form a flattened cephalic plate enclosed by a raised margin or cephalic rim.
The development of the cephalic rim is a useful character. In some species it is
rudimentary, in others it is high with a scalloped, notched or lobed posterior margin ;
but often there is only a pair of lateral notches and a single posterior one. Arwidsson
(1906) has also used the distribution of the papillae on the margin of the eversible
proboscis but as these are seldom visible other workers have not followed his example.
The body and number of segments. In most genera there are 18-24 setigerous seg-
ments followed by i-io achaetous preanals and then the pygidium. In these
genera the number of setigers and preanals is surprisingly constant but there are
other genera such as Macroclymene and Gravierella with 30-70 setigers and in these
the number of segments is not constant and in the latter genus a group of achaetous
segments may precede the last few setigers. The anterior segments up to about the
eighth arc usually short, the middle ones long and the last few again shorter. The
exact proportions vary with the state of contraction but the numbers and relative
sizes of achaetous preanals are useful. The parapodia are usually near the anterior
margins of the first few segments, then move back towards the middle of setigers 5-8
and from the ninth onwards they are at the posterior margin. The junction of
setigers 8 and 9 is very indistinct. The epithelium around the parapodia is usually-
glandular. The glands spread to form thickened circular bands or rings around the
anterior margins of the first few setigers or even o\er the whole surface setigers 3-7.
In the posterior region, however, the glands are limited to the swollen neuropodia.
In many species a ventral glandular streak extends from setiger 9 to the pygidium and
in some species it can be traced forwards to sedger i . Occasionally a dorsal glandular
streak is present on middle segments. In genera such as Rhodine the anterior margins
of some anterior segments extend forward as membraneous collars surrounding the
posterior margin of the preceding segment. Posterior segments may have posterior
collars.
Pygidium and anus. The shape of the pygidium and the position of the anus
provide valuable characters. In its simplest form found in Clymenura, the pygidium
is bluntly conical with a terminal anus sometimes closed by a swollen lobe or ventral
valve. Three hnes of development occur. In the first anal cirri appear ; when these
become numerous a circle of cirri is formed from which the anal cone protrudes.
6i6 I'OI.VCHAETA OF SOUTHERN AFRICA
This is the condition in Praxilklla. Fnrthcr development results in tlie formation of
an anal funnel rimmed with cirri into which the anal cone sinks and the ventral valve
is then reduced or lost. This ultimate sta<j;e occurs in Euclymcne and several other
genera. A second line of development seen in Lumhiiiivmene is the formation of a
flattened shield below the anus ; in Nicomache the posterior end becomes obliquely
truncate and the anus is dorsal. The third line of development is seen in Pftaloproclus.
Here the pygidium forms a petaloid plate around the central anus.
Stiae. 'I he notosetae are various forms of capillaries and seldom pro\'ide useful
characters. In juveniles, hastate setae are often present. In adults the tips may be
cylindrical or flattened to form marginal wings and these inay become incised to
form fine spinules, the whole tip then resembling a very slender feather. Such
feathered setae are more common in posterior segments. With the exception of
Rhodine the neurosetae are arranged in single rows. In juveniles the neurosetae first
appear as S-shaped hooks with a crest of denticles and a swelling in the shaft. In
the adult the basic form has a long shaft, a toothed crest and a tuft of tendons below
the main fang. In most genera the denticles or teeth abo\e the main fang are in a
vertical scries but in Maldane and Asrchii the denticles are arranged in transverse
arcs. Neurosetae may be absent on setigers 1-4 but are usually present and reduced
to acicular setae either with a few \estigial denticles or none at all. In this case the
neurosetae inay be represented by a single, smooth, curved spine.
Arwidsson (igo6) has divided the Maldanidae into fi\e subfamilies whose main
characters are summarised below.
LUMBRICLYMENINAE Arwidsson, 1906.
Nuchal gnioves cvnved. Clephalic ridge present but pt>orly defined. No cephalic
plate. Pygidium either conical with a ventral wiKe below the terminal anus or
shield-shaped with a dorsal anus. Segments withr>ut collars. Neuropodial hooks
from setiger i onwards and ahvays in single rows. The first fvw nemosetae may be
reduced to acicular spines Ijut later hooks have a \ertical scries of denticles abo\e the
main fang.
Genera LUMBRICLYMENE, CLYMENURA, NOTOPROCTUS, LUMBRICLYMENELLA,
PRAXILLURA.
RHODININAE Arwidsson, 1906.
Nuchal grooves slightly curved. ClephaUc keel present. No cephalic plate.
Pygidium conical with a terminal anus and a \cnlral valve. Some anterior segments
with anterior collars and some jirislerior ones with posterior collars. Neuropodial
hooks absent from the first few setigers but thereafter arranged in double rows.
Individual hooks with transverse arcs of denticles above the main fang.
A single genus RHODINE.
MALDANIDAE 617
NICOMACHINAE Arwidsson, 1906.
Nuchal grooves straight or curved. Cephalic keel well developed and arched. No
cephalic plate. Pygidium with the anus on a foliaceous plate or sunken in a funnel
rimmed with cirri. Segments without collars. Neuropodial hooks always in a single
row ; they are present in reduced numbers from setiger i and are usually simplified
to form acicular spines on the first three or four setigers. Later hooks with a vertical
series of teeth above the main fang.
Genera NICOMACHE, MICROMALDANE, PETALOPROCTUS.
EUCLYMENINAE Arwidsson, 1906.
Nuchal gi coves usually straight and parallel. Cephalic plate present, usually with
a well developed rim. Pygidium either with the anus sunken in a funnel ringed with
cirri or with the anus on a cone projecting from a ring of cirri and provided with a
ventral valve. Segments without collars except rarely on setiger 4. Neuropodial
hooks always in a single row. Neuropodial hooks occasionally absent from setiger i
but usually present in reduced numbers on the first three or four setigers. First few
hooks often sompUfied to form acicular spines. Later hooks with a vertical series of
teeth above the main fang.
Genera AXIOTHELLA, CLYMALDANE, CLYMENELLA, EUCLYMENE, GRAVIERELLA,
JOHNSTONIA, MACROCLYMENE, MACROCLYMENELLA, MALDANELLA, MICRO-
CYLMENE, PROCLYMENE, PRAXILLELLA.
MALDANINAE Arwidsson, 1906.
Nuchal grooves curved and divergent anteriorly. Cephalic keel low and broad or
high and arched. Cephalic plate present with a well developed rim. Pygidium a
slanting plate with the dorsal anus above it. No anal cirri. Segments without
collars. Neuropodial hooks always in a single row and absent from setiger i . Later
hooks all similar and have transverse arcs of denticles above the main fang.
Genera MALDANE, ASYCHIS, BRANCHIOASYCHIS.
Key to Subfamilies and Genera
Neurosetae start on setiger 5 and are arranged in double rows. (Setigers 2 and 3 with
collars s.-f. Rhodininae fig. 30.1.3) ...... RHODWE (p. 618)
Neurosetae start on setiger i or 2 and are arranged in single rows .... 2
Cephalic plate present though it may be poorly marked. Cephalic ridge low . . 3
Cephalic plate absent. Cephalic ridge well marked, (s.-f. Nico.machinae) (fig. 30. i.j)
Cephalic plate well defined and surrounded by a raised margin. Pygidium variable
Cephalic plate poorly defined. Pygidium conical, sometimes flattened ventrally. (s.-f.
Lumbricly-meninae) ...........
P>'gidium encircled by anal cirri with the anus on a projecting cone or sunken in a funnel
Neurosetae often present on setiger i. (s.-f. Euclymeninae) ....
Pygidium as a slanting plate with a dorsal anus above it. No anal cirri. No neurosetae
on setiger i . (s.-f. Maldaninae) . . . . . . . . . 17
6i8 POLVCHAEIA OF SOUTHERN AFRICA
5 Neuroseiae of setigers 1-3 acicular ; later ones as hooks with long, almost straight shafts 6
- Neurosetac of all setigers S-shaped with a swelling at the inflection. (Body small, possibly
juveniles of other genera) ...... MICROMALDANE (p. 6iCj)
6 Pygidium funnel-shaped with marginal cirri (fig. 30.1. k) . . NICOMACHE (p. 62:)
- Pygidium petaloid with a central anus ; no anal cirri (tig. 30.2.6)
PETALOPROCTVS (p. 622;
7 Neuropodial hooks of setigers 1-4 generally similar to those of succeeding segments but
have fewer teeth . " CLYMENURA (p. 62J)
- Neurosetae of setigers 1-3 or 1-4 acicular and markedly difi'erent from the hooks of
succeeding segments . ........... 8
8 Pygidium conical with a well marked ventral valve (fig. 30. 2. k). Setigers 1-3 with
acicularneurosetae LUMBRICLYMENE (p. 62 j)
Pvgidium a slanting plate with a dorsal anus. .Setigers 1-4 with acicular neurosetae
NOTOPROCTUS (p. e^y)
9 Neurosetae absent from setiger i . . . . . . MALDANELLA (p. b-zb)
- Neurosetae present on setiger i . . . . . . . . . . 10
10 Neurosetae of the first three to four segments are dentate crested hooks essentially similar
to those of later segments ...........11
— Neurosetae of the first three to four segments arc acicular and differ from those of later
segments ............. 12
11 Segments not very numerous (14-24) ; posterior ones not campanulate
AXIOTHELLA (p. 629)
— Segments very numerous (30 + ) ; posterior ones short and campanulate (fig. 30.4.6)
GRAVIERELLA (p. (,iQ)
12 Neurosetae of the first three setigers are acicular. Nuchal grooves straight ... 13
— Neurosetae of the first four setigers are acicular. Nuchal grooves curved PROCLYMENE*
1 3 A collar on the anterior margin of setiger 4 . . . . . CL YMENELLA*
— Segments without collars ........... 14
14 Rows of vascular cirri on the last few segments (fig. 30. 4. j) . JOHNSTONIA Ip. 630)
— No vascular cirri on the last few segments . . . . . . 15
15 Thirty setigerous segments or more .... MACROCLYMENE ip. (>},'i)
— Twenty-five setigerous segments or less . . . . . . 16
16 .\nus sunk in a pvgidial funnel rimmed witli cirri ; no enlarged ventral va!\e (fig. 30. 7. f)
EUCLYMENE (p. 634)
— Anus on a cone projecting from a circle of cirri ; a large \enlral valve (fig. 30. 7. n)
PRAXILLELLA (p. 642)
17 Cephalic keel well marked. Rim of cephalic plate notched laterally (fig. 30. 8. a)
MALDANE (p. 645)
— Cephalic ridge low and broad. Rim of cephalic plate very deeply incised laterally 'fig.
30.8.g) . . i3
18 Middle segments with numerous gilHilaments . . . BRANCHIONASYCHIS*
— Middle segments without gill filaments ...... ASYCHIS .p. b.\j)
RHODINE MAmgrcn, 1866
Head with a cephalic keel but no ce]3halic plate or raised rim. Nuchal grooxes
cur\ed. Anterior segments long with anterior collars on setigers 2 and 3. Glandular
bands well developed. Posterior segments with posterior collars. Rudimentary
achactous preanals present. Pygidium conical with a terminal anus and a well
developed ventral valve but no anal cirri. Notosetae are winged capillaries and
forms with flattened blades. Neurosetae absent on setigers 1-4 but present on
MALDANIDAE 619
setiger 5 and are arranged in double rows. Hooks with transverse arcs of denticles
above the main fang.
Type species : Rhodine loveni Mahngrcn, 1866.
Rhodine gracilior Tauber, 1 879
(fig. 30.i.a-d)
Rhodine gracilior Tauber, 1879 : 123 ; Arwidsson, 1906 : 74, pi. 2 figs. 53-58 ; pi. 7 figs. 237-241 and
pi. 8 figs. 242-243; Day, 1963a: 433, fig. 10 m-p.
Tube fragile. Body up to 50 mm. long by i.o mm. with a minimum of 23 setigers
and 5-10 achaetous preanals. Colour pale apart from a red or purple fourth setiger
(fig. 30. 1. a). Prostomium (fig. 30.i.b) forming an oval hood over the mouth with a
smoothly arched cephalic ridge and n-shaped nuchal grooves. No eye spots. Head
three times as long as broad with only a dorsal crest to mark the junction with seuger
I which is five times as long as broad. Setigers 2-5 progressively shorter, setigers 5-8
subequal and only twice as long as broad. Setigers 9 and 10 fused and setigers 11-16
about three to five times as long as broad. All subsequent segments decrease in
length and the achaetous preanals are much broader than long. Pygidium (fig.
30. 1. c) a short blunt cone with a terminal anus and a well developed ventral valve.
Setigers 2 and 3 with smooth-edged anterior collars. Setigers 3 and 4 completely
glandular, setigers 5-9 with the anterior half glandular but subsequent setigers with
glands restricted to a small area around the posteriorly situated parapodia. From
the 17th setiger onwards all segments have posterior collars, even the achaetous
preanals. Notosetae include narrow-winged capillaries and others with flattened,
sword-like blades. Neuropodial hooks appear in setiger 5 as double rows of hooks
arranged face to face. Each hook (fig. 30.i.d) with an expanded head bent back at
an angle to the shaft. A close-set cap of 15-20 denticles above the rostrum. Tendons
rudimentary and below them is a boss before the hook tapers to the shaft.
Type locality : North Sea.
Records: Cape (34/18/s and 34/22/d).
Distribution : Sweden (d) ; Nigeria (s) ; Angola (d) .
MICROMALDANE Mesnil, 1897
Small worms with few segments having the head bent downwards and without a
cephalic plate. Cephalic crest poorly defined. Nuchal grooves curved. Pygidium
with a shallow funnel whose margin is crenulate. No ventral valve. Notosetae
include hastatc-bladed capillaries and fine geniculate forms. Neurosetae are similar
in all parapodia, all avicular with a vertical crest of teeth above the main fang and
a swelling at the inflection of the S-shaped shaft. This genus probably represents a
juvenile stage o{ Mcomache.
Type species : Micromaldane ornithochaeta Mesnil, 1897.
620
POLVCHAETA OF SOUTHERN AFRICA
Fig. 30.1. Rhoilmc gracilior. (a) Aiilorior end. (b) Lateral view of head, (v.) Posterior end.
(d) Hook. Mkromalilane sp. (e) Head, (f) Posterior end. (r.) Hook. (11) Hastate
notoseta. J^'icomache Iwnbricalis. (i) Entire worm (natural size), (j) Dorsal view of liead.
(k) Posterior end. (i.) Acicular spine from setiger 2. (m) Normal hook, (n) Feathered
capillary, (o) Winged capillary. .\'icomache mossambka. (v) Anterior end. ((j) Posterior
end. (r) Normal hook, (s) .Xcirular spine from setiger 2.
MALDANIDAE 621
Micromaldane sp.
(fig. 30.i.e-h)
Micromaldane sp. Day, 1955: 531.
Body small with 22-23 setigers and one achaetous preanal. Head rounded in
front with a well defined cephalic ridge (fig. 30.1.6). Notosetae (fig. 30.i.h) hastate
with striated blades. Neuropodial hooks (fig. 30.i.g) S-shaped and swollen.
Pygidium (fig. 30.i.f) fijnnel-shaped with 10-12 triangular marginal cirri.
NICOMACHE Malmgren, 1866
Head with an arched keel but no cephaUc plate. Nuchal grooves S-shaped.
Segments 1-7 short with glandular rings, posterior ones longer. One or two achaetous
preanals in front of the pygidium which is funnel-shaped and rimmed with cirri.
Anus central and sunk in the funnel. Notosetae include winged capillaries and
finely spinulose forms. Neurosetae of the first three setigers are acicular spines and
succeeding ones are numerous hooks with a vertical series of teeth above the main
fang.
Type species : Sabella lumbrkalis Fabricius, 1 780.
Key to Species
I Twenty-two setigers : one achaetous preanal. Pygidium short and funnel-shaped
N. lumbricalis
- Thirty-seven setigers : one achaetous preanal. Pygidium long and cylindrical N.mossambica
Nicomache lumbricalis (Fabricius, 1780)
(fig. 30.i.i-o)
Sabella lumbrkalis Fahncius, 1780: i8g.
Nicomache lumbricalis : Fauvel, 1927: 190, fig. 66 a-i.
Body (fig. 30. 1. i) up to 160 mm. long. Prostomium (fig. 30.1.J) ill-defined, short
and broad merging into an arched cephalic keel. Ocelh present in juveniles. Nuchal
grooves short and S-shaped. Twenty-two to twenty-three setigers plus one ver>' short
achaetous preanal followed by the pygidial ring and funnel which is short and
rimmed with 15-25 equal anal cirri (fig. 30.i.k). Anus sunken, no ventral valve.
A single smooth acicular spine (fig. 30.1.I) in each of the first three neuropodia.
Subsequent neurosetae as numerous hooks each with a vertical series of four to five
teeth above the main fang and tendons below (fig. 30.i.m). Notosetae include
normal winged capillaries (fig. 30.1.0) and fine feathered forms (fig. 30.1.0).
Type locality : Greenland.
Records: Cape (from 30 i 7 i and 33 18 i, s to 34/21/i).
Distribution : Arctic ; North .\tlantic from Greenland (s, d, vd) to the English
Channel (e) ; North Pacific from the Behring Sea to N.\V. Japan and W. Canada to
Southern California (d, vd) ; Bouvet Is. (d).
622 POLVCHAETA OF SOUTHERN AFRICA
Nicomache mossambica Day, 1031
(fig. 30. 1 .p-s)
Nicomache mossambica Day, 1951 : 52, fig. 7 a-f.
Body u]) to j-i-O mm. long by 5 mm. Head (fig. 30.i.p) bent forward with a short,
arched cephalic keel dorsally. Nuchal grooves almost parallel. Thirty-seven setigers,
one minute achactous preanal and a long cylindrical pygidium rimmed with numer-
ous short cirri (fig. 30.i.q). First three segments short and glandular ventrally,
succeeding ones longer. Notosetac are narrow-winged capillaries plus a few fine
spinulose forms. Neurosetac of setigers 1-3 are two to four straight acicular spines
(fig. 30. 1. s) ; succeeding neurosetac are rows of hooks with a vertical scries of five
teeth above the main fang (fig. 30.i.r).
Type locality : Del.igoa Bay.
Records: Mocambiciue (26/32/i).
DisiRiEuiiON : A single record.
PETALOPROCTUS Quatrefagcs, 1865
Head with an arched keel but no cejihalic plate or raised margin. Nuchal grooves
short, divergent. Anterior segments short with anterior glandular rings, middle seg-
ments longer and the posterior ones again short. Small achactous preanal segments
may precede the pygidium which is a fohaceous plate with a central anus. Setigers
1-3 with acicular neurosetac, the succeeding ones with neuropodial hooks luu'ing a
vertical scries of teeth above the main fang. Notosetac include winged capillaries
and finely spinulose forms.
Type SPEC:IES : Pelalnproclin /ivv/co/fl Qiialrcfages, 1865.
Petaloproctiis terricola Quatrefagcs, 1865
(fig. 30.2.a-d)
Pi'latoprocli/i terricola Quatrefages, 1856 II : *J47 ; Fauvcl, 1927 : 194, fig. 68 a-I.
Body u]) to '200 mm. long. Head (fig. 30. 2. a) with a broad, semicircular pro-
stomium overhanging the mouth followed by a marked keel with short divergent
nuchal groo\'es on eithir side. Eye-spots present. Body of 22 setigers plus the
pygidium (fig. 30. 2. b). No acluu-rous preanal but two annuli below the pygidium.
Anterior segments short with glandular rings. In large specimens the last fi\e to
seven segments develop fleshy dorsal lobes which pr(ij<-ct backwards. Notosetac
include winged capillaries, fine capillaries with denticidate tips and spinulose forms
with spirid blades. Neurosetac of setigers 1-3 are smodlh, bluntly pointetl acicula
MALDANIDAE 623
(fig. 30. 2. d). Posterior ncurosctae have a vertical crest of four to five teeth above the
main fang (fig. 30. 2. c).
Type locality : France.
Records: South West Africa (23/14/s, 26/14/d and 26/15/i, s) ; Cape (31/17/i,
33/18/i and 34/18/i).
Distribution : Atlantic from the EngUsh C!hannel (i, s) to Morocco (i, d),
Ghana (s) and Angola (s).
CLYMENURA VcrriU, 1900
Prostomium well developed and followed by long parallel nuchal grooves.
Cephalic plate rudimentary or absent. Anterior segments with glandular rings,
the eighth with a ventral glandular patch. Achaetous preanals present. Anus on a
cone with an enlarged ventral valve, anal cirri few and elongate or absent. Notosetae
include both winged capillaries and feathered forms. The first few neurosetae
essentially similar to the hooks of succeeding segments though the denticles on the
rostrum are poorly developed. Later hooks with a vertical series of teeth above the
main fang and tendons below.
Type species : Clymcne cirrata Ehlers, 1887.
Clymenura tenuis (Day, 1957)
(fig. 30.2.e-i)
Leiochom tenuis Day, 1957 : 110, fig. 7 g-1.
Body up to 120 mm. long by 1.5 mm. broad. Prostomium (fig. 30. 2. c) bluntly
triangular in front and continuous with a faint cephalic ridge behind. OcelH
numerous and well marked. Nuchal grooves long and straight. Twenty-three to
twenty-five setigcrs plus o-l preanals followed by three to five annuli without
parapodial ridges. Pygidium (fig. 30. 2. f) with three slender anal cirri and a well
marked ventral valve which may protrude as a cone (fig. 30. 2. g). Glandular rings
on the peristome and setigcrs 2-7. A glandular patch on the ventrum of setiger 8.
Notosetae include broad-winged capillaries and feathered forms. Neurosetae either
absent from setiger 1 or represented by two to three hooks (fig. 30. 2. i) with two to
four denticles above the rostrum and poorly marked tendons. Posterior neurosetae
numerous with a vertical series of about six teeth above the rostrum and strong
tendons below (fig. 30. 2. h).
Type locality : Durban Bay.
Records: Natal (29/31/i); Mocambique (26/32/i).
Distribution : Only two records.
624
POLYCHAETA OF SOUTHERN AFRICA
Fig. 30. j. Pclaluproclui terrkola. (a) Head, (b) Posterior end. (c) Normal hook, (d)
Acicular spine of setiger 2. Clymemira tenuis, (r.) Anterior end. (f) and (o) Conlrat ted
and expanded appearance of posterior end. (h) Normal hook, (i) Acicular spine of
setiger i. LumbrUlynmw cylindrkaudata. (j) Anterior end in dorsal view, (k) Posterior end.
(l) Acicular spine, (m) Normal hook. Lumbrkhmcne minor, (n) Normal hook, (o) Dorsal
view of anterior end. (p) Posterior end.
MALDANIDAE 625
LVMBRICLYMENE Sars, 1872
Head without a cephalic plate or raised margin but with a broad prostomium
followed by a median dorsal ridge. Nuchal grooves markedly curved. Setigerous
segments elongated, the anterior ones without collars but with glandular rings.
Acicular neurosetae present in the first few (usually 4) setigers but each of the later
setigcrs has a row of hooks with a vertical series of denticles above the main fang.
Posterior end of the body tapered with a few achaetous preanal segments preceding
the conical pygidium. No anal funnel. Anus with a marked ventral valve.
Type species : Lumbricljmene cylindricauda Sars, 1872.
Key to Species
1 Pygidium cylindrical and anus terminal. ..... L, cylindricauda
- Pygidium flattened ventrally and anus dorsal ....... L. minor
Lumbriclymene cylindricauda Sars, 1872
(fig- 30-2-j-m)
Lubriclymene cylindricauda Sars, 1872 : 413 ; Arwidsson, 1906 : 40. pi. i figs. 15-25, pi. 7 figs. 219-221 ;
Day, 1963a: 432.
Body long and slender, up to 120 mm. long with 19 setigers and four to six
achaetous preanals of which only the first three are well marked. A brown bar
across the buccal segment and at each end of the first three setigers. Head (fig.
30.2.J) steeply arched with a median ridge above the broad prostomium. No eye-
spots. Nuchal grooves markedly curved, almost semicircular. Setiger i four times
as long as broad and succeeding ones even longer, an average segment in the middle
of the body being about seven times as long as broad. The last few setigers are
shorter and the achaetous preanals decrease progressively in length so that the last
is broader than long (fig. 30. 2. k). Pygidium bluntly conical with a marked ventral
valve below the anus. No anal cirri. The first six setigers have anterior glandular
rings which become progressively broader. In posterior segments glandular swellings
are hmited to the area around the setae but from setiger 11-14 there is a mid-ventral
glandular streak, .\otosetae are smooth-winged capillaries throughout. Neurosetae
of setigers 1-4 are stout, blunt, acicular spines (fig. 30. 2. 1), one per parapodium.
Neurosetae of all subsequent setigers are dentate-crested hooks. Setiger 5 bears a
row of about seven but later setigers have more. Hooks on setiger 5 with only two
to three teeth above the main fang, but each hook of later segments has a vertical
series of about four to six teeth above the main fang (fig.30.2.m).
Type locality : Norway.
Records: Cape (34/23/d).
Distribution : Norway.
f.L-6 I'OI.VCIHAETA OF SOUTHERN AI'RICA
Lumbriclymene minor Ardwidsson, 1906
(fig. 30.2. n-p)
Lnriibrithmc-nt' minor Arvvitlsson. 1906: 4(j. pi. i figs. 26-29, P'- 7 *V?s. 22^^-226; Fauvcl, 1927: 196,
fig. 68 k-q; Day, 1963: 366.
Body 25-30 mm. lung and about i.o mm. broad. Head (fig. 30.2.0) rounded
anteriorly without a well defined prostomium and dorsal crest. Nuchal grooves
poorly marked, curved, almost semicircular. No ocelli. Body of 19 setigerous
segments, three poorly marked achaetous preanals and a blunt, ventrally flattened
pygidium with a dorsal anus (fig. 30. 2. p). No anal cirri. 'I'hc setigerous segments
do not dilier greatly in length, each being three to four times as long as broad.
Intersegmental constrictions poorly marked and glandular bands though present on
the first si.x segments are not well defined. Notosetae are all winged capillaries
mostly with smooth margins but some arc striated distally. Neurosetae of setiger 1-4,
arc represented by a single, stout, smoothly pointed acicular seta per segment.
Subsecjuent neurosetae are rows of five to seven hooks, each with four to five teeth
in a vertical series aljove the main fmg and a few lateral denticles (fig. 30. 2. n).
The neck is short and there is a well defined swelling preceding the smoothly tapered
shaft.
Type locality : Sweden.
Records: Ca]5c (34/17/abyssal).
Distribution: Greenland (s, d) ; Sweden (d).
MALDANELLA Mcintosh, 1885
Prostomium well developed, \vith or without eye-spots. A well marked cephalic
plate with a raised margin. Nuchal grooves straight and parallel. Pygidium funnel-
shaped with marginal cirri. Anus sunk in the funnel. Achaetous preanal segments
present. Setiger i without neurosetae and setigers 2 and 3 with a reduced number
of simjjlified hooks ; later segments with a single row of hooks bearing a vertical
series of teeth above the main fang. Notosetae are winged capillaries and often
include feathered forms.
Type species: Maldaitella aiitarctka Mcintosh, \V,Q'\.
Key to .Spp:cies
I Two to three ncnroj)odial hu<jks on setiger 2. I'oiir short acliactous preanal segments
M, capcnsis
- 'I'weKe or more neiiropociial hooks on setiger 2. Tlirce achaetous preanal segments
M.Jibrillata
MALDAMDAE 627
Maldanella fibrillata Chambcrlin, 19 1 9
(fig- 30.3-f-k)
Maldanella fibrillata Chamberlin, 1919 : 413, pi. 72 figs. 1-6, pi. 73 figs. 1-2 ; Day, 1963 : 365.
Body up to 70 mm. long. Prostomium (fig. 30. 3. f) small and without eye -spots.
Cephalic plate (fig. 30.3.J) oval and slanting with a high, unnotched rim. Nuchal
grooves straight, half the length of the cephalic plate, their anterior ends continuous
with a sharply bent streak which curves back around the inside of the rim. Body
with ig setigers and three achaetous preanals the last of which is united to the pygidial
funnel (fig. 30. 3. k). Anterior segments short with glandular rims, middle ones long,
posterior ones decreasing, the last few sausage-shaped. Funnel with about 50 sub-
equal cirri ; anus sunk in the funnel and lacks a ventral valve. Notosetae of two
lengths but all are narrow-winged capillaries. IVo neurosetac on sctiger i. Setiger 2
with 12-20 hooks each with two to three teeth above the main fang (fig. 30. 3. g).
Setiger 3 and later ones with a row of about 30 hooks having a vertical series of three
to four teeth above the main fang (fig. 30. 3. h).
Type locality: 1,471 fathoms off Pacific coast of Panama.
Records: Cape (33/16/abyssa]).
Distribution: Pacific off Panama (abyssal).
Maldanella capensis Day, 1961
(fig. 30.3.a-e)
Maldanella capensis Day, 1961 : 523, fig. 1 1 h-k.
Body up to 45 mm. long by 2 mm. Prostomium (fig. 30. 3. a) broad with a curved
anterior margin and numerous eye-spots. Cephalic plate (fig. 30. 3. b) broadly oval
with a very low rim having deep lateral notches. Nuchal grooves short and straight.
Body with 19 short setigers, four very short achaetous preanals followed by the
pygidial ring and funnel (fig. 30. 3. c). Anterior segments hardly longer than broad,
later ones twice as long as broad. The four preanals together are shorter than the
last two setigers. Thirty short anal cirri. Anus sunk in the funnel and without a
marked ventral valve. Notosetae include narrow-winged capillaries and fairly
broad feathered forms. Setiger i without neurosetac. Setiger 2 with only two hooks
each having only three to four teeth (fig. 30. 3. d) ; setiger 3 with three to five hooks
and later segments with numerous hooks. Normal hooks (fig. 30. 3. e) with an acutely
curved main fang surmounted by a vertical series of about five teeth.
Type locality : Agulhas Bank, South Africa.
Records: Cape (32/28/s).
Distribution : A single record.
628
POI.VCHAETA or SOUTHERN AFRICA
I"ii,. '',o-3- MdhUimlla i:(!f'ai\i'i. (a) Anterior end. (u) Dorsal \ icw of head, (c) Poslerior
end. (d) Hook from seligcr \i. (e) Normal iiook. MahhincUa jdnillala. (f) Anterior end.
f<;j H<jok from setiger ■!. [u] Normal hook, (i) Notopodial capillary. (|) Head, (k)
Poslerior end. Axio/heUa qutidriniaculiihi. (i.) .Anterior end. (m) Posterior end. (n) Hook
in profile, fo) Hook in face view. Axiolhella jarli. (pj Anterior end. (Q.) I'osterior end.
(r) Hook (after Kirkegaard 1959).
MALDANIDAE 629
AXIOTHELLA Verrill, 1900
Head with a flattened cephalic plate with a raised rim. Nuchal grooves straight
and parallel. Anus sunk in a funnel rimmed with cirri or mere crenulations. No
ventral valve. Achaetous preanals sometimes present. Notosetae of two types.
Neurosetae essentially similar throughout from the first setiger onwards, sometimes
avicular, always with a vertical series of teeth above the main fang.
Type species: Axiothea calenata Malmgren, 1865.
Key to Species
I A small species with avicular hooks having a swelling in the S-shaped shaft. No achaetous
preanals ........... /4, quadrimaculata
- A large species with fairly straight hooks. Two achaetous preanals . . . A.jarli
Axiothella quadrimaculata Augener, 1914
(fig. 30.3.1-0)
Axiothella quadrimaculata Augener, 19 14: 70, pi. i fig. 10, text-fig. 10; Day, 1957 : iii, fig. 8 a-d.
A minute cylindrical worm about 5 mm. long with 19-20 setigers, no preanals and
a pygidial ring and scalloped funnel (fig. 30. 3. m). Prostomium (fig. 30. 3. 1) broadly
curved in front and continuous with a cephalic ridge behind. Nuchal grooves slightly
curved. Cephalic rim low and poorly marked. Anterior segments barred and brown
marks on head. Notosetae include hastate-bladed capillaries and a few feathered
forms. Neuropodial hooks avicular with five teeth in a vertical series above the main
fang and a swelling in the S-shaped shaft (fig. 30. 3. n).
Type locality : South-west Australia.
Records: ? Cape (33/17/s and 34/18/s) ; Mocambique Island (i).
Distribution : South-west Australia, New Zealand and subantarctic islands (i, s).
Axiothella jarli Kirkegaard, 1959
(fig. 30.3.p-r)
.^io/Af//aJar/i Kirkegaard, 1959: 56, figs. 11-13.
Body up to 35 mm. long. Prostomium (fig. 30.3. p) rounded in front with numerous
ocelh. Nuchal grooves straight and half the length of the cephalic plate. Cephalic
rim at first high, then notched and behind the notch is a low semicircular part
notched in the mid-dorsal line. The first eight setigers are short and the seventh
has a marked glandular 'ing. Number of setigers unknown. Two long achaetous
preanals precede the pygidial ring and funnel which is rimmed with 18 alternately
long and short cirri (fig. 30. 3. q). Neuropodial hooks of the first three setigers (fig.
30.3. r) with three to four teeth above the main fang and subsequent ones with a
vertical series of six to seven teeth. First three setigers with three, six and seven
G30 rOl.VCHAETA OF SOLTHKRN AFRICA
hooks ix-sjH-ctivcly, and subsequent ones with numerous hooks in a single row.
Notosctae include broad-winged capillaries and spinulose forms.
Type loi:ality : In 50 ni. off Angola.
Records: C^ape (33/i8;'s).
Distribution: Angola (s, d).
GRAVIERELLA Fauvel, 1919
C'c'i.ilialic pl.Ue with a raised rim. Nuchal groo\es straight and parallel. Body
with \ery numerous segments, the posterior ones being campanulate. No achactous
preanals, the region of regeneration being in IVont of the last few setigers. Pygidium
funnel-shaped with marginal cirri. Anus sunken and lacks a ventral valve. Neuro-
setae of the first three setigers reduced in number but essentially similar to all
succeeding ones. Hooks with a vertical series of teeth above the main fang.
Type species: Gravierella inultianmdala Fau\el, 1919.
Gravierella miiltiattttulata I"au\(I, 1919
(lig. 3o.4.a-e)
Grai-irnila muUhinmilala Fauvel, 1919: 4:^8, pi. 17 tigs. 58-69.
Body up to 80 mm. long by 1-2 nmi. Tube s.uuly, \ery fragile. Prostomium
(fig. 30. 4. a) well developed and conical with numerous ocelli. C'ephalic rim high
with lateral and posteiior identations. Nuchal grooves very long and parallel. Body
slender and elongate with 60-70 segments. No achactous preanals but with a region
of regeneration towards the posterior end. Pygidium (fig. 30. 4. b) with about 20
anal cirri which increase in length ventrally. First seven segments slightly longer than
broad ; segments 8-r:! longer. From setiger 14 onwards the segments are very short
and campanulate with backwardly directed collars. Neurosetae of setigers 1-3 are
three to five dentate hooks \ery similar to the more numerous ones of succeeding
segments which ha\e a \ertical series of five to sex'en teeth above the main fang
(fig. 30.4.1 1. Notoset.ie .ue winged capillaries (fig. 30. 4. e) and line feathered forms
(fig. 30.4.d).
Type locami v : M.uiagascar (Tulear).
Rec;()rds : Mocambicjue (26/32/i).
DisTRiuuTio.N : Madagascar.
JOHNSTONIA Qu.itrcfiges, 1865
111, id with a cephalic plate bordered by a raised rim. Xuchal grooves straight and
]i,u.dlcl. Anterior segments short, posterior ones long, the last few bearing rows of
vascular cirri. Sometimes an achactous preanal segment. Pygidium funnel-shaped
\s'ith marginal cirri and a central anus sunk in the funnel. No \entr.Ll \ ,iK e. Setigers
MALDAMDAE 631
1-3 with acicular neurosctac, subsequent ones with hooks with a vertical series of
numerous denticles above the main fang. Notosetae of two kinds including winged
capillaries and feathered forms. Anterior segments with glandular rings and posterior
ones with longitudinal belts.
Type SPECIES : Johnslnnia dvmenoides Quatrti^ages, 1865.
Key to Species
I Acicular neurosetae of setigers 1-3 with dentate apices. Vascular cirri simple (fit;. 30. 4. f)
J, clymenoides
— Acicular neurosetae of setigers 1-3 with smooth tips. Vascular cirri branched (fig. 30. 4. j)
J. knysna
Johnstonia clymenoides Qiiatrcfages, 1865
(fig. 30.4.f-h)
Johmlonia clymenoides Qualrefagcs, 1865 II : 245, pi. 1 1 figs. 10-15 ; Fauvel, 1927 : 184, fig, 64 a-h.
Body elongate, up to 200 mm. long, with 22 setigers and one preanal in front
of the pygidial funnel. Clephalic plate bordered with a lightly scalloped rim.
Prostomium broad ; ocelli present. Nuchal grooves straight and two-thirds the
length of the cephalic plate. Glandular rings on setigers i to 7 and four longitudinal
belts on setigers 9-15 situated on the dorsum, ventrum and the two sides. The last
five to six setigers with four rows of vascular cirri which are usually unbranched
(fig. 30.4.f). Achaetous preanal short, poorly marked. Pygidial funnel rimmed
with 20-30 anal cirri of which the ventral one is the largest. No ventral valve.
Notosetae include broadly winged and feathered capillaries. Neurosetae of setigers
1-3 have one to two vestigial denticles above the apex (fig. 30. 4. g) ; subsequent
hooks with numerous denticles in a vertical series (fig. 30. 4. h).
Type locality : France.
Records : South West Africa (23/14/d).
Distribution : Atlantic from the Enghsh Channel (i) and Morocco (i) to tropical
West Africa (s) ; Mediterranean (i).
Johnstonia knysna Day, 1955
(fig. 30.4.i-m)
Johnlonia knyma Day, 1955 : 429, fig. 4 e-j.
Body up to 180 mm. long by 4 mm., brownish in alcohol. Prostomium (fig. 30.4.1)
bluntly conical, with eye-spots. Nuchal grooves straight and extend over two-thirds
of the cephahc plate. Cephalic rim crenulate posteriorly. Body with 19 setigers, and
one to two achaetous preanals followed by the pygidial funnel (fig. 30.4.J). Setigers
1-3 fairly short, setigers 4-7 successively longer, setiger 8 short, setigers 9-19 about
twice as long as setiger 8, the 19th being four times as long as broad. The first
preanal is well developed and not much shorter than setiger 19 ; the second preanal
632
POLVCHAETA OF SOUTHERN AFRICA
Fig. 30-4- Gravicrella imdliannulata. (a) Anterior end. (b) Posterior end. fc) Normal hook.
(d) Tip of feathered capilhiry. (t) Winged capillary. Johutonia chmenoides (after Fauvcl).
(f) Segment with vascular cirri, (o) Acicular neuroseta of setigcr 2. (h) Normal hook.
Johnsfonia kriYsna. (i) Head, (j) Posterior end. (k) Branching vascular cirrus, (l) Acicular
neuroseta of setiger 2. (m) Normal hook. Mairochmene monilis (after Fauvel). (n) Anterior
end. (o) Posterior end. Macruclyninie saldauha. (p) Anterior end. (q,) Posterior end.
(r) Acicular neuroseta of setiger -z. (s) Normal hook.
MALDANIDAE 633
is either poorly marked or absent. Pygidium elongate, ending in a funnel encircled
by 20 cirri and a sunken anus with a small ventral valve. Glandular bands ex-
tensive on setigcrs 3-8. A narrow vential glandular streak from setiger 9 onwards
and a dorsal glandular streak from setiger 9 to 14. Vascular caeca (fig. 30. 4. k)
branched and arranged in four rows from setiger 14 to the first preanal. Notosetae
include broad-winged capillaries and fine spinulose forms. Neurosetae of setigers
1-3 as a single smooth, bent acicular spine per neuropodium (fig. 30.4.I). Later
setigers with numerous hooks, each with a vertical series of five to six teeth above
the main fang and well marked tendons below (fig. 30. 4. m).
Type locality : Knysna Estuary, South Africa.
Records: South West Africa (26/14/d) ; Cape (32/18/s, 34/23/e and 33/25/s).
Distribution : Endemic.
MACROCLYMENE \ cmW, 1900
Head with a slanting cephalic plate with a raised rim. Nuchal grooves straight
and parallel. Body with more than 30 setigcrs and sometimes an indistinct achaetous
preanal. First three setigcrs with one to three acicular neurosetae which may have
smooth or faintly dentate tips. Subsequent hooks each with a vertical series of five
to seven teeth above the main fang and chitinous tendons below. Anterior segments
with glandular bands. Posterior segments short, often campanulate. Pygidium
with the anus sunk in a poorly developed funnel whose margin is rimmed with cirri.
Type species : Clymene producta Lewis, 1 899.
Key to Species
I Acicular neurosetae of setigers 1-3 with smooth tips ..... Af. trtonilis
- Acicular neurosetae of setigers 1-3 with vestigial teeth (fig. 30. 4. r) . . M. saldanha
Macroclymene monilis (Fauvel, 1902)
(fig. 30.4.n-o)
Clymene monilis Fauvel, igoa : 8g, figs. 31-38.
Body up to 120 mm. long by 2 mm. Prostomium (fig. 30.4.n) ovoid. Nuchal
grooves two-thirds of ccphaHc plate. Cephalic rim with a marked lateral notch and
five to eight triangular lobes posteriorly. Thirty-one or more setigers followed by the
pygidial ring and funnel (fig. 30.4.0) which is encircled by 16-20 unequal anal cirri.
No achaetous preanal segments. Setigers 1-3 each with a single neuropodial spine.
Setigers 4-7 with anterior glandular rings. Middle segments twice as long as broad,
posterior ones campanulate. Notosetae include many broad-winged capillaries, a
few fine-winged capillaries and slender feathered forms in posterior segments.
N
Cj4 I'oi.vcii \r. r.\ or southern ArRK:.\
Aciciilar iicurosctac of sctigcrs 1-3 stout and abruptly bent near tlicir snionlli
pointed tips. Posterior hooks with a vertieal series of six Id seven teeth abcnc the
main fang.
Type locality : Senegal, western ,\friea.
REt:oRDS : Not recorded from southern Afrie.i.
Distribution: Senegal (i), Nigeria (i).
Macroclymene saldanha (Day, 1955)
(fig. 30.4.P-S)
Clymenc [PraxiUelta) siilclmiha Day, 1955: 428, fig. 4 a-d.
Body about 55 mm. long by 1.3 mm. and creamy yellow in spirit. Pi'ostomium
(fig. 30. 4. p) triangular, pointed and without eye-spots. Nuchal grooves straight
and extend over four-fifths of the cephahc plate. CcphaUc rim well developed and
high anteriorly, notched laterally and again posteriorly. Body with 35-40 setigers,
a doubtful achaetous preanal or possibly none and a small pygidium (fig. 30.4. q).
Setigers 1-3 short, sctigcrs 4-8 longer, and subscc|uent ones progressively shorter
again until the last is only twice as long as broad. Pygidium shorter than the last
setiger, and lacks a funnel-shaped expansion, having merely a ring of 10-20 short
cirri around the sunken anus. Glandular bands faint on setigers 2-3, well marked on
setigers 4-7 and faint on setiger 8. A narrow ventral streak from the first setiger to
the pygidium. Notosetae include (a) stout broad-winged capillaries ; (b) slender
spinulose capillaries ; and (c) slender smooth capillaries. Neuropodial hooks of
setigers 1-3 are straight with three obscolesccnt teeth above the blunt rostrum
(fig. 30. 4. r). Later hooks are well developed with a vertical series of 5-7 teeth above
the main fang and well marked tendons below (fig. 30.4.S).
Type locality : Langebaan Lagoon, South ."MViea.
Records: Gape (33/18/i).
DisTRiBi'TioN : A single record.
EUCLYMENE \'errill, 1900
Head with a slanting cephalic jilate with a raised rim. Nuchal grooves straight
and parallel. Up to 25 setigerous segments and foiu' achaetous preanals. The first
three setigers with one to three aeicular neurosetae which usually have smooth tips
or occasionally vestigial denticles ; subsequent neurosetae are numerous hooks with
a vertical series of five to seven denticles above the main fang and chitinous tendons
below. Pygidial funnel rimmed witli ( iiri. Anus sunk in thi' fumiel .uid has no
enlarged ventral valve. Glandular rings on anterior segmi'uts and sonu'timcs
glandular streaks on posterior ones.
Type species : Clymenc ocnlcdii Glaparcde, 1863.
MALDANIDAE 635
Key to Species
Acicular neurosetae of setigers 1-3 with vestigial teeth on the convex side of the apex.
(Cephahc rim not crenulate posteriorly) ........ 2
Acicular neurosetae of setigers 1-3 with smooth apices ...... 3
Two achaetous preanals and a circular ridge (fig. 30.5.6). Ocelli present
E, oerstedi (p. 635)
Only two achaetous preanals. Ocelli absent ...... E. sp. (p. 636)
Four achaetous preanals ........... 4
Three achaetous preanals. (Nineteen setigers. Cephalic rim crenulate posteriorly (fig.
30. 5. e)) .......... E. lombricoides (p. 636)
Two achaetous preanals ........... 5
Cephalic rim high and crenulate posteriorly. Nineteen setigers . E. lyrocephala (p. 638)
Cephalic rim low and a single notch posteriorly. Number of setigers unknown
E. cf. quadrilobata (p. 638)
Nineteen setigers ............. 6
Twenty-one or more setigers ........... 8
Cephalic rim crenulate posteriorly .......... 7
Cephalic rim with a pair of lateral notches and one posterior one . E. natalensis (p. 638)
A dorsal glandular streak from setiger 9 to 14 (fig. 30. 6. i). Ocelli present
E. glandtdaris (p. 639)
No dorsal glandular streak. No ocelli .....£. mossambica (p. 639)
Twenty-one setigers. Cephalic rim crenulate posteriorly. A dorsal glandular streak from
setiger 9 .......... E. annandalei (p. 641)
Twenty-four setigers. Cephalic rim with a pair of lateral notches and one posterior one.
No dorsal glandular streak ....... E. luderitziana (p. 641)
Euclymene oerstedi (Claparedc, 1863)
(fig. 30.5.0-q)
Clymene oersledii ClaparMe, 1863 : 28, pi. 13 figs. 8-13.
Clymene [Euclymene) oersledii: Fauvel, 1927: 173, fig. 60 a-i.
Body up to 100 mm. long. Prostomium (fig. 30.5.0) small. Ocelli present.
Nuchal grooves two-thirds of cephalic plate. Cephalic rim low with a single notch on
each side. Body slender with 19 setigers, two obvious achaetous preanals and circular
ridge preceding the pygidial ring and funnel (fig. 30.5.9). First preanal two-thirds
the length of the last setiger, the second much shorter, the last small and poorly
marked. Anus with numerous short anal cirri and 7-10 alternating long ones. Anus
sunken with a small ventral valve. Setigers 1-7 short with broad, white glandular
bands contrasting with the red posterior part of the segment. A broad mid-ventral
glandular streak on setigers 7 to 14. The first three setigers widi one to two acicular
neurosetae each having two to three vestigial teeth on the blunt apex (fig. 30. 5. p) ;
later neurosetae are numerous hooks with a vertical series of five teeth above the
main fang and well developed tendons below.
Type locality : Naples.
Records: ? Cape (32/17/d and 33/18/d).
Distribution : North Sea (i) and English Clhanncl (i) and ? tropical western
Africa; Mediterranean (i).
636 POI.VCHAETA OF SOUTHERN AIRUIA
Euclyniene cf. oerstedi
(fig. 3o.5.a-d)
Euclyniene cf. oersledii Day, 1963a: 429, lig. 10 g.
Number of sctigerous segments unknown. Prostoniiuni (fig. 30.5.3) bluntly tri-
angular and lacks ocelli. Nuchal grooves long and straight, equal to four-fifths the
length of the cephalic plate. Cephahc margin high, smooth laterally but with a
single median posterior notch. Segments increase in length after the fifth. Glandular
bands strongly marked on setigers 3-6. A faint mid-ventral streak from setiger i
onwards. Posterior end (fig. 30. 5. b) with two achaetous preanals preceding the
pygidial ring and funnel. First preanal twice as long as the second, the two together
shorter than the last setiger. Anal funnel with 18-24 '^'■"''' alternating in length and
a much longer ventral one. Anus sunken, no ventral \alve. Notosetae include
winged capillaries and fine, feathered forms. Setigers 1-3 with two to three aeicular
spines (fig. 30. 5. c) with abruptly bent tips bearing three to four small denticles. Later
hooks (fig. 30. 5. d) with six to seven teeth above the main fang and strong tendons
below.
Records: Cape (34/23/s).
Euclyniene lonibricoides (Qiiatrcfages, 1865)
(fig- SO-S-e-k)
Clymene lonibricoides Quatrefages, 1863 II : 236.
Clymene {Euclyniene) lombricoides : Fauvel, 1927: 172, fig. 59 a-i.
Body (fig. 30. 5. e) up to 150 mm. long by 9 mm. broad. Prostomium (fig. 30. 5. f)
broadly rounded in front with indistinct ocelli. Nuchal grooves straight and half the
length of the cephahc plate. Cephalic rim with the posterior part divided into about
10 lobes. Body with iq setigers, three preanals decreasing in length followed by the
pygidial ring and funnel. Anterior segments a little longer than broad, posterior
ones two to four times as long as broad. The three preanals together slightly longer
than the length of the last setiger (fig. 30. 5. g). Funnel with vtry numerous cirri
alternately long and short. Anal funnel sunken without a ventral valve. Setigers 2-8
with anterior glandular rings. Setigers 8 to 12 sometimes with a dorsal glandular
streak. Notosetae include winged capillaries (fig. 30.5.J) and feathered forms
(fig. 30. 5. h). Setigers 1-3 each with one to two smooth aeicular neurosetae (fig.
30.5.1). Later neurosetae are numerous hooks (fig. 30. 5. h) each with a \ertical scries
of five to six teeth above the main fuig and well marked tendons below.
Type EOc.\Errv : France.
REroRus : Sf)uth West .-\frica (26, 15/i and 28 iG's); C^apc (from 33/18/1, s to
34/21 i) ; Moscambicjue (26/32/i).
DisTRiDCTio.v : North Atlantic from Scotland (s) and the English Channel (i) to
Morocco (i) ; Mediterranean (i).
MALDANIDAE
637
Fig. 30.5. Euclymene cf. oersledii. (a) Anterior end. (b) Posterior end. (c) Acicular neuro-
setaofsetiger I. (d) Normal hook. Euclymene oersledii (alter Fauve\). (o) Head, (p) Acicu-
lar neuroseta. (q) Ventral view of posterior end. Euclymene lombricoides. (e) Entire worm
(twice life size), (f) Anterior end. (g) Posterior end. (h) Normal hook, (i) .Acicular
neuroseta of setiger 2. (j) Winged capillary, (k) Part of feathered capillar)'. Euclymene ci.
quadrilobata. (l) Head, (m) Posterior end. (n) Acicular spine of setiger 2.
6;sB POLYCIIAKTA OV SOUTHERN AKRICIA
Euclymene lyrocephala (Schinaicla, 18G1)
C/vmi7if />TOfc/i/ia/u Schmarda, iti6i : 15; Augcner, 1918: 483.
Body 80 mm. long. Prostomium ?. Nuchal grooves?. Cephalic rim high, cicnu-
lale posteriorly. Nineteen setigers, four short preanals plus the pygidium. Funnel
with 28 subecjual anal cirri. Aciciilar ncurosetac of lirst three setigers probably
smooth and acicular.
Type locality : False Bay, South Africa.
Records: South West Africa (26/15/i) ; Cape (33/18/i).
Distribution : No other records of this doubtful species.
Euclymene cf. quadrilobata (Sars, 1856)
(fig- 30-5-1-")
Euclymme cf. quadrilobata Day, 1963a : 430, fig. 10 h-j.
No complete specimen known but generally similar to Pseiidoclytnene (/iiadrilohala
(Sars) as described by Arwidsson, 1906. Prostomium (fig. 30. 5. 1) small and rounded.
Ccphahc plate almost circular with a very low rim with a pair of lateral notches and
one posterior median one. Nuchal grooves short and straight, less than half the
length of the cephalic plate. Number of setigers unknown. Glandular bands well
marked on setigers 4-7. Four achaetous preanals each half as long as the last setiger.
Pygidial ring and funnel (fig. 30.5.111) well developed with the anus sunken in the
funnel whicli is rimmed wilii 24 unequal cirri. No anal valve. Ncurosetac of
setigers 1-3 represented by a single stout spine with vestigial denticles (fig. 30.5.11).
Later ncurosetac numerous, each with four to five well marked teeth above the main
fang.
Records: Cape (34/26/d).
Distribution : ?
Euclymene natalensis (Day, 1957)
(fig. 3o.6.j-ni)
Clymene natalensis Day, 1957 : 107, fig. 7 c-d.
Body up to 120 mm. long by 3.5 nini. Prostomium (fig. 30. 6.1) rounded and with
a few ocelli. Nuchal grooves long. Cephalic rim high laterally and separated by a
notch or step from the lower posterior portion which is smooth apart from a median
posterior notch. Body with 19 setigers, two preanals and the pygidial ring and funnel
which has 20 subcqual anal cirri plus a longer ventral one (fig. 30.6.111). First
preanal almost as long as the last setiger, second only half its length. Anus without
a \entral valve. Glandular rings on setigers 2-7 and a median ventral streak from
setiger 9 onwards. No dorsal glandular streak. The first three setigers each with a
single smooth, acicular spine with a isenl tip (fig. 30.6.J). L.iter nein'osetae are well
MALDANIDA 639
developed with six teeth above the main fang and stout tendons below (fig. 30. 6. k).
Notosetae include numerous winged capillaries and a few fine feathered forms.
Type locality : Inhaca Is., Delagoa Bay.
Records: Cape (34/22/d) ; Natal (30/30/!); Mocambique (26/32/i).
Distribution : No other records.
Euclyntene glandularis (Day, 1955)
(fig. 30.6.e-i)
Clymene glandularis Day, 1955 : 427, fig. 3 a-e.
Body brown, up to 138 mm. long by 3.5 mm. Prostomium (fig. 30. 6. e) rounded
with numerous eye-spots. Nuchal grooves straight and three-quarter the length of
the cephalic plate. Cephalic rim with 8-10 crenulations posteriorly. Body with
19 setigers and two long achaetous preanals followed by the pygidial ring and
funnel (fig. 30.6.f). Setigers 1-7 increase in length, setiger 8 short, setiger 9 twice
as long and subsequent setigers progressively shorter until setiger 19 is less than twice
as long as broad. The first achaetous preanal is twice as long as the second, the two
together are about as long as setiger 19. Pygidial funnel with 25-30 subequal cirri.
Anus sunken and without a ventral valve. Setigers 2-8 with glandular rings.
Setigers 9 to 14 with a broad dorsal glandular streak (fig. 30.6.!). A faint mid-ventral
glandular streak from setiger 9 onwards. Notosetae include stout, winged capillaries
and fine spinulose forms. Neurosetae of the first three setigers are one to three stout
acicular setae with smooth curved tips (fig. 30.6.J) ; later neurosetae are numerous
hooks with a vertical series of five to six teeth above the main fang and well marked
tendon below (fig. 30. 6. k).
Type locality : Zwartkops Estuary, South Africa.
Records: Cape (from 34/18/i and 34/22/s to 33/25/e and 33/26/i) ; Natal
(30/30/i).
Distribution : Endemic.
Euclyntene mossambica (Day, 1957)
(fig. 30.6.a-b)
Clymene mossambica Day, 1957 : 109, fig. 7 e-f.
Body up to 70 mm. long by 1.6 mm. and pale brown in alcohol. Prostomium
(fig. 30. 6. a) rounded in front. No ocelU. Cephahc plate almost rounded and concave
with a tliick rim with 8-10 well marked crenulations posteriorly. Nuchal grooves
straight and three-quarter the length of the cephahc plate. Body with 19 setigers,
two long achaetous preanals followed by the pygidial ring and funnel (fig. 30. 6. b).
Anterior setigers short, sixth and seventh longer, eighth short, ninth and succeeding
ones long but the last few decrease again until the 19th is twice as long as broad.
First preanal two-third the length of setiger ig, second preanal much shorter and
640
POLYCHAETA OF SOUTHERN AFRICA
Fig. 30.6. Euclymene nwswmbica. (a) Anterior end. (b) Posterior cud. Euclymme glandularis.
(e) Anterior end. (f) Posterior end. (g) Acicular spine of setit»er 2. (h) Normal IiooIl.
(i) Dorsal view of setigers 8-9. Euclymene nalalenns. (j) Neuropodial spine of setiger 2.
(k) Normal hook, (i,) Anterior end. (m) Posterior end.
MALDANIDAE 641
poorly defined. Pygidial ring well marked, funnel with 20 short cirri and a longer
ventral one. Anus sunken and lacks a ventral valve. A stout glandular band at the
anterior margin of sctiger i. Glandular bands also present on sctigcrs 2-7. A narrow
glandular ventral streak from setiger g onwards. Notosetae include stout capillaries
with flattened blades and slender forms with feathered tips. A single neuropodial
acicular seta with a smooth bent tip in each of the first three setigers. Later neuro-
setae are well developed hooks each with a vertical scries of six teeth above the main
fang and well developed tendons below.
Type locality : Mocambique Island.
Records : Not recorded from southern Africa.
Distribution: Mocambique Island (i).
Euclymene annandalei Southern, 1921
(fig. 3o.7.a-d)
Euclymene annandalei Southern, 1921 : 648, pi. 28 fig. 22 A-G, pi. 29 fig. 22 H-K.
Body up to 80 mm. long, pale in alcohol. Prostomium (fig. 30.7.3) bluntly tri-
angular with numerous ocelli. Nuchal grooves straight and two-third the length of
the cephahc plate. Cephalic rim high anteriorly but lower and divided into about
eight crenulations posteriorly. Body with 21 setigers, two achaetous preanals
followed by the pygidial ring and funnel (fig. 30. 7. b). Anterior segments rather
short, setigers 3-8 being little longer than broad ; later segments longer but decrease
over the last few. First preanal well developed and about two-third the length of the
last setiger. Second preanal much shorter, and poorly developed. Funnel with 14
short equal cirri and a longer ventral one. Anus sunken in the funnel and without a
ventral valve. Faint glandular rings on setigers 1-3 ; broad bands on setigers 4-8.
A mid-dorsal glandular streak from setiger g and a narrow mid-ventral streak
from setiger 7 to the pygidium. Notosetae are mainly narrow-winged capillaries
anteriorly but include feathered forms posteriorly. Setigers 1-3 with a single acicular
spine with a smooth bent tip (fig. 30. 7. c). Subsequent neurosetae are numerous
hooks each with a vertical series of five to six teeth above the main fang and well
marked tendons below (fig. 30. 7. d).
Type locality : Chilka Lake, India.
Records : Not recorded from southern Africa.
Distribution: Tropical Indian Ocean (e).
Euclymene luderitziana Augener, 19 18
(fig. 30.7.c-h)
Euclymene luderitziana Augener, 1918: 481, pi. 6 figs. 144-145, pi. 7 figs. 186, 215, text-fig. 77.
Clymene luderitziana : Day, 1955: 427.
Body about 40 mm. long by 0.6 mm. Prostomium (fig. 30. 7. e) as a broad blunt
triangle with numerous eye-spots. Cephalic rim low with a pair of shallow lateral
notches and one median posterior one. Nuchal grooves convergent and three-quarter
642 POLYCIIAF.TA OF SOUTHERN AFRICA
the Icntjth of ihc cephalic phite. Body with 23-24 setigers, two short achactous
preanals Ibllowed by the pygidial ring and funnel (fig. 30. 7. f). Setigers 1-7 increase
in length, setiger 8 shorter than 7, setigers 9-19 subequal, setigers 20-24 shorter.
Second preanal much shorter and less distinct than the first, the two together shorter
than the last setiger. Pygidial funnel with 15-20 subequal cirri and a longer ventral
one. .\nus sunk in the funnel and without a ventral valve. iXotosetae include stout,
broadly-winged capillaries and fine ones with spinulosc blades. Neurosetae of
setigers 1-3 (fig. 30.7. g) as a single smooth, boldly bent acicular spine per neuro-
podium. Later neurosetae are numerous hooks (fig. 30. 7. h) each with a vertical
series of six to seven denticles abo\e the main fang.
Type locality : Luderitzbucht, South West Africa.
Records: South West .Africa (26/15,1, s) ; Cape (32/18,'i, s and 33/18/i, s, also
?34/i8/s).
DiSTRiBUTio.x : Endemic.
PR AXILLELL A YcniW, 1881
Head with a slanting cephalic plate with a raised margin. Nuchal grooves straight
and parallel. Up to 25 setigerous segments and four achaetous preanals. .Setigers 1-3
with two to four stout neurosetae often with vestigial teeth ; posterior hooks numerous
with a vertical series of five to seven denticles above the main fang and chitinous
tendons below. Pygidium with a ring of cirri of which the ventral one is enlarged
Anus on a protuberent cone and provided with an enlarged ventral valve. Glandular
rings on anterior segments.
Type SPECIES : Piaxilla piafterniiyui Malmgrcn, 1866.
Key to Species
1 Setigers 1-3 with two 10 four acicular neurosetae which have vestigial tceili ... 2
- Setigers 1-3 with one acicular neuroseta which has a smooth tip. . P. capensis
2 Nineteen setigers and four achaetous preanals ..... . P. practermissa
- Eighteen setigers and three achaetous preanals ...... P. affinis
Praxillella praetermissa (.Malmgren, 1866)
(fig. 30.7.1-1)
Pra.xilla /irae/ermiao yia\m^Ten, 1866: igi.
Praxillella praelerndssa: .\rwidsson, 1906: 192, pi. 4 figs. 136-143, pi. 9 figs. 294-296, pi. 12 figs.
361-36.1-
Clymene (Praxillella) praelermissa : Fauvcl, 1927 : 179, fig. 62 a-c.
Body up to 100 mm. long. Prostomium (fig. 30.7.1) triangular and blunt, without
ocelli. Nuchal grooves two-third cephalic plate. Cephalic rim low with a pair of
small lateral incisions and a posterior notch. Nineteen setigers, four long achaetous
preanals plus an indistinct pygidial ring and a short fuimel which has a ]3rotruding
anal cone with a large \cntial \alve (fig. 30.7.J). Anal ftmncl encircled by 20-30
MALDANIDAE
643
Fig. 30.7. Euclymene annandalei (after Soulhern) . (a) Head, (b) Ventral view of posterior end.
(c) Acicular spine of setiger 2. (d) Normal hook. Euclymene luderilziana. (e) .Anterior end.
(f) Posterior end. (g) .*\cicu!ar spine of setiger 2. (h) Normal hook. Praxillella praelermissa
(after Fauvel). (i) Head, (j) Posterior end. (k) Acicular spine of setiger 2. (l) Normal
hook. Praxillella cf. affinis. (m) Anterior end. (n) Posterior end. (o) Acicular spine of
setiger 2. (p) Normal hook.
644 P0I,YC:HAETA ok southern AFRICA
cirri of which ihc ventral one is the largest. Two to five acicular neiirosctae in the
first three setigers each with a blunt rostrum surmounted by three to five vestigial
teeth and a vestigial tendon below (fig. 30. 7. k). Later neurosetae numerous, each
with six to seven teetli abo\e the main fang in a \erticai scries and fine tendons
below (fig. 30.7. 1).
Type looalitv : Bahusia, Norway.
Records: No certain southern African record.
Distribution : Arctic ; North Atlantic from Norway to Spain ; Mediterranean.
? Praxillella capensis (Mcintosh, 1885)
Praxilla capensis Mcintosh, 1885 : 405, pi. 25 A fig. 8.
? Praxillella capensis: Day, 1955: 428.
Prostomium prominent but without eyes. C'ephalic rim with a pair of lateral
notches and one posterior one. Nuchal grooves straight and two-third the length of
the cephalic plate. Number of setigers unknown. Anal segments and pygidium
missing. Glandular bands on setigers 5-8. One pair of neuropodial acicular hooks
on the first three setigers. Each with a smooth slightly cur\'cd tip. Posterior neuro-
podial hooks with a single vertical series of six to seven teeth above the main fang.
Posterior notosetae include broad winged capillaries and narrow-bladed forms with
spinulose tips. This is a doubtful species.
Type loc:ality : 180 m. off Gape Town.
Records: Gape (34/1 8/d).
Distribution : Doubtful records from tropical western Africa (s).
Praxillella cf. affinis (Sars, 1872)
(fig. 30.7.m-p)
? Clymene affinis Sars, 1872 : 412.
? Clymene (Praxillella) affinis: F.iuvel, 1927: 180, fig. G2 f-1.
Praxillella cf. ajfinis : Day. 19633 : 430, lig. 10 k-1.
Body up to 23 mm. long without colour markings. Prostomium (fig. 30. 7. m)
bluntly triangular without eye-spots. Nuchal grooves four-fifths of cephahc plate.
Gephalic rim with a pair of faint lateral notches and a single posterior notch. Rim
high anteriorly in front of the lateral notches. Body with 18 setigers and three
achaetous preanals preceding the pygidium which bears a circle of 12 subequal
cirri and ends in a protuberant anus \vith a marked ventral valve (fig. 30. 7. n).
Setigers 1-8 two to three times as kmg as broad, setigers 9-15 about four times as
long as broad, later segments successively shorter, the three preanals being equal in
length to the last two setigers. Setigers 4-8 markedly glandular. A mid-ventral
glandular line from setiger 8 onwards. Setigers 1-3 each with two to three hooks per
neuropodium with three well marked teeth above the rostrum and a faint tendon
below (fig. 30.7.0). Later neurosetae are well developed hooks with a \-ertical series
MALDANIDAE 645
of five teeth above the main fang and obvious tendons below (fig. 30. 7. p). Notosetae
include a few winged capillaries and a few fine capillaries without obvious blades.
Type locality : Off Cape Columbine, South Africa.
Records: Cape (33/1 7/d).
Distribution : P. affinis is known from the North Atlantic (Norway (s) to southern
Ireland (i)) ; Japan.
M ALD ANE Gruhe, i860
Head with a well marked cephahc keel surrounded by a slanting cephalic plate
with a raised margin. Nuchal grooves short, straight or slightly curved. Anterior
segments without collars but with glandular rings. Pygidium with a dorsal anus
below which is a flat anal plate with a low rim which is notched laterally. Neuro-
setae absent from the first sedger but present on all later setigers. Hooks with a
transverse arc of denticles above the main fang. Notosetae include broad and
slender-winged capillaries and finely feathered forms.
Type species : Maldane glebifex Grube, i860.
Maldane sarsi Malmgren, 1866
(fig. 30.8.a-e)
Maldane sarsi Malmgren, 1866 : 188 ; Arwidsson, 1906 : 251, pi. 6 figs. 192-199, pi. 10 figs. 333-338 ;
Fauvel, 1927 : 197, fig. 69 a-i.
Body up to 100 mm. long and encased in a sausage-like mud tube. Cephalic keel
(fig. 30. 8. a) strongly arched and well defined in European forms, broader and more
indefinite in S. African forms. Nuchal grooves (fig. 30. 8. b) short, almost straight.
Cephalic rim low, deeply notched laterally but smooth posteriorly. Nineteen setigers,
two short, indistinct achaetous preanals plus the pygidium (fig. 30. 8. c). Anal plate
oval and slanting, rim chitinised and notched laterally but otherwise smooth.
Notosetae include winged capillaries with markedly tapered blades anteriorly and
winged capillaries plus a few spinulose forms which arc slender and faintly twisted
posteriorly. Neuropodial hooks (fig. 30.8.d, e) have the main fang surmounted by a
transverse arc of 1 2 large teeth above which is a cap of very numerous small dendcles.
Type locality : Sweden and Iceland.
Records: Cape (33/25/s) ; Natal (29/31/s, d, vd) ; Mocambique (24/34/s).
Distribution : Cosmopolitan from the Arctic to Antarctic mainly from deep
dredgings ; habitat mud.
ASYCHIS YJmhcvg, 1867
Cephalic keel low, broad and poorly marked. Nuchal grooves well developed
and curved. Cephahc plate with the rim deeply incised laterally to form two lateral
lobes and a cur\-ed posterior portion. Anterior segments short and wthout collars
646
POI.VCHAETA OF SOUTHERN .\FRU:A
Fig. 30.8. Maldane sarsi. (a) Anterior end. (b) Dorsal view of head, (c) Face view of
hook, (d) Posterior end. (e) Profile of hook. Asydin capemis. (f) Entire worm (twice life
size), (g) .\nterior end. (h) Profile of hook, (i) Posterior end.
but may have marked glantdular rings. Rutiimentary preanal segments present.
Pygidium with a dorsal anus, below which is a slanting oval plate with a low rim
which is either entire or notched laterally. Notosetae include winged and spinulose
capillaries. Xeurosctac absent from setiger I but present on all posterior setigers.
Hooks with a transverse arc of denticles above the main fang.
Type SPECIES : Asychi'. atlantkin Kinberg, 1867
Key to Species
I Cephalic rim deep, forming a pocket with a smooth margin and deep lateral incisions.
Dorsal papillae absent A. capensis
- Cephalic rim low with lateral notches. Two rows of dorsal papillae from setiger 6 onwards
A. dorsofilis*
* Proljably .Urchis Jotuijila shuiild be referred t() the germs Biiuiilnoa\ychis
MAI.DANIDAE 647
Asychis capensis Day, 1961
(fig. 30.8.f-i)
Asychis capensis Day, 1961 : 521, fig. 1 1 e-g.
Body (fig. 30. 8. f) up to 100 mm. long and encased in a toughened mud tube.
Prostomium (fig. 30.8. g) broadly semi-circular in front and without ocelU. Nuchal
slits strongly curved. Cephalic plate a long oval without an obvious cephalic keel.
Cephalic rim deep and divided into three sections by deep lateral incisions ; the
paired lateral flaps are smooth and the median posterior section forms a deep,
smoothly edged pocket covering half of the cephalic plate. Head and cephalic
plate speckled with brown when fresh. Body with ig setigers, one to two poorly
marked achaetous preanals and the pygidial plate (fig. 30. 8. i) which is vertical with
the margin strongly notched laterally and scalloped ventrally. Setigers 1-8 short,
9-13 long and 14-19 again short. Setigers 1-5 difi'usely glandular and setiger 6 with
an anterior glandular band. From setiger 7 onwards there are lateral glandular
ridges. Setiger i without neurosetae ; subsequent sedgers with a row of hooks
(fig. 30. 8. h) each with two transverse arcs of teeth above the main fang, the first arc
with five large teeth and the second with about 20 fine denticles.
Type locality : Saldanha Bay, South Africa.
Records: Cape (from 32/18/s to 34/18/s and 32/28/s). Habitat mud.
Distribution : Endemic.
NOTOPROCTUS Arwidsson, 1906
Head with a slanting plate without a raised rim. Prostomium broad and nuchal
grooves strongly curved. Up to 21 setigerous segments and four achaetous preanals.
No collar on setiger 4. Setigers 1-4 with 1-3 stout acicular neurosetae usually
without any sign of denticles or chitinous tendons; later setigers with a single row
of numerous hooks each with a single vertical series of teeth above the main fang
and chitinous tendons below. Pygidium with a dorsal anus below which is an
inclined oval plate with a smooth margin.
Type species: J^l'otoproctus oculatus Aiwidsson, 1906.
648 POLVCHAETA OF SOUTHKRN AFRICA
Family STERNASPIDAE Clarus, 1863
Body short, swollen and often ovoid or dumb-bell-shaped. It consists of few
segments. Anterior end invaginable with a small prostomium and the first three
segments provided with rows of acicular setae. Segment 7 with genital papillae.
The next eight segments not visibly setose and the final segments with a ventral plate
surrounded with setae. A terminal tuft of branchial filaments.
Records from southern Africa
Stcrnaspis scutata (Renier) .... 5 it's, — Nd
as Sterniupis sciitata \ar. africana Augener . 33AS
RFNLXRK.S
The single genus Sicrnnspis feeds on buried organic matter and burrows head
downwards in stifinuid. The digging organs arc short, stout spines on die first Uiree
segments. The hard anal plates cover the entrance of the burrow and the filamentous
gills can be extended into the water above.
STERNASPIS Om, 1821
Body sausage-shaped, often constricted in the middle. Few segments. Anterior
segments forming an introvert and bearing stout setae. Posterior end with a pair of
ventral horny shields with radiating bundles of setae at their edges. Filiform branchiae
arise from two posterior doisal plaques. Anus terminal.
Type species: Echiiwrhynchus scutatus Renier, 1807.
Sternaspis sctttata (Renier, 1807)
(fig. 3i.i.a-d)
Echinorhrnchiis sciilaliis Renier, 1807 : 34.
Slernn^pis scutala : Fauvel, 1927: 216, fig. 76 a-g.
Body (fig. 31. 1. a) swollen, about 20-30 mm. long \vith 20-22 segments of which
the first seven form an introvert. Surface veKety due to a dense covering of fine
papillae. Prostomium small, papillose and normally retracted into the intro\ert.
The first three segments each have lateral rows of about 12 acicular spines (fig.
3i.i.b). A pair of genital papillae on segment 7. The next eight segments have two
bundles of microscopic setae embedded in the body wall. Striated rhomboidal
shields on the \entrum of the posterior end each with i')-i7 bundles of capillary
setae arising on its outer edges. C^apillaries either fine and smooth (fig. 3i.i.d) or
stouter and pilose (fig. 31.1.C). Branchial filaments numerous, often rolled into
spirals.
Type locality: Mediterranean Sea.
Records: C:ape (33,'25;s) ; Xatal (31/29/s, d) -habitat mud.
Distribution : Cosmnpoliian from the .Arctic to the Antarctic in shallow to very
deep dredgings on mud.
OWENIIDAE A\D STERXASPIDAE 649
Family OWENHDAE Rioja, 191 7
Tubicolous worms encased in sandy tubes. Body elongate and cylindrical and
composed of relatively few segments. Head formed of a fused prostomium and
peristomium. It lacks antennae but is usually provided with a frilled food-gathering
membrane. Mouth terminal. Parapodia poorly developed. Notosetae are spinulose
capillaries and neurosetae are numerous long-shafted but minute hooks arranged in
transverse bands. No anal cirri.
Records from southern Africa
Owenia fusiformis Delle Chiaje . 26.\i\Vs, 27Mi, 28Mi, 32PiCs,
33CS, 34.-^5, 45Pi-\i, 48\VsdCs,
5iCsd
REM.\RKS
Owenia is a suspension feeder which gathers floating particles on a frilly membrane
around its head. There are no palps and the particles are carried along ciUated
grooves to tlie lips which presumably act as sorting organs although the details ha\e
not been worked out. The surface of the body is glandular and probably supplies
tlie tough, almost cartilaginous matrix of the sandy tube. The tube itself projects
well above the surface of the sand in which it is embedded.
Key to Gen-era
I Head rounded and without a feeding membrane or palps .... MYRIOCHELE*
- Head presided with a frilly feeding membrane (fig. 3 I.I. g) .... OWENIA
- Head with a pair of grooved palps ....... . MYRIOWENIA*
OWENIA Dellc Chiaje, 1844
Whole worm encased in a tough tube encrusted with imbricating sandgrains or
shell fragments. Prostomium fused to the achaetous buccal segment and provided
with a frilly food-gathering membrane which surrounds the terminal trilobed mouth.
The first three body segments short with capillary setae only, the rest elongated with
notopodial capillaries and neuropodial rows of minute long-shafted hooks.
Type SPECIES : Owenia fusiformis DtWe C)M^]e, 1844
Oxvenia fusiformis Delle Chiaje, 1844
(fig. 3i.i.e-j)
Ouenia fusiformis Delle Chiaje, 1844: 31 ; Fauvel, 1927: 203, fig. 71 a-f.
^Vhole worm encased in a tough cartilaginous tube strengthened by imbricating
shell fragments or sand grains (fig. 3 1 . i .e) . Body (fig. 3 1 . i .f ) diffusely glandular and
greenish brown; up to 100 mm. long with 20-30 segments. Feeding membrane
(fig. 3i.i.g) mounted on a trilobed base and incised to form six main divisions
surrounding the terminal mouth which has bilobed dorsal and ventral lips. Two
650
I'C.II.VCIIAKTA OF .SOUTHF.RX AFRICA
Fl(i. 31. 1 Slt'rna^pn Mutaln. (a") Ventral view ot'cntire utjrni (three times life size), (b) Anter-
ior aciciilar seta, (c) Posterior pilose capillary, (d) Posterior smooth capillary. Owenia
fusifoTjnis. (k) Sanely tube (twice natural size). (F) Dorsal \'iew ofentire worm (three times
life size), fc;) \'entrol-latcral \ low of anterior enil. (it) T/S mitldle segment, (i, Hook.
(j; Spinulose capillary.
OUEXIIDAE AND STERNASPIDAE 651
ocular marks at the base of the membrane. Thoracic region of three short setigers
bearing capillary setae. The first five abdominal segments much longer (fig. 3i.i.h)
and succeeding ones decreasing progressively in length. Notopodial capillaries with
spinulosc blades (fig. 31.1.J). Neuropodial hooks minute but long-shafted witli two
recurved teedi at the apex (fig. 31.1.1).
Type locality : Naples.
Records: South West Africa (22/34/s, 23/i4;'s, d and 26/14/s, d) ; Cape (from
32/18/s, d to 34/25/s) ; Natal (29/31/i, s) ; Mocambique (26/32/i and 23/35/e).
Distribution : Atlantic from Greenland (s, d) and Sweden (d) to North Carohna
(i, s, d and die Gulf of Mexico (i) ; the English Channel (i, s) then south along the
African coast to Angola (i, s, d) ; Mediterranean and the Red Sea (s) ; Indian
Ocean (i, s, d) ; northern Pacific from the Behring Sea to Japan ; Chile (s).
652
PtHA C:ilAl, lA OF S(U rill.RN AIRICA
FamiK FLABELLIGERIDAE S.iinl Joseph, 1894
15(idy with iL-lati\x-l\' lew segmcnls, sonirliims niarkrcilv t.i]icrcd posteriorly.
Segments essentially similar, surface often papillosi-. Proslomium indistinct and
both It and the buccal segment are normally retracted into a membraneous sheath.
When everted eight or more branchial filaments borne on a dorsal semi-circular
membrane and two large grooved ])alps become \isible. Setae of the first 1-4
segments are often elongated and directed forwards to form a cephalic cage pro-
tecting the buccal apparatus. Parapodia biramous but usually reduced to two
bundles of setae arising directK liom the bod\ wall. Xotosetae are simple annulaled
capillaries. Xeurosctae ma\' be similar to the notosetae or modified In form simple
or compound hooks. Blood green. Anus terminal.
Records from southern Africa
Brada villosa (Rathke) ....
Brada viUosa oi/na^i^ Day
Dijilocinui capensis Day ....
Flalnilinira affini'. Sars ....
as Tcclurclla Indalor Stimpson
as Pherusa tclraguna Schmarda
as C'hioraema tctragona (.Schmard.i) .
as FIdliiiliniiYi afjiiiis v.ir. mareiizi'lU'ii Mcintosh
as Fliihtl I igcid aflinis \ar. luctator (Stimpson)
as Flahtdligtia hiclalm (Stimpson) .
Pherusa lacvis (Stimpson)
as Slrlarinidcs lands (Stimpson)
as Siplionostomiim lacve Stimpson
as Triiphonia xatitholthlia Schmarda
as Flahi'lliuria xaiitlwliicha (Schmarda) .
as Slrlanoidrs .xanlholiichd (Schmarda)
Pherusa momoi (Day) ....
as Stvlarioides nwiiroi Day
Pherusa parmata (Grubi-) ....
as Slylariiddes jtaimalus Grube
Pherusa saldanlia Day ....
Pherusa suakafnana (.Vugener) .
as Sl)darinides sicahnpiarun .\ugener
as Shiarioides xanthalrieha (non Schmarda)
Pherusa trapiea (.\ugener)
as Sirlarinides troji'icus .^ugener
Pirnmis arennsus Kinberg.
as Tra/dionia eapensis Mcintosh
as Sivlarioides eapensis (Mcintosh) .
5 it's
isGi, 1301,
mCI, 26\Vs,
34^Vi
r,lCsd
5iC.sd
;i5Ci, 44C:i.
2C:i
4C;i
3C:i
i3Ci
36C:i
liWi
ibWi
33C:s
.-,,C:s
44C:i
->(:i
4c:i
i6\\i
i.-,c;d
5ic;s
^'-j'SiCs
~.\s
■27.\li, 4oXi
5ic:s
5 1 C^s
2G\Vis, 48C;d\Vs
ifjCid, (pp.)
2r,.\i
3x1, -,ic:s
loCa. ijC'.i
35C:i, 36Ci
(PP'
), 26\Vi
FLABELI.IGERIDAE 653
as Stjlarioides arenosus (Kinberg) . . . 4oNi, 44Ci
Piromis sp. . . . . . . . 5 1 Cs
Pycnoderma congoense Grube .... 33AS
BIOLOGICAL NOTES
The flabclligerids are deposit feeders and use their large frilly palps to collect food
particles from the surface. Possibly the dorsal semi-circle of branchial filaments
may assist in feeding in certain genera such as Diplocinus where they are usually
stout. Some of the genera are tubicolous while others creep about under stones or
burrow just below the surface of the sand. FlabelUgera ajfinis, for example, is found
under loose stones in rock pools and creeps around Uke a caterpillar by means of its
hooked neurosetae. Piromis lives in sandy pools on rocky shores and its sandy crust
provides good protection from small predators as it burrows through the surface
layers. The various species of Pherusa are more sedentary. P. laevis lives in mud-
fiUed crevices or the abandoned burrows of rock-boring bivalves. . Its head, pro-
tected by the cephalic cage of long setae is at the mouth of the tube and its tail is
often folded back against the body so that the faecal pellets can be expelled from
the mouth of the tube. P. swakopiana which is dredged from muddy bottoms also has
a slender tail folded forwards and it is presumed that it too lives in burrows in the
mud.
THE MAIN DIAGXOSTIC CHAR,ACTERS
Reviews of the family will be found in Haase (1915), Fauvel (1927) and Stop-
Bowitz (1948a).
The main diagnostic characters include the number and arrangement of branchial
filaments, the development of the cephaUc cage, the structure of the neurosetae and
finally the nature of the skin papillae and body covering.
The branchial filaments and the structure of the head region. The prostomium is ill-
defined and fused with the pcristomial segment to form the head or buccal apparatus.
In preserved specimens this is usually surrounded by a membraneous sheath and
retracted into the first few setigcrous segments. ^Vhen fully expanded however the
head is seen to include a pair of stout grooved palps on either side of the mouth and a
tongue-shaped or semi-circular cephalic hood which bears a number of branchial
filaments. Inside the cephalic hood and immediately above the mouth is a poorly
defined prostomial swelling which often bears two pairs of eye-spots.
The arrangement of the branchial filaments on the cephaUc hood has rarely been
described since the whole buccal apparatus is seldom extended sufl^icicntly for the
details to be clearly apparent. Nevertheless it is of considerable systematic impor-
tance and in doubtful cases dissection will provide additional evidence. In the
genus Piromis the cephalic hood is short and semicircular. In Brada and FlabelUgera
the thickened margin of the hood bears numerous irregular rows of branchial fila-
ments. In Phenia the margin is narrow and there is only a single row of filaments
though the inrolled lateral margins often give the impression of two lateral bunches
of filaments joined by a single dorsal row. In Diplocirrus there are relatively few
654 I'dlACll Al. lA (IF SOUTI I I'.RX Al RICA
branchiae (lo (ir less) and these arc arranged in an outer group of stouter cirriform
projections and two to foiu- inner ones which are usually but not always more slender
and filamentous.
The cephalic cage is formed of the elongated setae of the first few setigerous segments.
In most species of the genus Brada and in some species oi Diplocirrus these setae are
not obviously elongated and no cage is formed but in most species of Pherusa, Flahel-
ligera and Piromii the cage is well developed and protects the delicate branchial
filaments. The number of setigerous segments which take part in the formation of
the cage is a most useful systematic character and varies from one in Flahelligera
affinis to as many as six in Pherusa parniata. The cage setae are distinguished from
those of the body segments by the fact that they form a compact group of long,
forwardly directed setae and that they are all capillaries whether they are derived
from the notopodial or neuropodial bundle. In contrast, the setae of the normal
body segments are arranged at segmental intervals, they may be directed laterally or
dorsally and the neurosetae are often modified to form hooks. On this basis the
number of segments which contribute to the cephalic cage is most easily determined
by an examination of the neurosetae. It may be added that the number ofindixidual
setae which form the cage is constant within broad limits.
The neurosetae nj the hndy iegnwnls. The notosetae are all annulated capillaries which
are of little systematic value. The neurosetae however .u'c mucii more di\crse. In
Brada, Pycnnderma and Diplocirrus the neurosetae are slightly stouter tiian the notosetae
but not very different in structure. In Pherusa the neurosetae are simple hooks, in
Piromis they are pseudo-compound hooks often with bidentate apices and in Flahel-
ligera tliey are stout compound hooks with sim]ile apices. It should be noted that
not only the shape but also the number of hooks per parapodiuni is imporlanl.
'The skin papillae and hndv envering. The surface of the jjody is usually glandular
and may be covered by debris, a crust ofsand grains or by a translucent mucilaginous
coat which may even be toughened to a cartilaginous consistency. Projecting into
this body covering are skin [xipillac whose shape and distribution is of systematic
importance. Tlicy may be small, wart-like and arranged in a series of annular rings
or may be elongated and club-shaped. Those on the \'entrum are usually the shortest,
tiiiisc (in the dorsiun longer and tiiose around the setae and the cage longer still.
Ki:y to Genera
1 Bofiy ro\''rctl with a dear cartilaginous (iilicle. Neurosetae arc annulaleil < .T])i!!aries
PYCNODERMA (p. 6;,-,)
- Xo clear < artilaginous cuticle. Neurosetae either annulated ca])illaries or hooks . . 2
2 .Neurosetae are simple hooks. Branchiae filamentous and arise in a single marginal row
from the cephalic hood (fig. 32.:j.f ) PHERUSA (p. G3B)
- .Neurosetae are nrit simple hooks. Branchiae filamentous or (irrih>rm and arise in two or
several rows (fig. 32.i.b). ........... 3
3 ,'\ few stfnit branchiae in two distinct groups (fig. 32. 4. f). (Neurosetae annulatetl with
bent or minutely hooked tips) . . . '. . . DIPLOCIRRUS (p. (i(^)
- Numerous fine branchiae in several irregular rows ....... 4
FLABELLIGERIDAE 655
4 Body covered with a mucilaginous sheath (fig. 32.1.3). Neurosetae are stout compound
hooks FLABELLIGERA (p. 655)
— Body covered with adhesive papillae (fig. 32.i.k). Neurosetae with annulated shafts and
smooth tips. (Prominent nephridial papillae) .... BRADA (p. 656)
- Body encrusted with sand or debris (fig. 32.4.3). Neurosetae are pseudocompound hooks
and often bidenlate ......... PIROMIS (p. 663)
FLABELLIGERA Sars, 1829
Body flabby, translucent and covered with a thick mucilaginous sheath supported
by long pedunculate papillae. Branchiae filamentous and all similar ; the filaments
arise in se\eral irregular rows from the thickened margin of the triangular cephalic
hood. Cephahc cage well developed and formed of the elongated notopodial and
neuropodial capillaries of setiger i. Subsequent notosetae are all annulated capil-
laries. Neurosetae of setiger 2 and subsequent segments are one or more stout
jointed hooks.
Type species : Flabelligera affinis Sars, 1829
Flabelligera affinis Sars, 1829
(fig. 32.i.a-f)
Flabelligera affinis Sars, 1829: 31, pi. 3 fig. 16; Fauvel, 1927: 113, fig. 40 a-f; Day, 1961 : 505,
fig. 8 a.
Body (fig. 32. 1. a) up to 60 mm. long with about 50 segments entirely covered with
a thick mucilaginous sheath supported by long pedunculate papillae (fig. 32.i.d).
Cephalic cage rather short and formed by 50 — 50 capillaries of setiger i. Notosetae
of setiger 2 and subsequent segments are about six annulated capillaries (fig. 32.i.f)
supported by elongate club-shaped papillae. Neurosetae are one to two stout com-
pound hooks per segment (fig. 32.1.C) with transversely striated shafts and a jointed
brown apex (fig. 32.i.e). Two groups of 30 or more fine branchial filaments arise
from the thickened margin of the triangular cephalic hood in several irregular rows
(fig. 32. 1. b). Two short frilly palps. A median cephalic ridge on the inner side of
the cephalic hood with a pair of conical papillae at its base. Two pairs of large
indistinct eyes almost fused. Puckered lips around the mouth.
Type loc.\lity : Bergen.
Records: South ^Vest Africa (26 15/i, s); Cape (from 29/16/i and 34/18 i, s to
34/23, e, i and 32/28/i).
Distribution: Arctic; Atlantic from Norway (s, d) and Greenland (s, d) to the
English Channel (i, s), Canary Is. (i), Senegal (i, s) ; Falkland Is. (s, d) and
South Georgia (d) ; far northern Pacific (Bchring Sea to N.^V. Japan).
PYCNODERMA Grube, 1877
Body long, slender and covered with a tough cartilaginous sheath in which a few
elongate papillae are embedded. Cephahc cage formed by the setae of sedgcr I
although the setae of the next few segments are also elongate and directed forwards.
Two groups of fine branchial filaments. Notosetae are simple annulated capillaries.
656 POI.YCniAETA OF SOL THKRX ATRICA
Ncurosctae are annulated capillaries stouter than liic iKitosetae. ('rphalli hood with
numerous slender branchial filaments. A ]3air of shoit grooved palps.
Type species: Pj'cnoderma congoeruc Grubc, 1877.
Pycnoderma congoense Grube, 1877
(lis. :?-'-i-s-i>
Pycriodtrnm conoat'iist' Gvuhc, I1I77.1 : 540; Monro. 1930; 162. fi^. 63 a-cl.
Body (fig. 3'J.i.,e;) about 50 mm. long, and 2 mm. broad with about 70 segments;
it is not swollen anteriorly or markedly tapered posteriorly. Surface covered with a
cartilaginous cuticle from which long, fine, knobbed papillae (fig. 32.1.J) project
here and there. CVphalic hood with numerous slender branchial filaments in two
groups. A pair of short grooved palps and three well developed lips, one dorsal and
two \entro-lateral. Cephalic cage poorly developed and composed of tlic notosetac
and neurosetae of setiger i plus the notosetae of setigcr 2 but not the neurosetae,
though these hke the setae of the next few segments are directed forwards. Notosetae
of normal body segments are slender annulated capillaries with flagelliform tips
(fig. 32. 1. i). Neurosetae are similar but stouter (fig. 32.1.11).
Type locality : Congo coast.
Records : Not recorded from southern Africa.
DisTRiBUTio.N' : Tropical western Africa from Liberia (s) to Angola (s).
BRAD A Stimpson, 1854
Body more or less fusiform and maggot-like with few segments. .Surface papillose
and often covered with sand or debris. Cephahc hood poorly developed. Numerous
similar branchial filaments in three to foiu' irregular rows. A pair of palps. ,'\ntcrior
setae do not form a cephahc cage. Notosetae are feeble annulated capillaries.
Neurosetae stouter, with more closely annulated shafts and non-annulatcd ti]5S. A
pair of prominent ncphridial papillae on setiger 4 or 5 of the adult.
Type species: Siphiiiiiniiniuj nllnsuni Rathke, 1843.
Brada villosa capensis Day, 1961
(fig. 32.i.k-q)
Brada villom cafiemis Day, ig6i : 510, fig. 9 g-m.
Body (fig. 32. 1. k) maggot-like, about 15 mm. long for 36 segments ; shape broadly
cylindrical with truncate ends. Dorsal surface with long cylindrical pajiillae about
eight times as long as broad (fig. 32.1.0). Papillai' on parapodia even longer but
ventral papillae small, cylindrical, about three to four times as long as broad (fig.
32.i.n1. Whole surface sandy, especiaUy the dorsum (fig. 32.1.111). Setae of first
segment elongated but not forming a cage, aboui eiuhi in the notopodium and lour in
till' neuropodiuiii. Liiter par.ipudi.i with ab(.)ut six lung sleiidi'r notopodial capillaries
FLABELLIGERIDAE
657
Fig. 32.1. FlabeUigera ajfmis. (a) Entire worm (twice life size), (b) Anterior view of head.
(c) T/S body segment, (d) Details of elongate papillae embedded in the gelatinous coat.
(e) Hook, (f) Capillary seta. Pycnodermn congomse (from Monro, 1930). (g) Entire worm
(twice natural size) (cartilaginous sheath omitted), (h) Xeuroseta. (i) Notopodial capillary,
(j) Papillae embedded in sheath. Brada villosa capemis. (k) Entire worm (three times natural
size), (l) Anterior view of head, (m) T/S body segment, (n) Ventral papilla, (o) Dorsal
papillae, (p) Xeuroseta. (q) Notopodial capillarv'.
G58 rOI.VCHAETA OF SOUTHERX AFRICIA
annulatcd lo tlieir fine tips (fig. 32.1.C1) and abdut foiu' to five shorter, stouter neuro-
podial eapillarics with the proximal half closely anniilated and the distal half not
annulated (fig. 32.i.p). Setiger 5 with a stout ne])hridial pajiilla antero-vcntral to
the nenropodiiiin. Buccal apparatus (fig. :V--i-ll with a poorly developed cephalic
hood bearing about 30 to 40 short branchial Hlann-nls arranged in a dorsal arc three
to four deep, a pair of short, stout, grooved palps, an indistinct prostomial swelling
without eyes and a ventral mouth with two lateral lips and one ventral hp none of
which are swollen.
Type locality : Agidhas Bank, South Africa.
Recxirds : South West Africa (22/14/s, d) ; C\pr IVom 3'2/17/d to 35/20/d and
34'22;s).
Distribution : The subspecies B.r. capends is probably endemic.
PHERUSA Oken, 1807
Body cylindrical anteriorly but often narrowed to a well marked tail posteriorly.
Surface papillose and often covered with sand and debris. Introvert with a pair of
grooved palps, two or three cushion-shaped lips and a dorsal cephalic hood bearing
numerous similar branchial filaments along its edge. A well developed cephalic
cage formed by the elongated setae of the first two cjr three segments. Notosetae are
annulated capillaries. Xeurosetae of the head region are annulated capillaries but in
the body region they are simple hooks.
Type SPECIES : Am/i/iilritt- J>liimosa .Muller, 1776.
Key to Species
1 Simple hooks I'.'place the annulated capillaries in neiiropodium ;;.... 2
- Simple hooks replace the annulated capillaries in ncuropodium 4 . . . . 3
- Simple hooks replace the annulated ca|)illaries in neuiopodium 6 or 7. (A hard sandy
shield on the head) (fig. 32. 2. a) P. parmaf a {p. Gj''>)
2 Xeuropodium 3 with a slender elongated hook (fig. 3-'.j.g) ; stout hooks from scliger 4.
Skin with a few sinall wart-like papillae in two rows ... P. monroi (p. 660)
- Xeuropodium 3 with a short stout liook. Skin uitli numerous long papillae giving a
shaggy appearance ........ P. su'ako2>in>ia [p. 6G1)
3 Body with numerous papillae covering the surface ....... 4
- Body with two irregular rows of wart-likc papillae per segment . . P. laevis (p. 6bi)
- Body without skin papillae ; branchial filaments few and stout . . P. sp. (N.\D 30B)
4 Papillae long, giving a shaggy appearance. Cage setae numerous. Many branchial fila-
ments P- tropica (p. 6G3)
- Papillae short giving a velvety appearance. Cage setae few. Few branchial hlaments
P. saldanha (p. b()3)
Pherusa partnata (Grube, 1878)
(lig. 32.2.a-c)
.S7)'/,;no/r/,t /wrm./^Kt Grube, ifiyo: nj'J. l^l. n hg- i; I'"au\el, 1953: 34I1, fig. 179 b.
I'.odv cylindrical, about 30 mm. long for 60 segments but tapered posteriorly from
abciitt the 2^)th to form a narrow tail. Setiger i with a median bifid process on the
anterior margin. .\ well marked sandy shield on the head (fig. 32. 2. a) extending
FLABELLIGERIDAE
659
Fig. 32.2. Phertisa partnala. (a) Dorso-lateral view of anterior end. (b) Hook from middle
neuropodium. (c, c', c-) Capillary seta. Phertisa momoi. (d) Entire worm (four times
natural size), (e) Two body segments to show skin papillae, (f) Hook from middle of body.
(g) Elongate hook from neuropodium of setiger 3. Pherusa swakopiana. (h) Entire worm
(twice natural size), (i) Hook, (j) Details of anterior end. (k) Skin papillae, (l) Ventral
view of buccal apparatus (dissected).
66o l'()l.^ CIIAI. lA Ol' SOL'lIir.RX AFRICA
from sctigcr 2 lo 4. Body otherwise naked apart from two sparse rows of wart-likc
papillae per segment. ('.e])halic cage well marked and formed of 38 + 38 very fine
long setae of the first two segments. Setigcrs 3-6 do not form part of tiie cage but
each has three to fi\-e fine capillaries (fig. 32. 3. c) in both rami. A single stout simple
hook (fig. 3'J.'2.b) appears in the ncuropodium of setiger 6 or 7. Behind this the
number of hooks slowly increases to four at tlie i-nd of the tail but most segments
have two. The buccal apparatus includes two long grooved palps, a cephalic hood
rolled in at each side and bearing a single row of 15 + 15 branchial filaments on
either side of the cephalic ridge. Eyes not seen. One dorsal <uid two \entro-latcral
flattened lips around the mouth.
Type locality : Philippine Is.
Records: Natal (-'ij 31;!, s).
DiSTRiBUTiox : Indo-west-Pacific from Madagascar and Madras to Philippijic Is.,
and Xew Zealand.
Pherusa ntonroi (Day, 1957)
(fig. 32.2.d-g)
Slylarioides monroi Day, 1957 : 103, fig. 6 n-p.
Pherusa monroi : Day, 1961 : 505,
Body (fig. 32. 2. d) small, seldom exceeding 20 mm. for 60 segments. It is cylindrical
and tapered evenly to a distinct tail. Adherent sand grains on the first two segments
but thereafter the body is naked apart from two irregular rows of small wart-like
papillae per segment (fig. 32. 2. e). Gephalic cage \vell marked and formed by
33 + 33 setae of the first two sctigers. Notosetae of body segments are two to three
small capillaries per segment. The neuroseta of segment 3 is a single long slender
hook (fig. 32. 2. g) ; a stout simple hook appears on setiger 4 (lig. 32. 2. f) and there is
one (or occasionally two) on all succeeding neuropodia. A bifid process at the dorsal
entrance to the introvert. The buccal apparatus consists of a pair of stout frilled
palps, three short Ups around the mouth and a dorsal cephalic hood rolled in on
cither side and bearing a single continuous scries of 14-16 branchial filaments. Four
eyes on the cephalic ridge on the inner side of the hood.
Type loi:ality : Imbotje, southern Xatal coast.
Records: South West Africa (26/15/i, s) ; Clape (from 32/18/s and 34/ig/s to
32;28;s); Natal (si/^o/i)-
DisrRiBUTKjx : .Soutlu-rn Arabia (s).
FLABELLIGERIDAE 661
Pherusa swakopiana (Augener, 1918)
(fig- 32-2.h-l)
Slylarioides swakojiianus Augener, 1918 : 433, pi. 7 fig. 234, text-figs. 61-62.
Pherusa swakopiana : Day, 1 96 1 : 506.
Body (fig. 32. 2. h) up to 40 mm. long for 50 segments of which the anterior 20-25
form the stout, sausage -shaped body and the last 30 measuring 10 mm. form the
slender tail. Body covered with elongate papillae (fig. 32. 2. k) giving a shaggy
appearance. Cephalic cage (fig. 32.2.J) formed of the long setae of setigers i and 2,
each of the four rami contributing about six to eight to the total of 28 on each side.
Cage setae supported by very long papillae. Buccal apparatus (fig. 32. 2. 1) includes
a dorsal cephahc hood roUed in on each side and bearing a single continuous row of
about 100 flattened branchial filaments. No eyes. A pair of long grooved frilly
palps. Mouth with a long tongue-shaped dorsal lip above and two rather shorter
ventro-lateral lips below. The whole buccal apparatus is often retracted into a long
intro\ert.
Body segments with three to four fine notopodial capiUaries. Ncuropodial hooks
start on setiger 3 as one per segment until setiger 1 7 or 18 and up to four thereafter.
Each is stout and has a curved, pointed tip ; shaft finely striated (fig. 32.2.1).
Type locality : Swakopmund, South West Africa.
Records: South West Africa (22/14/i, s to 26,15,1); Cape (from 32,18,5 and
33/18/s, d to 33/25,/s) ; Natal (31/29/s).
Distribution : Morocco (s) ; Tristan da Cunha (s) ; Falkland Is. (s) ; South
Georgia (s).
Pherusa laevis (Stimpson, 1856)
(fig. 32.3.a-c)
Sifihonoslomiim lane Stimpson, 1856: 391.
Trophonia xanlhoirkha Schmarda, 1861 : 16, pi. ig fig. 165.
Slylarioides xatitholricha : Augener, 1918: 430, fig. 40.
Slylarioides laeris : Day, 1 955 : 42 1 .
Body rather small, seldom exceeding 30 mm. for 65 segments ; it is stout anteriorly
but tapers evenly to form a well marked trail. Adherent sand grains on tlic anterior
end but the rest of the body is naked apart from two irregular rows of rounded
papillae per segment (fig. 32. 3. b). Cephahc cage (fig. 32. 3. a) well marked and formed
of 44 + 44 setae of the first two segments. Setiger 3 with three to four fine capillaries
dorsally and two to three ventrally but these are separate from the cephalic cage. A
tentaculiform sensory appendage ending in three to four short processes above the
introvert. Buccal apparatus consisting of a cephalic hood rolled in on each side and
bearing a continuous row of about 35-40 branchial filaments. Two pairs of eyes. .\
pair of long groo\ed palps. Three cushion-like Ups, one above and two below liie
mouth.
Xotosctae of the fourth and subsequent segments are three to four small capillaries.
66-2
POI.VCHAETA (IV SOUTH KR\ AFRICA
Xcuropodial liooks start on sctiger 4. There is one per segment throughout the body
and each is a stout simple hook with faint transverse striations (fig. 32.3.0).
Type locaijty : False Bay, South Africa.
Records : .South'West .Africa {'2-2 14 i to 28 16 i, s) ; Clape (from 29/16/i to 34'18/i,
s and 34/24 i).
Distribution : Endemic.
a
; /
Va»^
Sk
,
•f^^':!'
^^^^^,
T^**^
Fig. 32.1. Pherusa laetis. (a) Lateral view of head, (b) Lateral view of two segments showing
papillae. Ic.j Hook. Pherusa Iwpicd. (d) Hook (after .Aii^ener, 1918). Phmisa ^aldanha.
(k) F.ntirc worm (three times life size), (f) .-\ntcrinr \ iew of buccal apparatus (dissected).
((;) .Setae and skin papillae, (u) Hook.
FLABELI.IGERIDAE 663
Phertisa tropica (Augener, 191 8)
(fig. 32.3.d)
Slylarioides tropicus Augener, 1918 : 437, pi. 6 fig. 147 ; pi. 7 figs. 220-221, text-fig. 63.
Body about 25 mm. long with 30 segments. It is sausage-shaped anteriorly with a
narrow tail. Surface shaggy due to a dense covering of long papillae. Cephalic
cage formed by 24-31 long capillaries of the first two setigers. Cephalic hood with
50-60 branchial filaments. Notosctac of body segments are five to eight capillaries
per bundle. Neurosetae of setiger 3 are capillaries. Stout neuropodial hooks start
on setiger 4, at first one to two, posteriorly three to four per segment. Each hook
has a shghtly cursed and pointed tip (fig. 32. 3. d).
Type locality : Landana, Congo coast.
Records : Not recorded from southern Africa.
DiSTRiBUTio.N : Tropical western Africa from Liberia (i) to Angola (i).
Pherusa saldanha Day, 1961
(fig. 32.3.e-h)
Pherusa saldanha Day, 1 96 1 : 508, fig. 8 b-d.
Body (fig. 32. 3. e) up to 30 mm. long for 65 segments. It is stout anteriorly and
tapers evenly to a slender tail. Surface velvety with very numerous, small papillae
(fig. 32. 3. g) and rings of slightly longer ones marking the segments. Cephalic cage
formed by a few stout setae of the first two setigers, and total about 17 on each side.
Cephalic hood (fig. 32.3.f) with 32 branchial filaments in a single marginal row.
Three well-marked cushion-hke hps. Two palps. Setiger 3 with one to two small
capillaries in each ramus. Succeeding segments with a single capillary in the noto-
podium and a single simple hook (fig. 32. 3. h) in the neuropodium.
Type locality : Saldanha Bay, South Africa.
Records: Cape (33/18/s and 34/22/s).
Distribution : No other records.
PIROMIS Kinberg, 1867
Body elongate and tapered posteriorly. Surface papillose \vith a sandy crust.
Buccal introvert with a pair of grooved palps and a cephalic hood bearing two groups
of numerous similar branchial filaments arranged in several irregular rows. Pro-
stomial lobe with four eyes. Cephahc cage well developed but poorly defined and
formed by the long setae of the first few segments. Notosetae are crenulate ca])illaries.
Neurosetae of the cage are similar but later ones are pseudocompound hooks ending
in unidentate or bidentate apices.
Type species: Piromis arenosus Kinberg, 1867.
664 POI.Vt;ilAi: lA OK SOUTHERN AKRICIA
Piromis arenosus Kinbcrg, 1867
(fig. ;v-'4.a-d)
Piromis arenosus Kinbcrs, liitiy: 358: Hariinan, nil'' : 1'7. pi- ij figs. ~~<)\ Day, iqGi: 509,
fig. 8 c.
Tropfuinifi cci/ifn^i\ .\U Intush. 1885 : 363. pi. 44 figs, --{i, pi. 23. \ tigs. 1-3.
S(vlttrioi(/i's cti/'tnsis : Mtmro. i't33: joj ; Day, K)34 ; tyj.
A large species reacliing cjo mm. with about 60 segments. Body (fig. 32.4.a)
evenly tapered posteriorly and without a distinct tail. The whole surface is encrusted
with sand and when this is peeled off each segment is seen to have a few cylindrical
papillae. There are four dorsally, four ventrally (fig. 32.4.13) and two on each side
between the rami of the parapodia. Cephalic cage poorly defined and may be
said to consist of the setae of the first two or three setigers although the setae of the
fourth .uc not much shorter and are also directed forwards. Setiger i has nine dorsal
and se\-en ventral capillaries, setiger 2 has seven dorsal and five \entral, setiger 3 has
about six dorsal and five long stout setae ventrally, some of which may end in
minute bidcntate hooks. There. iftcr .ill ii()ti)]iodi.i ha\e four to six capillaries and
all neuropodia have four to eight multi.irticuhite hooks (tig. 32. 4. d, d') which end
in either uuidentate or bidentate tips. The buccal .ipparatus 1 lig. 32.4.ci is ntr.ictile
and there is a dorsal tcntaculiform process ending in two to six ]5apillae on setiger i
over the introvert. Buccal sheath low and membraneous. C^ephnlic hood tongue-
shaped and bears on its ventral surilice two elongate groups of about too tentacular
filaments. Indi\idual filaments cylindi-ical and ringed \vith cilia. Paljjs long and
groo\"ed. Prostomial loljc with four e\es. .-V semicircuhir clurs,il lip o\erhanging the
mouth and two \eiUro-later.d ones.
Type loi Aiarv: Port Xatal ( -^-- Durban), South .Xfrica.
Re( ords : C^ape ; from 53 iS i, s to 32'28,i, s) ; Xatal {from 31 29/i to 29 31/i).
DiSTRiBUTio.N : Red Sea; ? Uruguay (s).
Piromis sp.
Piruiuis sp. Day, Ii)6i : jog.
.\ damaged specimen with .in inconijdrie head but no sign ol'a ciplulic cige. .\
thin film of mud and mucus I'orms a delicate coat which co\i-rs the IjocIv. Below this
a scattering of conical pajiilLu', longer later,illy and mme numerous \cntr,illy.
Xotosetae are four to fi\e stout capillaries with long internodcs between .mnulalions
and fine fl.igelliform tips. Xeurosetae are f >iir to li\-e ]iseudocompound hooks ending
in falcate unident,ite blades.
DIPLOCIRRUS n \isc, Kji-, (emeiul.)
Body c-liing,ile and gently tapered. Posterior segments well defined. Surfice
pa]3illiise and with adherent s.ind or debris. ]5uccal appar.itus with a pair of '^frooxed
palps, three lips and a ceph.ihc hood bearing a fi-w stout tentacels along its edge anp
a second inner group which are usually more slender. Prostomial lobe with four
FLABELLIGERIDAE
665
Fig. 32.4. Piromis arenosus. (a) Entire worm (twice life size), (b) Ventral view of head end
with the sandy coat removed, (c) Ventral view of the dissected buccal apparatus, (d, d')
Xeuropodial hook. Diplocirrus capensis. (e) Worm with posterior end missing (four times
life size), (f) Anterior view of dissected buccal apparatus, (g) Papilla from the ventral
surface, (h) Papilla from the dorsal surface, (i) Xotoseta. (j, j') Neuroseta.
666 I'()LVt:HAKT.\ OF SOUTIIERX AIRICA
eyes. Cephalic cage \aiial)lc, sonictiincs abscnl. Rolli iiotosclac and iicindsctac arc
boldly aiinulatcd, the iiotosetac ha\iiig fine hair-like tips and the neurosetae stouter
with bent or minutely hooked tips. .\e]5hridial papillae sometimes seen on setiger 5.
Type SPECIES : 'liojtluiiiia s.Iauca Malmgren, 18G7.
Diplocirriis capensis Day, 1961
(fig. 32.4.e-j)
Diplochtin cal'itiiii Day. 1961 : 509, fig. g a-f.
Body (fig. ;52.4.e) arenicoliform, about 12 mm. long, with well defined posterior
segments and the whole surface covered with flask-shaped papillae which are longer
dorsally (fig. 32.4.h) than ventrally (fig. 3-'.4.g). No cephalic cage. No visible
nephridial papillae. CephaUc lobe (fig. 32-4-f) semicircular with a median ridge
bearing eyes at its base. Eight stout Isranchial cirri of which six arise from the margin
of the tentacular lobe and two of the same size or slightly stouter arise from its inner
surface. A pair of grooved palps twice as stout as the tentacles. Mouth Ncntral with
poorly detined lips. Both notosetae (fig. 32.4.1) and neurosetae (fig. 32.4.)) are
annulated but the neurosetae are stouter than the notopodial capillaries and end in
minutely hooked tips.
Type LotiALrrv : (JlT Port Elizabeth, South Africa.
Records: Clape (from 34/23/s, d to 34/25/d).
Distribution : Endemic.
SABEI.L ARIIDAE
667
Family SABELLARIIDAE Johnston, 1865
Tubicolous worms Living in dense sandy tubes. Head crowned with an opcrciiknn
of golden palcae. Prostomium indistinct and hidden between the opercular lobes
which may be completely fused dorsally but always remain separate ventrally and
bear numerous buccal cirri on either side of the mouth. A cirriform lobe and a pair
of grooved palps hidden in the opercular cleft in front of the mouth. Body divided
into four regions: First region of two short uniramous segments with ventral capil-
laries only. Second (parathoracic) region of three to four biramous segments with
dorsal cirriform gills, stout oar-shaped notosetae and capillary neurosetae. Third
(abdominal) region of numerous biramous segments with dorsal cirriform gills,
notopodial serpuliform uncini and neuropodial capillaries. Fourth (caudal) region
of rudimentary achaetous segments reflected forwards under the abdomen.
Records from southern Africa
33Ci. 35Ci, 36Ci,
4oNi, 44Ci, 480,
51CS
4Ci
loCi, I iWi, i3C;i,
i6Wi, 26\Vi
39^V'i
4oNiPi, 44Ci, 45Pi
I Pi, 45Mi
I yMi
— Ps
4oNi
32Cd
55Ca
— Ns
27Mi, 28Mi,
4oNiPi, 45Pi
26Ai
32Pi
— Ns
SGCs, 4oNi, 44Ci,
48^Vs, 5iCsd, — Nsd
26Ai, 48\Vs
26VVS
n strong sandy tubes attached
to various objects. Most of them grow on rocks but the smaller species may be
Gunnarea capensis (Schmarda)
as Hermella capensis Schmarda .
as Sabellaria capensis (Schmarda)
as Pallasia capensis (Schmarda) .
Idanthyrsus pennatus (Peters) .
as Pallasia pennata J'eters .
as Cryptopomatus geayi Gravier .
Lygdamis indicus Kinberg
as Lygdamis indicus var. gihiiristi Day (nor
Mcintosh) .....
Lygdamis murata var. gilchrisli (Mcintosh)
as Tetreres murata var. gilchrisli Mcintosh
Phalacrostemma elegans Fauvel
Sabellaria intoshi Fauvel
as Sabellaria spinulosa \'ar. intoshi Fauvel
as Sabellaria guinensis Augener .
as Sabellaria spinulosa var. gilchrisli Mclntosl
Sabellaria peclinaia Fauvel
Sabellaria spinulosa alcocki Gravier .
Sabellaria spinulosa eupomatoides Augener
as Sabellaria eupomatoides Augener
Sabellaria spinulosa fucicola Augener
as Sabellaria fucicola Augener
BIOLOGICAL XOTE.S
The sabellariids are suspension feeders and live
668 I'(ii.vc:h.\eta or sol' riii'.RX .\iric:a
found on the shells of living molluscs or on the fronds of algae. Although single
specimens are often found, most species are gregarious and Gmmarca capensis forms
sandy reefs covering hundreds of square metres on the shores of the western Cape.
The tropical Idanthvrsin pcnniili/s is less prolific but still encrusts many rocks on the
Natal coast.
(hnmaira reefs seem to be best developed at the level of low water neaps and the
surface of the reef is quite dry at low water of springs. At this time individual tubes
are plugged with a crown of golden paleac or llattened setae which cover the surface
of the head and serve both to protect the worm from predators and prevent dessica-
tion. ^\■hen the tide rises and tire tubes are submerged, the head is pushed out and
arched back so that the two opercular lobes are distended to reveal numerous buccal
cirri around the mouth. These bear rings of cilia and catch suspended food particles
which are conveyed along grooves to the mouth. The mechanism whereby the
heavier silt particles are eliminated is not known but the grooved palps seem to be
mainly concerned in tube building. Each tube is lined with a black secretion and
closed at the posterior end. The slender tail end of the worm is bent forwards
against the body so that the faecal pellets are voided from the mouth of the tube.
THE MAIN DIAGNOSTIC CH.-\RACTERS
Revisions of the morphology and systematics of the family Hermellidae ( = Sabcl-
lariidae) are given by Johansson (1927) and Hartman (1044b). While the most
important systematic features are found on the head, the number of parathoracic
segments bearing oar-shaped setae is of generic importance and the number of teeth
on the uncini differs slightly. Though the prostomium is indistinguishable and
hidden in the cleft between the opercular peduncles, it may have a free cirriform
projection or this may be absent. A pair of grooved palps is always present in front
of the mouth but the munber of lobes bearing the buccal cirri ditfers from species to
species and is constant within limits. The opercular lobes may be quite separate
dorsaUy or fused to varying degrees and their ends which bear the paleae may be
long, fleshy and sloping or vertically truncate and entirely covered by the paleae.
There may be a pair of stout dorsal hooks at the base of the opercular cleft or only
small acicular setae immediately behind the external row of paleac or none at all.
The opercidiun itself consists of one to three rows of paleae (external, middle and
inner rows) and the shape of the indi\idual paleae is constant apart from wear and
the degree of development of the median tooth in the external row ; thus the dorsal
ones are often better de\eloped than the lateral ones.
Key to Genera
1 One rowof iMleae (fig. :i3.i.b) PHALACROSTEMMA [p. be>j)
- Two or tliree rows of paleae ........... 2
2 Three rows of paleae. (Three parallioracic segments) ...... 3
- Two rows of paleae ........•••• 4
3 Middle row of paleae forming a cone wliich ci>nccals ilie inner low PHRAGMATOPOMA*
- Middle row of paleae not concealing the inner row (tig. 33.1. i) . SABELLARIA (p. hyi)
4 Opercular peduncles completely fused dorsally and lack hooks or acicular setae
GUNNAREA (p. 673)
- Opercular peduncles not fused and have a pair of stout hooks basally (fig. 33. 2. j) . . 5
SABELLARIIDAE 669
5 Three parathoracic segments. Outer paleac bipinnate or serrated (fig. 33. 2. k)
IDANTHYRSUS (p. 675)
- Four parathoracic segments. Outer paleae smooth . . . . L YGDAMIS (p. 675)
PHALACROSTEMMA Marcnzcller, 1895
Opercular peduncles elongated. Opercular crown formed by a single circular row
of long palaca and a few short acicular setae. A pair of stout hooks at the dorsal
junction of the opercular lobes and one to three cirri at the ventral junction. A pair
of grooved palps. Few buccal cirri.* The first two segments with fine neuropodial
capillaries, then four biramous parathoracic segments followed by about 12 abdom-
inal segments with uncini in the notopodial pinnules and capillaries in the neuro-
podia. Caudal region smooth and achactous. Cirriform branchiae from the second
setiger to the middle of the abdomen but may be lacking on parathoracic segments.
Type species: Phalocrostemma cidariophilum Marenzeller, 1895.
Key to Species
I Opercular peduncles very long. A single large median cirrus between the opercular
peduncles (fig. 33.i.b) ......... P, elegans
- Opercular peduncles short. Two to three short cirri between the opercular pedimcles
P. cidariophilum*
Phalacrostemma elegans Fauvel, 191 1
(fig- 33-i-i-g)
Phalacrostemma elegans Fauvel, igi la : 3, fig. 3 ; Fauvel, 1914a : 270, pi. 24, figs. 1-16 ; Day, 1963 :
367-
A small abyssal species (fig. 33. i .a) about 1 7 mm. long, living in a tube of foramini-
feran shells. Opercular peduncles (fig. 33.1 .b) very long and separate, each bearing a
single circular row of very long, spirally serrated paleae (fig. 33.i.g) and one to two
much shorter stouter paleae (fig. 33.i.d). A pair of short stout hooks (fig. 33.i.e)
at the dorsal origin of the opercular peduncles. A single large median cirrus between
the opercular lobes and a pair of grooved palps. No true buccal cirri but four cirri-
form papillae on the ventral surface of each opercular lobe. The first two setigers
bear capillary neurosetae only, the next four parathoracic segments bear notopodial
oar-shaped setae (fig. 33.1.C) and neuropodial capillaries and the final lo-ii
abdominal segments bear six-toothed uncini (fig. 33.i.f) in the notopodial pinnules
and capillaries in the neuropodia. Cirriform branchiae from setiger 2 to about the
sixth abdominal segment. Tail achaetous and reflected forwards.
Type locality : 1,968 metres off Madeira.
Records: Cape (34/17/a).
Distribution : Madeira (a) .
*Fauvel (1927, p. 212) stated in his definition of the genus: "Pas de paqucts dc tentacles filiformes autoiir
de la bouche". There are certainly no large groups of buccal cirri but thciT are a few homologous buccal
cirri in P. elegans. It is difficult to understand how P. cidariophyllum, the type species of the genus, can feed
without them.
(i70
rOI.VCHAKTA OF SOITHERX AIRIOA
Fig. 33.I. Phalacro'ilcmma elegant (after Fau\-f*I, 19141. fAi Lateral \'iew of body (four times
life size), (b) N'entrai view of anterior end. {cl CJar-shaped seta, (d) Short, stout palea
from opercular crown, (e) Hook from opercular peduncle, (f) Uncinus. (o) Serrated
palea from opercular crown. Sabellaria inloshi. (n! Head end protruding from tube.
(li Operculum. {]) Outer palea in plan, (k) Middle palea in plan, (l) Middle palea in
profile. IM) Inner palea in profile. Sabellaria sjnmiloia cujmmaioidii (from .\ugcncr). (n) End
of outer palea. (nj Middle palea.
SABELLARIIDAE 671
SABELLARIA Savigny, 18 18
Opercular peduncles mainly fused. Operculum almost at right angles to body.
Opercular crown formed of three rows of golden paleae, the external row flattened
with toothed ends, the middle and inner rows both geniculate. No stout hooks at the
base of the peduncles but acicular setae may be present dorsally behind the external
row of paleae. A median cirrus may be present in front of the mouth. Three para-
thoracic segments with oar-shaped setae.
Type species; Sabella alveolata Linnaeus, 1767.
Key to Species
1 Inner paleae with serrated margins. (Outer paleae with tapered, denticulate ends) S.pectinata
- Inner paleae with smooth margins .......... 2
2 Middle paleae short and boat-shaped, never with a long spike. External paleae seldom
with a barbed central tooth in adults ........ S. intoshi
- Middle paleae with a long spike projecting from one end. External paleae always with
a long barbed central tooth [S. spinulosa subspp.) ....... 3
3 .Spike of middle paleae hooked and claw-like (fig. 33.1.0) . . S. spinulosa eupomatoides
- Spike of middle paleae straight or only slightly curved (fig. 33. 2. b) .... 4
4 Inner paleae spoon-shaped, .'\cicular setae serrate ..... S. s./ucicola
- Inner paleae produced at one end (fig. 33. 2. c). Acicular setae smooth . S. s. alcocki
Sabellaria pectinata Fauvel, 1928
(fig- 33-3-i-k)
.Sabellaria pectinata Fauvel 1928: 163, fig. 3a-g; Fauvel, 1932: 210; Fauvel, 1953: 396, fig. 206 a-g.
Body up to 8 mm. long, pale in alcohol. Tube fragile. Outer paleae (fig. 33. 3. i) num-
ber about 15-20. They are elongate with a series of about 12 teeth which increase in
size from small denticles midway along the blade to long sharp spines at the distal
end. Central spine nor markedly longer than the rest. Middle paleae (fig. 33.3.J)
much smaller than the outer ones, asymmetrically spoon-shaped with the tips pro-
duced. The surface is covered with a series of scaly ridges which form cusps at the
edges except on the asymmetrical flange which overlaps the next palea in the row.
Innermost paleae (fig. 33. 3. k) essentially similar to the middle row but the tips are
longer, sharper and directed inwards. In this case the ridges form spinules towards
the tip. Two to three smooth spines in the dorsal groove between the opercular
peduncles. A median cirriform lobe. Buccal tentacles remarkably long and stiff.
Type loc.\lity : Gulf of Mannar, India.
Records: Natal (29/3i;s).
Distribution: Tropical Indian Ocean.
G7'J POI.VCIIAETA OF SOUTHERN' AFRICA
Sabellaria intoshi Fau\cl, 1914
(I'lK- ;53-i-ti-'»)
Sabellaria spimihna var. inloihi Fauvcl, 1914: 139. pi. 8 figs. 50-53.
Sabellaria guinmsis Augener, igi8: 498, te.xt-fig. O2.
A large species reaching 60 mm. Tubes very hard, often forming gregarious
masses. Outer paleae (fig. 33.1.J) end in fi\c unequal teetli of which the median is
longer, usually smooth but in a few dorsal palac it may be barbed in juveniles.
Middle paleae (fig. 33.1.!;, 1) short, broad and asymmetrically boat-shaped, over-
lapping one another laterally. Inner paleae (fig. 33.i.m) longer and tapered but
essentially similar to the middle ones. Two to three smooth acicular setae in the
dorsal groove between the peduncles. A short median cirriform lobe ventrally
between the opercular lobes followed by a ridge and a pair of slender grooxed palps
in front of the mouth. About 12 rows of numerous buccal cirri (fig. 33.1.1). Abdo-
minal imcini with two rows of seven teeth.
Type LOf.Ai.iTV : Sao-Thome, Gulf of Guinea.
Records; Natal (29/31,1,5); Mocambique (26/32/i).
Distribution : Eastern Atlantic from the English Channel to tropical western
Africa, Angola (i).
Sabellaria spiniilosa fucicola (Augener, 1918)
(fig. 33.2.a)
Sabellaria fucicola Augener, 1918 : 496, pi. 6 fig. 158, text-fig. 81.
Sabellaria spinulosa fucicola: Day, 1961 : 525.
A small gregarious species about 5 lum. long with the tubes attached to algae.
Outer paleae broad and end in five to eight teeth of which the median is long and
barbed. Middle paleae (fig. 33. 2. a) with the base produced on one side as a long
erect spike which is almost straight. Inner paleae pointed with spoon-shaped ends.
Three to four fine spinulosc setae in the shallow dorsal cleft between the opercular
peduncles.
Type loc.\lit\'; South \Vest Africa.
Records: South ^Vest Africa (26/15/i, s).
Distribution : No other record.
Sabellaria spinulosa alcocki Gravicr, 1906*
(fig. 33.2.b-c)
Sabellaria alcocki Gravier, igoGd : 298, pi. 8 figs. 1 1-23.
A small gregarious species up to 20 mm. long with about thirty setigerous seg-
ments. Opercular crown almost at right angles to body with a deep dorsal notch
between the opercular lobes. About fifteen external paleae to each lobe, each broad
* According to Ilartman (1044), Gravicr's species is close to, if not identical with i'. bella Griibc 1870.
SABELLARIIDAE 673
and ending in five to seven teeth of which the median is always elongated and
barbed. Middle paleae (fig. 33. 2. b) about twelve teeth on each side, each with a
cup-shaped base with the outer edge produced into a long, erect, almost straight
spine. Inner row of about twelve paleae (fig. 33. 2. c) generally similar to those of
the middle row but with much shorter spines directed towards the centre. Numerous
buccal cirri arranged in about eight rows. A median cirrus and a pair of grooved
palps only shghtly larger than the buccal cirri are situated on the roof of the oper-
cular cleft in front of the mouth.
Type locality: Indian Ocean, 8°23N/76°28E, i86 metres.
Records: South ^Vest Africa (26/15/s); Cape (?32/i8/s and 34/22/s, 34/24/i to
33/27/s); Natal (30/30/i, s and 29/3 i/s, d).
Distribution : Eastern Atlantic from the English Channel (i) to Africa (Senegal
(s) ); Persian Gulf and tropical Indian Ocean (i,s, d) ; Indo-China; California.
GUNNAREA Johansson, 1927
Opercular peduncles completely fused dorsally and anteriorly. Opercular crown
formed of two rows of paleae set at right angles to body. No dorsal hooks or acicular
setae between opercular peduncles. A pair of grooved palps but no free median
cirrus, only a ridge on the roof of the opercular cleft. Buccal cirri arranged in num-
erous rows. Three parathoracic segments bearing large, oar-shaped setae.
Type species : Ilermella capensis Schmarda, 1861.
Gunnarea capensis (Schmarda, 1861)
(fig. 33.2.d-i)
Hcrmella capensis Schmarda, 1861 : 23, pi. 23 fig. 171.
Sabellaria capensis : Mcintosh, 1885 : 418, pi. 25A figs. 24-25, pi. 26A figs. 1 1-12.
A large gregarious species forming massive reefs (fig. 33. 2. d) between tide marks.
Operculum (fig. 33. 2. f) with an external row of twenty to twenty-five golden paleae
on each side, each palea (fig. 33. 2. h) with a broad blade set at right angles to the shaft ;
the edges are smooth and the distal end of the dorsal series are rounded and formed
of two broad flattened incurving teeth; the ventral series has simple tips. Inner
row with fifteen to twenty wedge-shaped paleae (fig. 33. 2. g) completely covering
the truncate ends of the united opercular peduncles. Ventral cleft between opercular
peduncles (fig. 33. 2. e) with fifteen to twenty rows of buccal cirri and a pair of small
grooved palps on the roof of the cleft immediately in front of the mouth. Median
cirrus reduced to a ridge. Abdomen of 45 segments. Abdominal uncini with six teeth
per row (fig. 33.2.1).
Type locality: Table Bay, South Africa.
Records: South \Vest Africa (from 22/141 to 26/15'i); Cape (from 29,16,! to
32/28/i); Natal (from 31/29/i to 28/32/i) - Abundant.
Distribution: Endemic.
'■>7+
poiA t.iiAi:! A oi- SOL riii.R.N Ai rr;a
,-. - f>
I'lr;. 'i'i.C!. Salifllinia ifiirmloifi Jncicota. (A) Middle palea. Sahctlaria spiriiilota ahocki. (b)
Middle palea. (cj Inner palea. Gimnarea capemis. (d) A few sand tubes from reef.
(f. i \'entral view of head with buccal cirri cut short on one side, (f) Dorsal view of operculum.
Cr;) Lateral view of inner palea. (h) Plan view of outer palea. (i) Uncinus. Idanlhr'uis
jimnatui. (jj Dorsal view of anterior end. (k; Outer palea. (i.) Inner palea. (m) Opercu-
lar hook. (N) Uncinus.
SABEI.LARIIDAE 675
IDANTHYRSUS Kinberg, 1867
(including CRYPTOPOMATUS Gravier, 1909)
Opercular peduncles elongated but not completely fused ; opercular crown oval
and often slanting in relation to the body. A pair of stout hooks at the dorsal
junction of the peduncles. Opercular crown consisting of two rows of golden paleae
which do not cover the ends of the peduncles. External row of paleae long and bear
lateral tcedi. Opercular cleft with numerous rows of buccal cirri on the sides and
a pair of grooved palps plus a median cirrus on the roof in front of the moutJi. Three
parathoracic segments bearing large, oar-shaped setae.
Type species: Idanthyrsus armatus Kinberg, 1967.
Key to Species
I External paleae plumose with curved shafts and slender lateral denticles . /. pennatus
- External paleae as serrated spikes with straight shafts and tapered denticles . /. armatus*
Idanthyrsus pennatus (Peters, 1855)
(fig- 33-2-J-n)
Sabellaria (Paltasia) pennata Peters, 1855: 613.
Idanthyrsus pennatus : Johansson, 1927: 88.
Opercular crown (fig. 33.2.J) with two clearly separated rows of long paleae.
External paleae (fig. 33.2.!:) with curved shafts and slender denticles giving the
general impression of a feather or a palm leaf. Inner paleae (fig. 33. 2. 1) smooth
with tapering tips. One to two pairs of stout dorsal hooks (fig. 33. 2. m) at the bases
of the opercular peduncles. About 15 rows of buccal cirri on the sides of the opercular
cleft. A pair of grooved palps and a small median cirrus continued as a ridge on
the roof of the opercular cleft. Three parathoracic segments with narrow oar-like
setae with frayed tips. Uncini (fig. 33. 2. n) with seven teeth. A large species reaching
50 mm. which makes solitary tubes or small reefs between tide marks.
Type locality : Mocambique Is.
Records: Natal from 30/30/ to 27/32/i) ; Mocambique (26/32/1), Mocambique
Is. (i).
Distribution : Ascension Is. (i) ; Tropical Indo-west-Pacific from India (i) and
Madagascar (i) to New Caledonia (i) and Japan (i).
LYGDAMIS Kinherg, 1867
Opercular peduncles elongate and not fused. Opercular crown long and slanting
with respect to the body ; it is formed of two rows of paleae which are rather small
so that the fleshy ends of the opercular peduncles are exposed. A pair of stout dorsal
hooks at the base of the opercular peduncles. Inner margins of opercular peduncles
with several rows of buccal cirri. A median cirrus and two large grooved palps in
front of mouth. Four parathoracic segments bearing oar-shaped setae.
Ty-pe species : Lygdamis indicus Kinberg, 1867.
67'3
POLVCHAETA OF SOUTHERX AFRICA
db/
Fic. 33.3. L]\i^(/(iitiis miirata lar. iiUcfuhti (after Mcintosh, iO'-4)- (a) Outer palea. (b) Inner
palea. Lygdamis indkus. (c) Lateral view of anterior end. (u, eJ Edge-on view and profile
of uncinus. (f) \'cntral view of head with the palp and buccal cirri removed from the left
side, 'r.i Outer paiea. fuj Inner palea. Sahrllaria pechnata. (i) Outer palea. (j) Middle
palea. (k) Inner palea.
SABELLARIIDAE 677
Key to Species
I Inner paleae ending in pointed tips (fig. 33. 3. b) . . . . L. miirata gilchristi
- Inner paleae stout with blunt tips (fig. 33. 3. h) . . . . . . . /.. indicus
Lygdaniis murata gilchristi (Mcintosh, 1925)
(fig- 33-3-a-b)
Tetreres murala var. gilchristi Mcintosh, 1925 : 76, pi. 10 figs. 10-12.
Operculum a long oval formed of outer and inner rows of paleae mainly imbedded
in the flesh. Outer paleae (fig. SS-S-a) slightly tapered and end in blunt tips. Inner
row (fig. 33. 3. b) shorter, swollen in the middle and end abruptly in pointed tips.
Four parathoracic segments with oar-shaped setae. Abdominal uncini with eight
teeth. Tube broad and composed of sand grains and foraminifcran shells. A deep
water species.
Type LOCALITY : South Africa, 33°25S/i7°24E in 476 metres.
Records: Cape (33/i7;'d).
Distribution : A single record only.
Lygdamis indicus Kinberg, 1867
(fig. 33-3-c-h)
Lygdamis indicus Kinberg, 1867 : 350 ; Johansson, 1925 : 8, fig. 2, 2-7.
Operculum (fig. 33. 3. c) oval and slanting. External paleae (fig. 33. 3. g) about 25
on each side ; each smooth, straight and tapered to the recur\ed tip ; margins finely
striate in juveniles. Inner paleae (fig. 33.3.11) about 16 on each side; each much
stouter with a blunt end. A tapered median cirrus and a pair of large grooved palps
in front of the moudr (fig. 33.3 f). About 8-10 rows of slender buccal cirri on the
inner sides of the opercular peduncles. Abdominal uncini (fig. 33. 3. d, e) with two
rows of eight to nine teedi each.
Type locality : Bangka Straits, East Indies.
Records: Natal (30/30/i, s).
Distribution : Tropical Indo-west-Pacific.
6-ii P01.^ CIIAK TA or SOUTllKRX AIRICA
Family PECTINARIIDAE Quatrcfagcs, i86r,
'l'ul)ii (ilous worms ciu ascil in In-c, tapcnd, s.iiidy tubes open at botii ends. Body
slioit wiih ail aiitiiior opi iculuni and three distinct regions. Prostomiuni indistinct.
'J'lie buccal scgmenl ibrnis a broad opeicular plate on the dorsal siuface of tlic head
and bears a Ian of stout paleae anteriorly. A cephalic veil derived from the pro-
stomiuni liirnis a hood in Iront ol'the mouth wliic h is sinrounded by numerous buccal
tentacles. Two pairs of tentai ular cirri, the first on the buccal segment and the
second on segment .! which is very short. Segments ■] and 4 usually bear lamellaled
gills. Segments 5-7 short and uniramous with notopodial cajjillarics only. The
next 12-15 segments arc biramous with geniculate notosetae dorsally and rows of
uncini \entrally. TJie last few (? 5) segments form a flattened plate or scaphc with
liociks at its base.
Records from southern Africa
I'fclinntia (Ainjihict(iie) cdjien^is (Pallas) . . GCli, igCi. .) [.Cli,
51CS, 56\Vs
as Nereii criindiaria capensis Pallas, 1778
as Sabella capensis Linnaeus, 1 788
as Sabella chrysodon Linnaeus, i 788
as Sabella indica Linnaeus, i 788
Pectinaria {Pectinaria) papilloma ClauUery . . 4oPi
Pectinaria {Lagis) koreni Malmgren . . .jSW'sd
Pectinaria {Lagis) koreni cirrata Day . . .'jGXs
Pectinaria (Lagis) neapulitana Claparede . . fiiC's
as Pectinaria pseiidoknreni Day . . . 44Ca, 45Ni
RIOLOGKIAL NOI l.S
Pectinaria is a deposit feeder and constructs a chimney-shaped tube most of which
is buried in the sand. The tube is r|uite free and may be constructed of sand grains,
sponge spictiles, foraminiferan sliells or ex-en shell fr.igments. It is cylindrical and
often fiintly curved like a hollow lusk. Onh the open li]i of the tube projects aho\e
the surface of the sand and the respiratory current is drawn down the tube, o\er the
tail of the worm and along the body to the gills. The liead is provided with a thick
fleshy operculum and a row of flattened setae (paleae) which are used for bui row-
ing. Behind these is the mouth surrounded by ninnero\is grooved buccal tentat les
which gather food particles buiied in the sand and help in the cfmstruclion ol the
tube.
'II IE MAIN DIAGXOSTIC: ClHARACilERS
Reviews of the faniih will be found in Hi'ssle (nji 7), Fainel (i()2 7), Nilsson (11)28)
and Hartman (1941).
.•\s noted by Fativcl, the fusion of the anterior segments makes it diflicult to deter-
mine their homologies and the first uncigerfius segment has been ix'ckoned as
the 7th, 81I1 or c)th b\- diH'eieiit workers. The \ lew adojited here is that the
PECTINARIIDAE 679
first uncigerous segment is the 8th but luckily the exaet number of anterior
segments is not significant fi-om a taxonomic point of view. The most useful char-
acters are the opercular margin, the cephalic veil, the number of uncigerous
abdominal segments, the number of tooth rows on the uncini, the separation of the
scaphc from the rest of the body, and, on the species level, the nature of the tube.
The operculum is a stout muscular plate which forms the antero-dorsal surface of
the head. It is surrounded dorsally and laterally by a raised opercular margin or rim
which may be smoothly continuous or divided into a number of triangular lappets or
"teeth". Vcntro-laterally the operculum bears the first pair of tentacular cirri and
ventrally it gives rise to a series of 20-30 golden paleae by means of which the worm
burrows in the sand. The paleae are uniform in shape and their number varies with
the size of the worm. They are of little taxonomic importance.
The cephalic veil ( = antennular membrane of Hartman) is a curved membrane
which bears eyes in juveniles and lies immediately behind the paleae and protects
and encloses the grooved buccal tentacles. In the genus Petta the cephalic veil has a
smooth margin but in Pectinaria the margin bears numerous long papillae and the
veil is said to he fringed. The relations between the veil and the operculum are well
shown by Hessle. In Pectinaria auricoma the veil is completely free from the operculum
and forms a ventral semicircle around the front and sides of the numerous buccal
tentacles. In Pectinaria koreni the veil is merely an anterior arch whose ends are
fused to the operculum at the bases of the tentacular cirri. In Pectinaria neapolitana
there is an intermediate condition ; the veil is a semicircle extending around the
sides of the buccal tentacles as in P. auricoma but there are low bridges attaching the
veil to the bases of the tentacular cirri. Such a veil is said to be partially fused to the
operculum.
A low ventral ridge unites the first pair of tentacular cirri behind the mouth.
The second pair of tentacular cirri are united by a more distinct ventral ridge
usually incised to form glandular lobes but occasionally produced to form triangular
papillae. Two achactous segments follow with lamellate branchiae laterally and
glandular pads ventrally, then the first three setigers with capillary notosetae and
glandular ventral pads but no ncurosetae. This marks the end of the thorax.
The abdomen consists of 12 or 13 segments provided with notopodial capillaries and
neuropodial uncini and behind these, 0-2 segments which lack uncini but may possess
small bundles of notopodial capillaries. The capillaries are winged, some are said
to have smooth tips and others spinulose tips, but probably all are spinulose under
high magnification ; they arc not of systematic importance. The uncini on the other
hand provide valuable characters. They are provided with both major and minor
teeth abo\e a horseshoe-shaped gouge. The major teeth may be arranged in one,
two or even three to four vertical rows and Hartman has separated Cistenides with
one row of teeth from Pectinaria with two or more rows of teeth. These are regarded
here as subgenera. Unfortunately the number of teeth is difficult to determine
unless the uncini are viewed edge -on.
URo I'OLVCHAKTA OF SOL'THIiRX AIRICA
The scaphe is a sliort foliaccous caudal region which is large and well dclined in
Pectinaria but small and poorly defined in Ptlla. It bears a number of short, acicular
scaphal hooks where it joins the abdomen and an anal Ugule with an anal cirrus termin-
ally. The number of scaphal hooks is somewhat variable within an individual
species, e.g. 3-4; 6-10; 13-15 on each side but the shape and ornamentation of
the scaphe is of some value.
The tube is always conical and open at both ends. In some species it is almost
straight and in others it is obviously curved. .Some species use irregular sized sand
grains, others small uniform sand grains and some even select short fragments of
sponge-spicules which are arranged like bonded bricks in the construction ol the
tube.
Key to Genera
I M.Trgiii of ceph.-)Iic veil smooth. Scaphe indistinctly separated from ihe abdomen. PETTA*
- Margin of cephalic veil with cirriform projections. Scaphe distinctly separated from
abdomen PECTINARIA
PECTINARIA Savigny, 1818
Head with an opercular plate having a smooth or dentate margin and a row of
paleae ventrally. Cephalic veil with marginal cirri and may be free from or united
to the opercular plate. Two pairs of tentacular cirri and numerous buccal tentacles.
Usually two pairs of lamellate gills. Neuropodial uncini on 12-13 segments starting
from setiger 4. Notosetae are winged capillaries with denticulate tips. Uncini
scrpuliform with a gouge and numerous teeth. Scaphe separate from the abdomen
and lacks eyes.
Type species : Nereis cylindmria hclgica Pallas, 1 766.
Key to Scbgenera and Species
1 Cephalic veil partly or entirely fused to the operculum. (Operctil.ir rim smootli. Twehe
segments with uncini. (.Subgenus LAGIS)) ........ 2
- Cephalic veil free from the operculum (fig, 34.2.6) ....... 4
2 Cephalic veil partly fused to the operculum {lig. 34.1.C) ... P. iL.) neapolitana
- Cephalic veil entirely fused to the operculum (fig. 34. 1. h) ...... 3
3 \'entral flange joining second pair of tentacular cirri with f'.-io blunt lobes
P. (Z..I koreni koreni
- X'entral flange joining second pair of tentacular cirii with 15-20 long papillae
/*. (Z..) koreni cirrata
.\ Opercular rim dentate (.Subgenus AAJPHICTENE ). Thirteen segincnts with imcini.
Tube made of si^ongc spicules (lig. 34.2.b^j . . . . . . P, (/I.) capensis
- Opercular rini smooth ............ ;,
") L'ncini with major teeth in a single row (Subgenus CISTENIDES\ . (.\o .South .African spp.)
- l'ncini with major teeth in two or more rows (Subgenus PECTINARIA). (Thirteen
segments with uncini. .Scaphe papillose) . . . . . . P. (P.) papillosa
PECTINARIIDAE 681
Pectinaria (Lagis) neapolitana Claparede, 1870
(fig. 34.i.a-f)
Peclinaria ?ieapolilana Claparide, 1870: 123.
Pectinaria pseudokoreni Day, 1955 : 432, fig. 5 a-c.
Body (fig. 34. 1. a) tapered and up to 25 mm. long. Cephalic rim smooth. Veil
(fig. 34.1.0) partly Rased to operculum being attached by a low fold to the base of
the first tentacular cirrus and also continuing posteriorly along the sides of the buccal
cirri, .\bout 12 velar cirri. Posterior to the mouth the ventral surface of the second
tentacular segment forms a transverse ridge with about 10 blunt triangular lobes.
Three anterior segments bearing notosetae only. Twelve abdominal segments with
both notosetae and unciiii followed by two achactous prescaphal segments. Five
scaphal hooks. Scaphe (fig. 34.i.e) oval with a lobed margin. Anal ligule with a
small anal papilla. Uncini (fig. 34.i.f) with two to three rows each with about eight
major teeth and three to four minor teeth preceding the basal gouge. Notosetae
mainly with denticulate tips (fig. 34.i.d). Tube composed of coarse sand grains
irregularly arranged (fig. 34.i.b).
Type locality : Naples.
Records: South West Africa (26/14/d and 26/15/s) ; Cape (from 33/18/s to
34/23/i, s and 33/25/e, s).
Distribution : Mediterranean.
Pectinaria (Lagis) koreni koreni Malmgren, 1 865
(fig. 34-i-g-h)
ifljgiV A'oreni Malmgren, 1865: 360.
Pectinaria (Lagis) koreni: Fauvel, 1927: 221, fig. 77 a-i.
Cephalic rim smooth. Ten to fifteen golden paleae on each side. Cephahc veil
(fig. 34. 1. h) completely fused to the operculum and does not continue beyond the
point where its edges are fused to the bases of the first tentacular cirri. About 20
velar cirri. Posterior to the mouth the vcntrum of the segment with the second pair
of tentacular cirri bears eight to ten blunt triangular lobes. Three anterior segments
with notosetae only. Twelve abdominal segments with notopodial capillaries and
uncini followed by two achaetous prescaphal segments. Notosetae include smooth-
winged capillaries and others with saw-edged tips. Uncini (fig. 34. i .g) pectiniform
with three to four rows each with six to eight major teeth and four indistinct minor
ones preceding the basal gouge. Three pairs of scaphal hooks. Scaphe oval with five
lateral scaUops on each side of which the first three bear papillae. Anal hgule
broader than long with a small anal cirrus. Tube almost straight and composed of
sand grains of varying size.
Type locality : Stavanger, Norway.
Records: South West Africa (23/14/d).
Distribution : Eastern Atlantic from Scotland (s) and Norway to Morocco (s)
and tropical western Africa (s) ; Mediterranean.
682
poi,Yc;n.\i;TA of southerx airic.a
Fi(^ 34-I- Pnliiiarii! (Lagi^) nra/mlilana. (a) Entire worm (three times natural size). IB! Part
of tube, (c) \'entral view of head, (d, d'i \otopodial capillary and details of tip. (e)
Scaphe. (f) Unrinus. Pectinatia (Lagii) koreni kureni (after Fauvel). (c) Unc inus. (h)
N'entral \"iew {if head. Pi-ilinaria {Lagis) korrrii cirrala. (i) Unc inus. (j) X'entral \ iew of
head.
Pectinaria [Lagis] koreni cirrata Day, 1963
(fig. 34.i.i-j)
Pettinarin il.agh) korcru c'nrnta Day. iriG^a : 434. fitr. 11 a.
A small species about 10 mm. long. 10 -\- ro palcac. (3piinilar lim high and
smooth. \'eil completely fused to the operculum and ending whefe its sides arc
fused to the bases of the first pair tentacular cirri. Fourteen velar cirri. First pair of
tentacular cirri slender and longer than the paleae ; second pair shorter. Three
PECTINARIIDAE 683
small lobes inside the \eil and just mcdiad to tentacular cirrus i. Posterior to the
mouth the ventrum of the segment bearing the second pair of tentacular cirri forms
a ridge bearing 15-20 slender cirri (fig. 34.1.J). Second pair of gills smaUer than the
first. Three anterior segments with notosetae only. Twelve abdominal segments
with both notosetae and uncini followed by two achaetous prescaphal segments.
Uncini (fig. 34.1.1) with two rows each with six to seven major teeth and several minor
ones above the gouge. Scaphal hooks four to six on each side. Scaphe oval with a
flanged margin incised to form a median and five to six lateral lappets. Anal ligule
tongue-shaped widi a small anal cirrus. Tube composed of coarse sand grains.
Type locality : Dredged off Natal, South Africa.
Records: Natal (29/32/s).
Distribution : A single record.
Pectinaria [Amphictene) capensis (Pallas, 1776)
(fig. 34.2.a-d)
Xereis cylindraria capensis Pallas, 1776 : 1 18, pi. 9 figs. 1-2.
Pectinaria {Amphictcne) capensis Mcintosh, 1904 : 76, pi. 7 figs. 35-36 ; Day, 1955 : 432 ; Da\', 1963a ;
433-
A large species reaching 90 mm. (fig. 34. 2. a). Cephahc rim (fig. 34. 2. c) with about
20 serrations. Eleven to fifteen golden paleae on each side. Cephahc veil joined to
the operculum medially but quite free from the bases of the first pair of tentacular
cirri. Two clavate papillae and a conical lobe between the sides of the veil and the
base of the first tentacular cirrus. Twenty to thirty velar cirri. Both pairs of tenta-
cular cirri shorter than the paleae. Ventrum of the segment bearing the second pair
of tentacular cirri incised to form about 12 square lappets. Lateral swellings dorsal
to the origin of the second tentacular cirrus and first pair of gills. Three anterior
segments with notosetae only. Thirteen abdominal segments with both notosetae
and uncini followed by one achaetous prescaphal segment. Seven to eight scaphal
hooks on each side. Scaphe oval with two pairs of marginal cirri proximally and one
pair distally. Anal hgule shield-shaped with a slender anal cirrus often detached.
Notosetae include a row of broad, smooth-winged capillaries and a row of capillaries
with a basal spur and a spinulose blade. Uncini (fig. 34. 2. d) with two rows of six to
eight major teeth and a crowded group of four rows of about eight minor teeth pre-
ceeding the basal gouge. Gouge large and horseshoe-shaped in plan. Tube (fig.
34. 2. b, b') straight and composed of short lengths of sponge spicules arranged like rows
of bricks or occasionally of small, even-sized sand grains regularly arranged in rows.
Type locality : Cape of Good Hope.
Records: South-West Africa (26/i5;'s and 28;i6/s) ; Cape (from 32/i8;s and
33/18/i, s to 34/18/s).
DiSTRiBLTio.v : ? Red Sea.
684
I'OLVCHAETA Ol- SOUTHERN AFRICA
Fig. :^4.'2. IWlinana {Amphiclcne) aifxmis. l.\) Entire worm (twice life size), fn. h^) Tube wllli
part magnilied to show details, (c:) Dorsal view of head, (d) Uncinus. Pniitnuia {Pccliiiaria)
piif'ili'iui. /Kj Wntral view of head with the veil fcilded down over liie mouth, (f) Dorsal
\ iew of head, (c) Uncinus.
PECTINARIIDAE 685
Pectinaria [Pectinaria) papillosa CauUery, 1944*
(fig. 34.2.e-g)
Pectinaria papulosa Caullery, 1944: 71 ; Day, 1951 : 55, fig. 8 a.
A large stout species reaching a length of 60 mm. and 16 mm. across the operculum.
Rim of cephalic plate smooth (fig. 34.2.f); 13+13 golden paleae. Cephalic veil
(fig. 34. 2. c) joined to the operculum medially but quite free from the bases of the
tentacular cirri laterally. About 46 velar cirri. Both pairs of tentacular cirri smaU,
the second pair being lateral in origin and mounted on a flange which extends from
the dorso-lateral surface across the ventrum immediately behind the mouth. The
ventral part of the flange has a smooth margin. The second setiger also has a
prominent ventral ridge. Three anterior segments with notosetae only. Thirteen
abdominal segments with both notosetae and uncini followed by one prescaphal
segment with only a small bundle of notosetae. Three to four strongly curved
scaphal hooks on each side. Scaphe oval, edges scalloped, surface papillose. Anal
ligule cordate. No anal cirrus (? lost). Notosetae stout with narrow hispid wings
and spinulose tips. Uncini (fig. 34. 2. g) with two rows of about seven major teeth
followed by two to three rows each with three minor teeth preceding the basal
gouge. Tube composed of coarse, irregularly arranged sand grains.
Type locality : East Indies.
Records: Mocambique (26/32/i).
Distribution : East Indies.
*\'ery close to P. antipoda Schmarda.
686 l'( iIA (. 11 \1 1 A (>1 SOL riir.RX Al RICA
Family AMPHARETIDAE Malnit;,-,-,!, iHGy
riil)icolous worms li\ing in fragile tubes of sandy mud. Bnd\ tapered and di\ided
into two regions. Prostomium \vell de\'elopcd, often with eyes but without sensory
appendages. Mouth with numerous buecal tentacles for deposit feeding. Pcri-
stomium and the next segment achactous and often partly fused. Segments III-\'I
primiti\ely bear segmentally arranged dorsal branchiae but these are often reduced
in number and concentrated in a transverse row across segment III. Similarly one
or more bundles of setae may be reduced or absent from segments IIl-VI. Segment
\'II and 10-13 succeeding thoracic segments bear both notopodial capillaries and
neuropodial uncini. Abdominal segments without milosriae but with imcini.
Pygidimn often with anal cirri.
BIOI.OWUAl. XOIES
The ampharetids are deposit feeders which gather food particles from the surface
of the sand or mud by means of buccal tentacles which can be extruded from the
mouth. They construct fragile tubes of mud or sometimes sand grains and attach
them to sponges, compound ascidians or the shells of li\-ing molluscs. They are
seldom found on stones and very few ampharetids li\e in shallow ^\ater but they
become common with increasing depth.
THE MAIN DIAGNOSTIC CHARAC IKRS
Re\'iews of the family and keys to the genera will Ik- found in Hessic (1917),
Chamberhn {1919), Fauvel (1927), Usehakov (1957) and Day (1964). It is im-
]3ortant to note that Hessle and Usehakov regard the paleal (or first branchiferous)
segment as segment II whereas the other workers c[uoted rc^gard the paleal segment
as segment III. This is also the \iew adopted in the present work.
The important taxonomic characters include the structure of the prostomium,
the nature of the buccal tentacles, the branchiae, the setation of segments III to \T,
the number of uncigerous thoracic segments, the nimilxr of abdominal segments
and the structure of their parapodia, the shape of the un< iiii and the development of
the anal cirri.
The lu'iid and biictal tentacles. The prmtomium (or tentacular membrane of Hessle)
is a flattened hood ri\ er the mouth and bears eyes, nuchal slits and sometimes a pair of
"landular ridges which di\erge outwaifls towards the antero-lateral margins. The
buccal tentacles arise from the upper lip and may be retracted into a special cavity
above the true mouth. The tentacles themselves arc papillose or smooth with a
groove along one side as in the Terebelhdae. In a few spec ies the tentacles are borne
by an exersible probosciform lobe or tongue (fig. ;',",. 2. a).
The hranchiae, nephridia and sclaliim ofsegmenti III-Vl. The lour brandii.d sigments
arc ofti-n ti-Iescoped or |5artially fused and some of the branchiae and bundles of setae
may be missing. Primitively there are four pairs of segmentally arranged gills as is
shown by the blood \-esscls Ijiit in man)' genera tlie gills are concentrated to form a
transx'erse row mounted on a branchial ridge across segment III. Fiuther the nimiber
AMPHARETIDAE 687
of paired gills may be reduced from four to three or even two. The individual gills
are usually smooth and tapering but in certain species they may bear rows of small
lamellae or papillae. The number of nephridia may be reduced in the same way as
the gills, and in Sabellides the enlarged anterior pair of nephridia open on a pair of
prominent nephridial papillae mounted on the branchial ridge between the right and
left groups of gills. In Anobothrus the two nephridia open on a single median papilla.
In the subfamily Melinninae the neurosetae of the bi anchial segments take the
form of minute acicular setae deeply embedded in a lateral fold which slants upwards
and backwards from below the mouth to the sides of segment \T. The two folds are
united by a dorsal crest across segment VI. In the subfamily Ampharetinae neuro-
setae are entirely lacking from the branchial segments and even the notoseate
may be reduced or lacking from some segments. Segment IV very often lacks noto-
setae and is fused to segment III. Segment III itself may lack notosetae and even
segment \'. On the other hand segment III may develop enlarged notosetae called
paleae which project forwards on either side of the gills.
The uncigerous thoracic segments. Segment \TI always bears both notopodial
capillaries and neuropodial uncini so that it is the most reliable landmark on the
body. The number of uncigerous thoracic segments is constant for each genus and
thus a most useful taxonomic character. The notopodia are usually simple conical
projections bearing bundles of winged capillaries, but in Amphicteis and certain other
genera each notopodium has a small terminal papilla or notopodial cirrus which is
quite distinctive. The neuropodia are small square uncigerous pinnules below the
notopodia. Like the notopodia they may also bear superior papillae though these
are seldom so obvious as those on the notopodia.
The abdomen and the anal cirri. Abdominal segments may be disdnguished from
thoracic ones by the lack of notosetae. In most species the endre notopodium is
lacking but in some it persists in a rudimentary form for a few segments or on all
segments to the pygidium. Occasionally the notopodial cirrus persists even though
the notopodium itself is lacking. The neuropodium is present as an uncigerous
pinnule on every abdominal segment ; in fact it tends to elongate so that the last few
uncigerous pinnules are the longest of the whole body. The same is true of the
superior papilla of the neuropodium which is usually minute on the thorax but may
become a long cirriform organ after the third abdominal segment. This is well
sho\vn by Sabellides octocirrata. It is important to distinguish this neuropodial cirrus
from the rudimentary notopodium above it by examining the parapodia at the junction
of the thorax and abdomen.
The number of abdominal segments varies from over 50 in many species of the
subfamily Melinninae to as few as 10 in some of the Ampharednae. The approxi-
mate number is of systematic value in the Ampharetinae. The pygidium may be
quite simple with only a few indistinct papillae around the anus or it may develop
long anal cirri. Commonly there is only one pair of these but two, three or even four
pairs may be present.
The setae. As noted, the notosetae of segment III may be enlarged to form con-
spicuous paleae or they may remain small or may be absent. \Vith few cxcepdons
fi88 POI.VCHAE TA OF SOUIHERX AIKICiA
llic notosctac of subscqiKiit ihoracir segments arc winged capillaries whicli are so
uniform in structure that they are of little systematic \'aluc. In several genera of the
Melinninae however, the notosetac of segment IV are modified to form a pair of
stout postbranchial hooks. Again in a few genera of the Ampharetinae one or more of
the posterior notopodia is elevated above the rest and the notosetac arc modilied -
usually the blades develop spinulcs.
The neurosetac of the branchial segments of the Melinninae are minute and acicu-
lar and deeply embedded in the flesh. In the Ampharetinae, neurosetac arc entirely
absent from segments III to VI. Subsequent thoracic segments all bear uncini.
These arc toothed plates which arc (juadrangular to triangular in outline with one,
two or more series of teeth above the small rostral point and basal prow. In many
species the rostral point is like a small tooth and may be mistaken for the lower-
most of the series but in species of .Meliiina it is obviously an attachment organ and
in species with two or more series of teeth the rostral point is always single and median.
The basal prow may project forwards as a continuation of the base or may be short
and curve upwards so that its blunt apex almost meets the rostral point. The number
of teeth above the rostral point is very constant in the thorax but usually increases
in the abdomen. There is a single vertical scries of teeth in the Melinninae and in
several genera of the Ampharetinae. Other genera have two or three series but
rarely more.
Key to Subfamilies
I Segments III-Vl (or III-V) with fine acicular neurosetae. Post-branchial hooks may
be present. No paleae ........ Melinninae (p. 68g)
- Segments III-VI without neurosetae. No post-braiuhial hooks. Paleae may be present
A.MPHARETINAE (p. Gg3)
Subfamily MELINNINAE Clhamberlin, igig
Buccal tentacles never papillose, usually smooth with a groove along one side.
Paleae absent. One or two pairs of stout notopodial hooks may be present behind the
gills. Small acicular neurosetae embedded in segments III-V and sometimes VI.
Uncini always with a single series of teeth. Numerous (20-go) abdominal segments.
Records from southern Africa
Isolda pulchella Muller ....... 56CS
holda ivhvdahensis Augener ...... 26Ai
Melinna cristata (Sars) ....... 48Cd
Melinnopsides capensis (Day) ......
as Mditinopsis capensis Day ...... 44^^'!
Key to Genera
1 Stout notopodial hooks behind the gills (lig. 35.i.i) ....... 2
- Notopodial hooks absent ........... 4
2 Four pairs of gills . ............ 3
- 'I'hrcc ]")airs of gills (one smootli, two pennate) ....... IRAN A*
AMPHARETIDAE 689
All gills smooth MELINNA (p. 689)
Some gills smooth, some pennate (fig. 35.i.k) ...... ISOLDA (p. 691)
Four pairs of smooth gills MELINNOPIS*
Three pairs of smooth gills MELINNOPSIDES (p. 692)
MELINNA Malmgren, 1866
Buccal tentacles smooth with a groove along one side. Four pairs of smooth gills.
A dorsal crest across segment VI. Segments III-VI with notopodial hooks on
segment IV and notopodial capillaries on segments V and VI. Fine acicular neuro-
setae on segments III-V and sometimes on VI as well. Fourteen uncigcrous thoracic
segments and 30-50 abdominal ones. Thoracic uncini with a single series of teeth.
Type species : Sabellides cristala Sars, 1 85 1 .
Key to SpEcrES
1 Numerous buccal tentacles (fig. 35.1. i). Membrane across segment V'l crenate . . 2
- One large buccal tentacle (fig. 35.1. a). Membrane across segment \T smooth
M. monoceroides
2 Neurosetae present on segment \'I. Branchial filaments united for less than half their
length ............ M. cristata
— Neurosetae absent from segment \'I. Branchial filaments united for more than half their
length ............ M. palmata*
Melinna monoceroides Fauvel, 1936
(fig- 35-i-a-b)
Melinna monoceroides Fauvel, 1936: 93, fig. 12 a-d.
Body up to 17 mm. long. Prostomium (fig. 35.1.3) trilobed, without eyes or
glandular ridges. A single greatly enlarged tentacle arising from the upper lip.
Branchiae in two groups of four, three of which are united for half their length and
arranged in a semicircle around the fourth. Individual gills long and smooth,
projecdng far beyond the prostomium. Segments III-V (and possibly VI) with fine
neuropodial acicula. Segment IV with stout notopodial hooks ; segments V and VI
with fine notopodial capillaries. Transverse dorsal ridge across segment VI forming
a deep pocket with a smooth margin. A total of 14 uncigcrous thoracic segments
starting from segment VII. Thoracic uncini (fig. 35.i.b) with a single series of
five to six teeth above the very small rostral point and basal prow. Number of
abdominal segments unknown. Abdominal neuropodia without superior cirri.
Pygidium unknown.
Type locality : Dredged in 224 m. off Morocco.
Records : Not recorded from southern .Africa.
Distribution : Morocco (d) ; Angola (s).
(><|0
roi,vc:iiAi:iA ui .soliiikkn airuia
I-"n;. 3'|.i. .MiCninti minw(iroidc\ (niodificd from I'.uivcl, K)'i'j). (a) Anlcrior cikI. (b) 'I'horaric
iiiu inus. Mtlinna crlstala. (r.) Post-hraiichial liouk. (D, e) Edge-on and prolilc (A ihoratic
iini iitus. (I) Xcuropodial acicular seta. {<•) Winged imtopodial capillary, (n) Entire
worjn Mhn-c times life si/e). (i) Anterior end. (j) Abdominal parapodium. holda
j'uhhiUa. (K) Anterior end witli tentacles fully extruded fgills on right side omitted).
(l) AI)d'iniinal segment showing parapodium. (m, n) Profile and edge-on view of thoracic
Lincinns.
AMPHARETIDAE 691
Melinna cristata (Sars, 1851)
(fig- 35-i-c-j)
Sabellides cristata Sars, 1851 : 205.
Alelinna cristata : Hessle, 1917: 92; Fauvel, 1927: 237, fig. 83 i-n.
Body (fig. 35. 1. h) tapered, up to 50 mm. long. Prostomium with eye-spots.
Buccal tentacles smooth with a groove along one side (fig. 35.i.i). Branchiae in two
separate groups of four each, united by a web for less than half their length. Segments
III-VI all with fine neuropodial acicula (fig. 35.i.f) embedded in the flesh. Segment
IV with stout notopodial hooks (fig. 35.!.c) ; both segments V and VI with noto-
podial capillaries (fig. 35.i.g). A total of 14 uncigerous thoracic segments starting
from segment \'II. Transverse dorsal ridge across segment VI with a dentate
margin. Thoracic uncini (fig. 35. 1. d) with a single series of three to four teeth above
the small rostral point and a basal prow and sometimes with a crest of three to
five teeth (fig. 35.i.d). Thirty to fifty abdominal segments with uncigerous pinnules
and small rudimentary notopodia (fig. 35.1.J). No anal cirri.
Type locality : 550 metres off Norway.
Records: Cape (32/1 7/d).
Distribution : Arctic ; North Atlantic from Greenland (s, d, vd) and Norway
(vd) to the EngUsh Channel (e, s) and North CaroUna (s) ; boreal North Pacific
from Alaska to N.W. Japan ; subantarctic (d).
ISOLD A MuUer, 1858
Buccal tentacles smooth with a groove along one side. Four pairs of giUs of which
two are smooth and two are pennate. A dorsal crest across segment VI. Segments
III-VI with notopodial hooks on segment IV and notopodial capillaries on segments
V and \'I. Fine acicular ncurosetac on segments III-V and sometimes on VI as
well. Twelve to thirteen uncigerous thoracic segments and 25-36 abdominal ones.
Thoracic uncini with a single series of teeth.
Type SPECIES : Isolda pulckella MiiWer, 1858.
Key to Species
I Thirteen uncigerous thoracic segments. Inner two pairs of gills with two rows of long
lamellae (fig. 35. 1. k) .......... I. pulchella
- 'I'wehe imrigerous thoracic segments. Inner two pairs of gills with many minute lamellae
(fig. 35.2.3) . . . . . . . . /. zvhydahaeitsis
Isolda pulchella MiiUer, 1858
(fig- 35-i-k-n)
Isolda pulchella Miiller, 1858: 219; .^ugener, 1918: 517, pi. 7 fig. 229, text-fig. 88; Day, 1963a:
434-
Body up to 45 mm. long, brown in alcohol with barred tentacles. Prostomium
snout-like. Eye-spots minute. Buccal tentacles smooth with a groove along one side
(fig. 35. 1. k). They arise from a horizontal shelf in the roof of the mouth. Stout
fio:: POI.VC.HAETA OF SOUTHERN AFRICA
lateral folds embrace the branehiferous region. Trans\eise ridge across segment VI
with a smooth margin. IJrauchiac in two groups of four, each united basally ; inner
two pairs of gills \vith l\\(i rows of long lateral lamellae. Fine acicular ncurosctae on
segments III-\' but not \'I. Stout noto]5odial hooks on segment IV. Small noto-
podial capillaries on segments V and \'I. A total of 13 uncigerous thoracic segments
starting from segment VII. Thoracic uncini (fig. 35.i.m, n) with a single series of
five to six teeth above the small rostral point and projecting basal prow. Abdomen
with 32 or more segments each with a square luicigerous pinnule above which is a
small papilla (fig. 35.1.I).
Type i,o(_:ahty : St. Catherine Is., Brazil.
Reccirds : Cape (34,'i;3/s).
Distribution ; North Carolina (s) ; Brazil ; S.W. Australia (s) ; Burma.
Isolda Tivhydahaensis Augener, 1918
(fig. 35---^-ci)
hoUa whrdahacnsis Augener, igi8: 514, pi. 7 lig. 216, text-fig. 87.
A small species about 5 mm. long. Head (fig. 35. 2. a) snout-like with a triangular
prostomium. Eyes not seen. Tentacles short, grooved and mounted on a protrusible
"tongue" arising from the roof of the mouth. A pair of lateral folds embrace the
branehiferous region. Dorsal ridge across segment VI with a smooth margin.
Branchiae long and slender, arranged in two groups of four which arc united basally.
Inner two pairs of branchiae stouter than the two outer and beset with numerous
minute lamellae (fig. 35. 2. c). Fine acicular ncurosctae embedded in segments
III-V but not VI. Segment IV with stout notopodial hooks, segments V and VI
with notopodial capillaries. A total of 12 uncigerous thoracic segments starting
from segment VII. Thoracic uncini (fig. 35. 2. b) with a single series of fi\e to six
equal teeth above the small rostral point and projecting prow. Abdominal uncini
usually with two rows of teeth. Abdominal segments numerous and bear square
uncigerous pinnules (fig. 35. 2. d).
Type uot;ALiTY : Whyda, tropical western Africa.
Records : Not recorded from South Africa.
Distribution : Western Africa from Morocco (s) to Angola (i, s, d).
MELINNOPSIDES Y><\) , 1964
Buccal tentacles smooth with a groove along one side. Three pairs of smooth gills.
Xo dorsal crest on segment VI. Segments III-VI without notopodial hooks on
segment I\' but with notopodial capillaries on segments V and \\. Fine acicul.ir
neurosetae on segments III-V but not VI. Ten luicigcrous thoracic segments and
about 30 abdominal ones. Thoracic imcini \\ illi a single series of teetlu
Type speues : Mi-liniinjisii capensis Day, 1955.
AMPHARETIDAE
693
Meliiinopsides capensis (Day, 1955)
(fig. 35.2.c-f)
Melinnopsis capensis Day, 1955 : 433, fig- 5 d-g.
A small species about 6 mm. long. The worms are gregarious and live in sandy
tubes attached to corallines. Prostomium (fig. 35.2.f) arched with a rounded anterior
margin. Two to three pairs of eyes. Three pairs of smooth cirriform gills all united
basally to form a continuous row of six across segment III. Segments III-V (but
not VI) with fine ncuropodial acicula embedded in the flesh. Segments V and VI
with notopodial capillaries. No notopodial hooks on segment IV. No dorsal crest
on segment. VI. Ten uncigcrous thoracic segments starUng from segment VII.
Thoracic uncini (fig. 35.2.6) with a single vertical series of five teeth above the small
rostral point and basal prow. Twenty-eight abdominal segments with square
uncigcrous pinnules. No rudimentary notopodia.
Type locality : Plcttenbcrg Bay, South .\frica.
Records: Cape (34/23/i).
Distribution : Endemic.
Subfamily AMPHARETINAE Chamberlin, 1919 (emend. Day, 1964)
Buccal tentacles either smooth with a groove along one side or papillose. Paleae
present or absent. No notopodial hooks behind the gills. Neurosetae absent from
segments III-VI. Neuropodial uncini start on segment \TI and may have one or
more series of teeth. Few (8) to many (60) abdominal segments.
Records from southern Africa
Ampharele aaitifrons (Grubc) .
as Ampharele kerguelensis (non Mcintosh)
Ampharele agulhasensis (Day) .
as Lysippe agulhasensis Day
Ampharele capensis (Day)
as Lysippe caperuis Day
Amphicteis gunneri (Sars)
Gljphanostomum abjssalis Day
as NeosabelUdes cf. elongatus
Phyllncomus hiltoni (Chamberlin) .
as Schistocornus hilloni C'hamberlin
Sabellides capensis Day ....
as Sabellides sp. ....
Sabellides (Pterampharete) luderitzi (Augener)
as Pterampharete luderitzi Augener
Sabellides oclocirrata (Sars)
Samythella qffinis Day ....
5iCs
48\Vsd
51CS
51CS
33CS, 48CS, 5iCsd
55Ca
45Pi
5iCis
44Ci
26\Vis,48\Vs,5iCs
51CS
56Cd
694
P(^I,^■(;II \i. I A of soi thern Africa
I'n^. !')■-■ I^iilila irhri/iihenui. (a) Aiiici icr end witli "tiniK"'" extriidi-d (gills im Icfl sitle
iiiiiiind . 'h) Thoraric uncinus. Ir.) Tip of bippnnatc l)r.mi lii.j. (d! Abdiiminal segment
and ])arai)<tdiun"i. Alcliiino/nidcs ca/it'risis. (h ) Tliora' u uininus. ( i- , Anterior end.
Am/jliKleii fiimneri. (g) Entire worm (three limes life size), (h) Tlioracic segments sliowing
notopodial rirri. (i) Anterior end. (j) .Abdominal segments and parapodia. (k, l) Profile
aiifl edyc-on \ iew of thor.u ic line iims. (m) Winged (apilkiry. (n) Palca.
AMPHARETIDAE 695
Key to Genera
1 Four pairs of gills ............. 2
- Three pairs of gills ............ 8
- Two pairs of gills (first row of uncini very long) .... AUCHENOPLAX*
2 Glandular ridges on prostomium (fig. 35. 2. i). (Xotopodial cirri present (fig. 35.2.11)) . 3
- No glandular ridges ............ 4
3 Fourteen uncigerous thoracic segments ...... AMPHICTEIS (p. 695)
- Eleven uncigerous thoracic segments ........ AMAGE*
4 Fourteen uncigerous thoracic segments ......... 5
- Thirteen uncigerous thoracic segments ........ LYSIPPE*
- Twelve uncigerous thoracic segments ......... 6
- Eleven uncigerous thoracic segments. (Two nephridial papillae on the branchial ridge
(fig. 35.3.a)) . . . SABELLIDES (p. 696)
5 Notopodial cirri present PHYLLAMPHICTEIS*
- Notopodial cirri absent LYSIPPIDES*
6 Tentacles papillose. Thoracic uncini with two vertical series of teeth (fig. 35. 4. c)
AMPHARETE (p. 699)
- Tentacles smooth with a groove along one side. Thoracic uncini with a single vertical
series of teeth ............. 7
7 One median papilla on the branchial ridge. Specialised notosetae in some posterior
notopodia SOSANE*
- No nephridial papillae on the branchial ridge. No specialised posterior notosetae
PHYLLOCOMUS (p. 702)
Fourteen uncigerous thoracic segments, (.\bdominal notopodia present. No glandular
SAMYTHA*
. MICROSAMYTHA*
ridges on prostomium
Thirteen uncigerous thoracic segments
Twelve uncigerous thoracic segments
Eleven uncigerous thoracic segments
Tentacles papillose. (No dorsal ridge on segment \T)
Tentacles smooth. (Thoracic uncini with a single vertical series of teeth)
SAMYTHELLA (p. 704)
9
GLYPHANOSTOMUM (p. 703)
NEOSABELLIDES*
AMPHICTEIS Gruhc, 1851
Prostomium with a pair of divergent glandular ridges. Buccal tentacles smooth
with a groove along one side. Four pairs of gills. Segments III-IV without neuro-
setae. Segment III often with paleae, segments I\'-VI with notopodial capillaries.
Fourteen uncigerous thoracic segments. Notopodial cirri present. Thoracic uncini
with a single \ertical series of teeth. Thirteen to nineteen abdominal segments,
sometimes with rudimentary notopodia as well as uncigerous pinnules.
Type species : Amphicleis gunneri Sars, 1835.
Amphtcteis gunneri (Sars, 1835)
(fig- 35-2.g-n)
AmphitriU gunneri Sars, 1835 : 50, pi. 9 fig. 30.
Amphtcteis gunneri: Fauvel, 1927: 231, fig. 80 a-k ; Day, 1961 : 527.
Body (fig. 35. 2. g) up to 60 mm. long. Prostomium (fig. 35.2.!) quadrangular with
prominent glandular ridges. Buccal tentacles rather short, smooth and grooved
along one side ; they arise from a horizontal shelf in the mouth which is occasionally
Ppfi POLYCHAETA OF SOUTHERN AFRICA
protruded. Two groups of four to five ocelli. Four pairs of smooth tapered gills
arranged two in front and two behind on the branchial ridge with a median gap
between right and left groups. Notosetae of segment III as large wingless paleae
with smoothly tapered tips (fig. 35. 2. n). Segments I\'-VI with notopodial capil-
laries (fig. 35. 2. m). Fourteen uncigerous thoracic segments starting from segment \'I I.
Papilliform dorsal cirri on both notopodia and neuiopodia (fig. 35.2.11). Thoracic
uncini (fig. 35. 2. k, 1) with a single vertical scries of four teeth above the tooth-like
rostral point and basal prow. Fifteen abdominal segments with square uncigerous
pinnules (fig. 3.').2.j) bearing a superior papilla. Rudimentary notopodia on all
segments in the form of pedunculate lobes. They originate progressively further and
further above the uncigerous pinnules (fig. 35. 2. g). Pygidium with a pair of long
anal ciiri.
Typed locality : Dredged olf Norway.
Records: South-west Africa (26'i5;s) ; C^ape (from 32,17/d to 3418 s and
34 25 s) ; Natal (31,29 s and 30 30,5).
Distribution : Atlantic from Greenland (s, d, vd, al and Norway (s, d) south to
North C!arolina (s, d), Morocco (s, d) and Ghana (s, d) ; Mediterranean; tropical
Indian Ocean (d, \xl) ; Kerguelen (s).
SABELLIDES Milne-Edwards, 1838
(including PTKRAMPHARETE Augener, 1918)
Prostomium without glandular ridges. Buccal tentacles papillose. Four pairs of
gills, arranged three in line and one behind on the branchial ridge. A pair of
nephridial papillae on the branchial ridge between the two groups of gills. Segments
III-\'I without neurosetae. Segments \' and VI with notopodial capillaries,
segment I\' usually fused to segment III and without notosetae, but notosetae often
present on segment III. Ele\en uncigerous thoracic segments. No spcciahsed
posterior notosetae. Notopodial cirri absent. Thoracic uncini with one or two series
of teeth. Between 1 1 and 18 abdominal sgcmcnts with uncigerous pinnules but with-
out rudimentary notopodia.
Type species: ? Sahella octocirrata Sars, 1835.
Key to Species
Gills papillose (fig. 33. 3. a and g). Thoracic uncini with two series of five teeth fsubuenus
Pterampharete] .......... S. (P.i hideritzi
Gills smt)oth. Thoracic uncini with a single vertical series of teeth ffig. 3",. It 'itrd k) -2
Posterior abdominal neuropodia with a long superior cirrus (lig. 35.3.1). Nephridial
papillae on branchial ridge minute ....... S. octocirrata
Pc)Sterior abdominal neuropodia with only a minute superior papilla. Xei:)hridial pajiillae
on branchial ridge large (lig. 35. 3. 1} ....... S. capensis
AMPHARETIDAE 697
Sabellides {Pterampharete, luderitzi (Augcncr, 1918)
(fig- 35-3-a-g)
Plerampharele luderitzi Augener, 1918 : 505, pi. 7 figs. 222-224, text-fig. 84 ; Day, 1961 : 527.
Body about 20 mm. long. Prostomium (fig. 35.3.3) spade-shaped. One pair of
eyes. Numerous tentacles beset with capitate papillae (fig. 35. 3. e). Four pairs of
papillose gills (or three inner pairs papillose and the outer pair smooth) arranged in a
line on the branchial ridge with a pair of small nephridial papillae between them.
Notosetae of segment III as eight to nine well developed paleae with finely tapered
tips. Segment IV achaetous and fused to segment III. Segments V and VI with
notopodial capillaries. Eleven uncigerous thoracic segments (fig. 35.3. c). Thoracic
uncini (fig. 35. 3. f, g) with two vertical series of five teeth each above the small
rostral point and recurved basal prow. Abdomen with 1 1 segments bearing uncini
on pinnules which become triangular after the first two (fig. 35. 3. d). A minute
superior papilla above the uncini. Pygidium with a pair of long anal cirri.
Type locality : Ludcritz, South West Africa.
Records: South West Africa (23/14/3, d t028/i6 Is); Cape (from 32/17/d to
34/18/s and 34/23/s).
Distribution' : Endemic.
Sabellides octocirrata (Sars, 1835)
(fig- 35-3-h-k)
Isabella octocirrata Sars, 1835: 51.
Sabellides octocirrata: Hessle, 1917 : lOI ; Fauvel, 1927 : 232, fig. 81 a-g.
Length up to 10 mm. Prostomium (fig. 35. 3. h) a triangular hood over the mouth
pinched in at the sides. One pair of eyes. Buccal tentacles pennate. Four pairs of
long smooth gills arranged obliquely on the branchial ridge with a minute pair of
nephridial papillae in the narrow median gap between them. Segment III with four
to five notopodial capillaries which are much smaller than those which follow.
Segment IV achaetous and fused to segment III. Segments V and VI with noto-
podial capillaries. Eleven uncigerous thoracic segments starting from segment \TI.
Thoracic uncini (fig. 35.3.J, k) with a single series of three to four teeth above the
rostral point and recurved basal prow. Abdomen with 14-17 segments which bear
long neuropodial cirri (fig. 35.3.1) above the rows of uncini after the first two segments.
Abdominal uncini with eight teeth arranged in two to three series. One pair of anal
cirri.
Type locality : Norway.
Records; South ^Vesl Africa (26 15s); Cape (from 32/18/s to 34 i8s and
32/28/s).
Distribution : North Atlantic from Greenland (s) and Sweden (d) south to the
Canary Islands (s) ; Mediterranean ; ? .Alaska,
s
ego
rOLVClIAE I'A OF SOUTHERN AFRRIA
Fig. 35.3. SabelUdes (Plerampharete) luderilzi. (a) Anterior end. (b) Part of papillose gill,
(c) Thoracic parapodia. (d) Fifth abdominal parapodium. (e) Tip of buccal tentacle.
(f, o) Profile and edge-on view of thoracic unciniis. Scibrllides octocirrata. (h) Anterior end.
(i) Fourth and fifth abdominal parapodia. (j, k) Profile and edge-on view of thoracic uncinus.
SabdUdes caffmsis. (l) Anterior end. (m) Fourth and fifth abdominal parapcjdia. (n, o) Pro-
file and edge-on view of thoracic uncinus.
SabelUdes capensis Day, 1961
Sabrllides ctipenwi Day, 1961 : 528, fig. 12 a-h.
Length up to 12 mm. Prostomium (fig. 35. 3. 1) a .speckled, flattened hood over
the mouth. One pair of eyes. Buccal tentacles papillose. Four pairs of smooth gills
arranged three in front and one behind on the branchial ridge with a wide median
gap between thetn in which there is a pair of ver) long nephridial pa|)il!ae. Segment
AMPHARETIDAE 699
III without setae. Segment IV achactous and fused to segment III. Segment V and
VI with small bundles of notopodial capillaries. Eleven uncigerous thoracic seg-
ments starting from segment VII. Glandular ventral pads continue to the ninth
uncigerous segment. Thoracic uncini (fig. 35. 3. n, o) with a single vertical series of
three to four teeth above the shorter rostral point and recurved basal prow. Eleven
to fourteen abdominal segments with square uncigerous pinnules each bearing a
minute papilla above the uncini. One pair of anal cirri.
Type locality : Dredged off Cape Town, South Africa.
Records: South West Africa (26/15/s) ; Cape (from 29/16/i to 34/1 7/d and
34/25/5)-
Distribution : Endemic.
AMPH ARETE Malmgren, 1866
Prostomium without glandular ridges. Buccal tentacles papillose. Four pairs of
gills. No nephridial papillae on the branchial ridge. Segments III-VI without
neurosetac. Notosetae of segment III usually enlarged to form paleae, notosetae of
segment IV usually absent, notosetae of segments V and VI always present. Twelve
uncigerous thoracic segments. Notopodial cirri absent. Thoracic uncini usually
with two series of teeth. About 12-15 abdominal segments which have uncigerous
pinnules but usually lack rudimentary notopodia.
Type SPECIES : Amphicteis aculifrons Gruhc, i860.
Key to Species
1 Paleae markedly larger than the capillaries of uncigerous thoracic segments (fig. 35.4.3). 2
- Paleae not larger than the capillaries of uncigerous thoracic segments .... 3
2 Abdominal segments with long superior cirri on the neuropodia (fig. 35. 4. d). Paleae
have smoothly tapered tips (fig. 35.4.6) ...... A. acutifrons
- Abdominal segments without superior cirri on the neuropodia. Paleae have abruptly
tapered tips (fig. 35. 4. 1) ........ A. kerguelensis
3 Conspicuous notopodial lobes above the first four uncigerous pinnules of the abdomen
(fig. 35.4. p, q). Gills extend well beyond the prostomium . . A. agulhasensis
- No notopodial lobes above the uncigerous pinnules (fig. 35. 4. j) on the abdomen. Gills do
not extend beyond the prostomium . . . . . . . .A. capensis
Ampharete acutifrons (Grube, i860)
(fig- 35-4-a-O
Amphicteis acutifrons Grube, 1 860 : 1 09.
Ampharetf aculi/roiis : Hessle, 191 7: 96.
Ampharete grubei Fauvel, 1927 : 227, fig. 79 a~p.
Body up to 80 mm. long. Prostomium (fig. 35.4.a) a triangular hood over the
mouth, and pinched in at the sides. One pair of eyes. Tentacles clearly pcnnate.
Gills smooth and extend well beyond the prostomium. They are arranged two in
front and two behind on the fused third and fourth segments. Segment III with
7O0 l'OIAC:HAi: r.\ Ol SOLTIIKKX .\iru;a
10-15 large palcac whiih taper e\enly to line tips (fit;. 35. 4. e). Segment IV witiuiut
setae but may have an achaetous notopodium. Segments \' and VI witii small
capillaries. Twelve uncigerous thoracic segments starting from segment VII.
Thoracic notosetae (fig. 35. 4. f) are winged capillaries. Thoracic uncini (fig. 35. 4. b,
c) with two Ncrtical series of five teeth above the poorly developed rostral point and
basal prow. TweKe abdominal segments bearing dorsal cirri on the uncigerous
pinnules (iig. 35. 4. d). Rudimentary notopodia absent. Pygidium with several anal
cirri, including one long pair.
TvPK mcAiirv : (ireenland.
Records: Clapc (34 17 d and 34'2f,'s).
DisTRiBi'TiON : Arctic; North Atlantic from (ireenland (s, d) and Sweden (d)
south to North Clarolina (s) and Morocco (s, d) ; Mediterranean; Noith Pacific
from the Hehring Sea to N.W. Japan and southern C!aliiorni,i.
Anipharete kerguelensis Mcintosh, 1085
(tig. 354-k-l)
Aiuphanic l:crgucleniis Mrlntosh, 1883: 42G, pi. 47 Hg. m, ]il. 2liA figs. 22 2 | ; Hcssic, njiy: mo.
Body up to 18 mm. lung. Prostomium a triangular hood o\cr the mouth. One
pair of eyes. Tentacles pi-nnatr. (iills lung ;uid smooth, extending well beyond the
prostomium. SegmeiU 11! with se\eu to eight large ]3aleae which end in blunt tips
with minute filiform jirocesses (fig. 35.4.1). Segment IV with an achaetous noto-
podium. Segments \' and \'l witli normal capillaries. Twelve uncigcnjus thoracic
segments starling from segment \'II. Thoracic uncini (fig. 35.4.k) with two vertical
series of si.x teeth above the short rostral point and ncurved basal prow. Twelve
abdominal segments. No dorsal cirri on the neur(i]iodia and Ud rudimentary noto-
podia. Pygidium cue iicled by muuiTous small p.ipill.ie.
Type locvlitv: Kcrguelcn.
Records: ?South ^\'est Africa (•J3/i4's, d).
Distribution : Antarctica ; South Georgia ; Kcrguelcn (d) ; New Zealand ;
? tropical western Africa.
Anipharete agttlhasensis (Oay, 1961)
(fig. 35.4. m-r)
Lvii/i/if (igiilluiuini\ U.iy, i')<ji : 529, fig. 12 c-k.
Body slender, up to if, mm. long. Prostomium (fig. 35.4.111) curved in front and
]Mnched in at the sides. One pair of e\es. Buccal tentacles long with two mws of
capitate papillae. Pour pairs of gills willi a wide midiaii g.ip between them, (iills
long, e.\t<'nding well beyond the prostomium. "i'.ileae" on segment III of the same
size as the normal ca]jillaries of later thoracic segments. Segment IV achaetous and
fused 1(1 m;; mi lit 111. Segment \' with a small buiidli- of ( .ipill.n ies and segment \'I
AMPHARETIDAE
701
f"'<=- 35-4- Ampharete acutifroiu. (a) Anterior end. (b, c) Profile and edge-on view of thoracic
uncinus. (d) Fourth and fifth abdominal parapodia. (e) Palea. (f) Winged capillary.
Ampharete caperuis. (g) Anterior end. (H, i) Profile and edge-on \ievv of thoracic uncinus.
(j) Fifth abdominal parapodium. Ampharete kerguelensis (after Mcintosh), (k) Profile of
thoracic uncinus. (l) Tip of palea. Ampharete aguikasensis. (m) Anterior end. (n, o) Pro-
file and edge-on view of thoracic uncinus. (p, Q, r) First, fourth and eighth abdominal
parapodia.
702 POLYCMIAi: TA Ol' SOITIIERN AI'RICA
with normal ones. Twelve uncigcrous ihoracic segments starting from segment VH.
Thoracic uncini (fig. 35. 4. n, o) with two vertical series of four lecth abo%c the small
rostral point and recurved basal prow. Twelve abdominal segments ; the first four
(fig. 35. 4. p, q) having large notopodial lobes above the square uncigerous pinnules
and the last eight (fig. 33. 4. r) without notopodia but with paddlc-shapcd uncigerous
pinnules. Pygidium with a jiair of long anal cirri lUid 10 low papillae around the
anus.
Type loc:alii v : Agulhas Bank, South Africa.
Records: Clape (34/22, s, d and ? 34/23/s).
DisTRiBLTioN : Eudemic.
Artipharete capensis (Day, ig6i)
(fig- 33-4-.S-J)
Lni/>/ic capc'iiis Day, 1961 : 530, fig. 12 h-o.
Body tapered, up to 20 mm. long. Prostomium (fig. 35. 4. g) pointed and arched
down over the mouth. One pair of eyes. Buccal tentacles beset with long papillae.
Four pairs of gills on the branchial ridge with only a narrow gap between them. Gills
short and do not extend beyond the tip of the prostomium. "Palcae" on segment III
rather smaller than the normal capillaries of later segments. Segment I\' achaetous
and fused to segment III. Segment V with weak capillaries and segment \T with
normal ones. Twehe uncigcrous thoracic segments starting from segment \'II.
Thoracic uncini (fig. 35.4.11, i) with two vertical scries of four teeth above the small
rostral point and recurved basal prow. Twelve abdominal segments without any
rudimentary notopodia. Uncigerous pinnules (fig. 35.4.J) roughly triangular
throughout with only a minute dorsal ]3apilla abo\e the uncini. Pygidium with a
pair of anal cirri.
Type locality : Saldanha Bay, South Africa.
Records : South West Africa (26/14/d) ; Gape (from 32/1 7, d to 34/18/s).
DisTRiBUTio.x ; Endemic.
PHYLLOCOMUS Giu\n , 1878
(including .S'(.7//.S77>a>.\/t".S' C:hamberlin)
Prostomium without glandular ridges. Buccal tentacles smooth witli a groove
along one side. Four pairs of gills. Segments III-\'I without neurosetae. Notopodial
capillaries present on segments I\', V and \T. Twelve uncigerous thoracic segments
without specialised notosetae. Xotopodial cirri present or absent. Thoracic uncini
with a single series of teeth. Xinnerous (over 30) abdominal segments.
Type SPECIES : Phrllucomiis crocea Gruhc, 1878a.
AMPHARETIDAE 703
Phyllocomus hiltoni (Chamberlin, 191 9)
(fig- 35-5- a-li)
Schislocomus hiltoni Chamberlin, 1919a: 17; Fauvel, 1932: 219, pi. 8 figs. 15-19; Fauvel, 1953:
41 1, fig. 216 a-e ; Day, 1957 : 112.
Body (fig. 35. 5. a) up to 50 mm. long, speckled with brown dorsaUy. Prostomium
(fig. 35. 5. b) a flattened lobe over the mouth with a rounded anterior margin. No
glandular ridges. Eyes absent. Numerous buccal tentacles which are smooth with
a groove along one side ; they arise from a hood-shaped membrane which is retractile
into a pocket-shaped cavity below the prostomium and above the true mouth.
Branchiferous ridge well developed. Four pairs of gills of which the first and outer-
most is smooth, the second and medial is unilamellate (fig. 35. 5. c) and the third and
fourth lateral pairs are bilamellate. The first and second are on the branchial ridge
and the third and fourth arise above segments V and VI. Segment III without setae
of any sort. Segments I\'-VI with notopodial capillaries but no neurosetae. Twelve
uncigerous thoracic segments starting from segment \TI. Notosetae (fig. 35. 5. h)
are all normal winged capillaries. Uncigerous pinnules on the thorax (fig. 35. 5. d)
with a superior papilla. Thoracic uncini (fig. 35. 5. e, f) with a single vertical series
of five to six teeth above the small rostral point and recurved basal prow. Abdomen
with about 50 segments, each with a rudimentary notopodial lobe and an uncigerous
pinnule with a superior cirrus ; in addition, the anterior abdominal segments have
an intermediate triangular projection between the minute rudimentary notopodium
and the uncigerous pinnule (fig. 35. 5. g). Pygidium with a circle of short cirri around
the anus.
Type locality : California.
Records: Natal (30/30/s) ; Mocambique (23/35/e).
Distribution : India (s) ; Southern California.
GLYPHANOSTOMUM Levimen, 1883
Prostomium without glandular ridges. Buccal tentacles ( ?) smooth. Three pairs
of gills. Segments III-VI without neurosetae. Segment III sometimes with paleae
and segments I V-VI always with notopodial capillaries. Eleven uncigerous thoracic
segments. Notopodial cirri absent. Thoracic uncini with two or more series of teeth.
Twelve to twenty-five abdominal segments without rudimentary notopodia but with
uncigerous pinnules.
Type SPECIES : Samjtha pallescenslhtc], 1878.
Glyphanostomum abyssale sp. nov.
(fig- 35-5-i-m)
Neosabellides cf. elongatus : Day, 1 963 : 367.
Body slender, up to 30 mm. long and encased in a narrow, closely ringed mud tube
(fig. 35. 5. m). Prostomium (fig. 35.5.1) as a flattened hood o\er the mouth. One pair
of eyes. Buchal tentacles weakly papillose. Two groups of three gills each, in a
704 P()I,VC:ilAl, r A OF SOI rilKRX Al ru:a
transverse row aeross segment II I with a median gap between lliem. Individual gills
long and smooth extending well beyond the ti]5 of the prostomiimi. Segment HI
\vithout setae. Segment l\ with a small bundle ofeapillaries and larger bundles on
\' and \'I. Eleven uneigerous thoracie segments starting on segment \'1I. The first
few segments are short but later ones are three times longer than broad. Fourteen
\i-ntral pads. Thoraeie uncini (fig. 35.^,.!;, 1) with two vertieal series of four leelh
abo\e the small rostral point and reeui\cd basal jirow. Thirty-two abdominal
segments, eaeli with an o\al uneigerous jjinnule l)earing a minute superior pajjilla
above the row of uncini (fig. 35.5.J). No rudimentary notopodi.i. Pygidium with
six to eight tapered anal cirri and a larger pair of ventral lobes.
Type locality: 2/269 metres west of Clape Town.
Records: C^ape (34/16/abyssal).
DiSTRlBlTlo.N' : \o other record.
SAMYTHELLA \'errilh 1873
Prostomium without glandular ridges. Bucchal tentacles smooth and not borne
on an elongate tentacular lobe. Three pairs of gills. Segments III-VI without
ncurosetae. Notopodial capillaries prt'sent on segnniils 1\' -\'I and sometimes on
III as well. Twelve uneigerous thoracic segments. Xutopodial cirri may be present
on the last few thoracic segments. Thoracic imcini with one oi" two series of teeth.
Up to 36 abdominal segments.
Type species: Samvthella clons,ata \'crrill, 1873.
Samythella affinis Day, 1963
Samylhtlla iifjinii Day, 19633: 435, fig. 11 b-c.
Length up to 10 mm. Prostomium (fig. 35. 5. p) a llattened hood over the mouth.
No eyes nor glandular ridges. Buccal tentacles smooth. Three pairs of smooth gills
in a continuous transverse row across segment ill without a median gaji liitween
them. Individual gills tapered and project far in Iront ol'the prostomium. Scgimnt
111 with paleae hardly larger than the normal c.ipillaries of tm( igcrous segments.
Segments IV and V with small capillaries and segment \T with normal ones.
Twelve uneigerous thoracic segments starting from segment \'II. All notosetac are
normal winged capillaries (fig. 33. 5. r) and then' ,ire no specialised posterioi- notosetae.
I hirteen glandular ventral pads. Thora( i( uncini (fig. 3f,.f",.n, o) with a single
Ncrtic al s<ries of fi\<- teeth abf)ve the rostral |ioiiil and reciu\<-d basal pi'ow. .Abdomen
of I I segments of which the first few bear small square uneigerous tori and later ones
have longer pinnules which are expanded distally (fig. 35. 5. q). No rudimentary
notopodia nor neiuopodi.tl i irri.
Type ecxality : Oil' Port Elizabeth, South Ahica.
Records: f^ape (34 23^).
DisiRiiiUTiox : .'\ single record.
AMPHARETIDAE
705
Fig. 35.5. Phyllocomus hilloni. (a) Entire worm (tlirce times life size), (b) Head with details
of tentacles, (c) Lamellate gill, (d) Thoracic parapodium. (e, f) Profile and edge-on
view of uncinus. (g) Parapodium from middle of abdomen, (h) Winged capillary seta.
Glyphanostomum abyssalis. (1) Anterior end. (j) .\bdoniinal parapodium. (k, l) Edge-on
and profile of uncinus. (m) Part of mud-tube. Samylhelta riffinix. (n, o) Edge-on \icws and
profile of thoracic uncinus. (p) Anterior end. (cj) .Vbdominal parapodium. (r) Winged
capillary seta.
7o6 I'OI.VCHAF. TA OF SOUTIII-.RN AFRICA
Family TEREBELLIDAE (;rubc, 1851
Tubicolous worms willi soft tapered bodies di\'idcd into two regions and encased
in mucous tubes encrusted witli sand or mud. Head with numerous grooved food-
gathering tentacles which are not papillose nor retractile into the mouth. Thorax
with biramous segments and often gills and lateral lobes on segments 2-4 and
glandular pads \cntrally. Notosctae are winged capillaries. Neurosetac either as
long-shafted hooks or uncini which may be avicular with arcs of denticles above the
main fang or pectinate with a single vertical series of teeth. Abdomen with numerous
segments which lack notopodia and notosctae but have neuropodia and ncuropodial
imcini. Pygidiuni without anal cirri.
BIOl.OGICiAI. NOTES
Tcrebellids are highly adapted deposit feeders. They all ha\c grooved buccal
tentacles and many of them can extend these out over the substratum for a distance
equal to the length of the body. They usually live in cpiiet areas such as lagoons,
rock pools or crevices where the organic particles settle and these ar<' picked up by
the sticky tentacles and conveyed along the groove either by ciliary action if the
particle is small, or by muscular contraction of the whole tentacle if the particle is
large. Details have been worked out by Dr. R. Phillips Dales and he has shown that
the food-laden tentacle is wiped on the upper lip which acts as a sorting device.
Apart from a few exceptions to be noted later, the terebellids are all tubicolous
and some of the larvae build tubes of diatom frustules while floating in the plankton.
The adult tube is usually attached to the side of a rock and extends back into a
crevice or down below the level of the sand. The building materials are not carefully
selected and the result is an untidy and rather fragile tube composed of sand, mud,
fiagments of shell or sponge spicules. At deep levels foraininiferan shells are com-
monly used. Only a few genera such as Lanke and Loimia coinmonly build tubes
away from rocks. Lanke often builds a stout tube which projects well above the
surface of the inud and decorates the flattened end with projecting lingers of sand
grains. Loimia occasif)nally builds its tube on open sandy beaches and the whole
tube is then shaped like a cork-screw so that it breaks before it pulls out of the sand.
Pohcinus and a few other genera such as Amaeana do not make tubes at all and creep
about naked. Polycinus li\'es among branching organisms such as algae, hydroids
and bryozoa and pulls its body around with its tentacles. It is noteworthy that the
Polycirrinae are the only group which lack gills. The one exception is the American
genus Enoplohranchus which lives on black mud. Its branchiae are cjuitc dilicrent
from those of other tereliellids and are vascular outgrowths of the notopodia.
Another unusual terebellid is Aitacama which burrows througli the silt by means of a
papillose proboscis below the mouth.
THE MAIN DIAGNOS'lIC: C:HARAtnER.S
The family Terebellidac, originally defined by Grube, was revised by M.ilmgren
(186^)) and his system of classification has been accepted with minor modifications
by all later workers. Useful discussions will be found in Saint Joseph 1894, Hesslc
TEREBELLIDAE 707
191 7, Chambcrlin 191 9 and Fauvel 1927. The most important revision is tliat of
Hessle who describes the anatomy and discusses the relation of the TerebeUidae to
the Trichobranchidac [sic], Amphictenidae ( = Pectinariidac) and Ampharetidae.
Recent work on feeding habits and the structure of the mouth and buccal tentacles
will be found in Dales 1955.
The most useful taxonomic characters include tlie structure of the head, the
number and the type of branching of the gills, the development of lateral lobes on
the first few segments, the distribution and nature of the setae, the development
of die glandular ventral pads and die number of ncphridia.
The structure of the head. Since the prostomium is not distinct in adult Terebelhds
there has been considerable doubt as to the segmental homologies of anterior
structures. Hessle (191 7) draws comparisons between the TerebeUidae and Ampha-
retidae. He notes that the buccal tentacles were originally arranged in two arcs on
either side of the mouth opening as shown by their innervation, but states that in
modern forms they are arranged in a continuous semicircle above the mouth. Above
the tentacles again there is, in the Amphectenidae and the Ampharetidae, a flattened
projecting lobe which Hessle terms the "Tcntakelmembran" and between the ten-
tacles and the mouth there is a curved ''Oberhppe". Below the mouth Hessle states
that : "Ein frei vorstehcndcr Hautsaum des Buccalsegments dient auf der Ventral-
seite als Unterlippe". Actually, as Dales (1955a) has shown, the Terebelhds have
not one lower lip but four, all derived from the buccal segment or the projecting
surface of the stomadaeum. Dales also says, however, that there is one upper lip and
that the tentacles arise from the prostomium.
The homologies of the Tcntakelmembran in the Ampharetidae have been dis-
cussed by me (Day, 1964), and it was concluded that this flattened lobe which
contains the brain and cephalic eyes is the prostomium. The buccal tentacles of
the Ampharetidae arise from the roof of the mouth and may be retracted into it. In
some species (e.g. Amphicleis gumieri (fig. 35. 2. i) ) it may be seen that they arise from
the margin of a horizontal septum which divides the mouth into a tentacular pocket
above and a food passage below. In Isolda whjdahensis (fig. 35.2.3) the same septum
is greatly elongated and clearly issues from the mouth. It bears short grooved
tentacles at its end so that these organs are not prostomial in origin but are derived
from the stomadoeum.
The buccal tentacles of the TerebeUidae are obviously homologous with those of
the Ampharetidae but neither the tentacles nor the upper lip which bears them is
retractile into the mouth. In Trichobranchus glacialis (fig. 36.1.3) which belongs to
the primitive subfamily Trichobranchinae Unking the Ampharetidae to the Tere-
beUidae, the prostomial fold which bears the eyes is separated from the long, frilly
upper Up which bears the tentacles. In the subfamily Polycirrinae the upper hp is
similar to that of Trichobranchus but in the more advanced subfamiUes Thelepinae and
Amphitrilinae the tentacular lobe is short and fused to the prostomium. The combined
structure is Uke a stout coUar which bears tentacles on its antero-ventral surface and
eye-spots on its postero-dorsal surface. Below this collar-like tentacular lobe is a second
upper lip in the shape of a glandular hood o\'erhanging the mouth opening. We may
summarise by saying that in distinction to the Ampharetids where the roof of the
7o8 I'OIA CllAK TA OI' SOL'THKRX AFRIOA
buccal cavity and tentacles nia\ he lAcrlcd al will, the ti ribrllids ha\(' the lining of
the buccal cavity permanently exerted Id limn twn upper lips and lour lower ones.
'//;<■ bianchiae. rypicallv there are three ]5airs of gills on segments 2, 3 and 4. In
dittcrent genera or even ditlerent species of the same genus these may be reduced to
two pairs or e\en one. In the subfamily Polycirrinae there are no gills at all though
this is not surprising sinci- they are not tubicolous. It may be added that as gills
are often lost and readily regenerated, the gills are often very unequal in size.
The gills may branch dichotomously or the gill fdaments may be arranged in
whorls to form a terminal tuft or "pom-pom" on a single main stem. This is a useful
taxonomic character in the genus Pisla but the relati\e lengths <il the filaments is not,
since the whole <;ill is contractile. In the subfamily Thelepinae the gills are not
branched but arise as simple filaments directly from the body w.ill. Somewhat the
same condition occurs in species oi Amphitiilv where the branchial trunk is extremely
short and the filaments are elongated.
Lattial lolics. The buccal segment may develop a prominent siiell-like lower lip, a
conical proboscis (e.g. Artacama), or wing-like lateral lobes. Similar lateral lobes are
developed from segments 2, 3 and 4 by several genera and are important taxonomic
characters. In species of Pisla and some other genera, the large lateral lobes on tiie
buccal segment may meet and fuse \entrally and then |iioject forward as a sort of
basal sheath to the tentacles. All lateral lobes are extremely glandiil.ir and function
mainly in the construction of the tube.
7 lie (lishU'iilinii mill iinline nf the setae. As shown l)y \\'ils(]n (192H) the setae first
appear on segment 2 during de\-elopment but these and others may be lost later so
that in the adult the first setigerous segment may be the second, third or fourth and
in Hauchiella there ,ire no setae at .ill. L'su.illv the neurosetae appear on a later
segment than the notosetae and the distiibution of both types of setae is of generic
importance. Typically there are about 17 bundles fif notosetae and their presence
distinguishes the anterior or tliinaeie leiiinn from the posteiior or ahiloniinal region,
although in some genera, e.g. 'Ilielepiis .nid 1 erehella the notosetae extend onto the
abdominal segments. It is of interest to note that many genera of the family Ampha-
retidae also have 17 bundles of notosetae. The identification of the first setigerous
segment is easy in genera ih.it bear gills for the first gill is always on segment 2, but
when- sills are absent as in Polreiirii^ and its allies the nnmliering of this segment is
m"ve dillicult. In this connection it may be rememlieicd that the liiitacles arise
from the stomadaeum and not from the prostomium. Some e.iih desi riptions arc
confusing for this reason.
With few exceptions the notosetae are iciinied ea/iillaries consisting of a basal shaft
and a terminal I'liiile with Literal flanges or icings on eithc'r side of a central axis.
( )ccasionally the- wings are incised or \eriiiteil ,is in ( ert.iin spe< ies of Polyeirriis but
usually the wings are smooth. The tij) of the axis may be smooth and in this case
the whole seta is referred to as a "smooth tifipeel eafiillarv" or the distal part of the axis
ni.iy de\e|o]i a saw-edged median ridge in xvhich case the seta is said to lia\c a
ileiitiiiilate ti/i. In Terehrlla the wings ol" the posterior notosetae .ire reduced or even
absent .iiul the denticulate tip is enl.irged iinlil the eiitiri' blade is dentii ul.iti-.
TEREBELLIDAE 709
The neurosclae arc usually arranged in a single vertical row on the uncigeruus ridges
or lori of the subfamilies Polycirrinae and Thelepinae. In the Amphitritinae how-
ever the posterior thoracic segments have their neurosetae [uncini) arranged in
alternating rows with one uncinus facing one way and the next the opposite way or
in two rows either face to face or back to back. On abdominal segments the un-
cigerous ridges become more prominent and often project as uncigerous pinnules. Only
very marked differences in shape are of systematic importance.
The most primitive form of neuroseta is the acicular hook found in the thorax of
the Trichobranchinac. These have long shafts and a curved rostrum surmounted
by a crest of obscure denticles. In most terebellids, however, the neurosetae are
uncini, or flattened S-shaped tooth-plates with a main fang or rostrum arising from
a broad flattened base and surmounted by a series of small teeth or denticles. These
denticles may be arranged in a single vertical series {pectinate uncini) or in a series of
transverse arcs [avicular uncini). In the latter case the number and arrangement of
the denticles may be expressed by a dental formula in which the number of teeth in
each arc is stated e.g. MF : 3-5 : 7-12 where the first arc above the main fang has
three to five teeth and the second seven to twelve. In the genus Thelepus the identifi-
cation of species depends on the shape of the basal part of the uncinus. This projects
forward as a basal prow upon which the attachment button is mounted in different ways.
To see the full details, the uncinus should be examined in edge-on-view and in profile
after treating with 5"o KOH.
Glandular ventral pads. Scattered glandular cells may be found all over the body
but speciahscd concentrations of giant glandular cells are limited to certain areas.
These are known as ventral pads or cushions and occur on the ventrum of most of the
thoracic segments. The outline of the pad changes as the worm extends and con-
tracts but the number is reasonably constant. They are found in the Amphitritinae
and Thelepinae but are absent in the Trichobranchinae and in greatly reduced
numbers in the Polycirrinae. In the latter family the pads are small and limited to
a midventral groove of the first few segments but the lateral body wall of the first
10-15 segments forms glandular swellings below the notopodia and this makes it
diflicult to find the rows of uncini.
Nephridia. Hessle (191 7) described these structures in detail and uses them in his
system of classification. This has not found favour with later workers because it
demands dissection and the nephridial papillae on which the nephridiopores open
vary in size according to the sexual state of the animal. None the less in the difficult
genus Poljcirrus the number of nephridia is an important character. The most
satisfactory method is to remove the dorsal body wall from the anterior thorax,
take out the gut and then count the nephridia which show as opaque white sacs.
GENERIC GROUPINGS
As mentioned earlier, most workers have adopted Malmgren's system of classi-
fication with minor modification. Saint Joseph (1894) stressed the importance of
the uncini and Hessle (191 7) the importance of the nephridia. The importance of
710 I'dlACHAETA Ol' SOLTHERN AFRICA
these slructuiTS is not disputed but they do not provide a practical means of identi-
fying large collections. In Fauvel (iQ'jy) a wide variety of external characters arc
used and tiie nephridia only become im|5orlant in dislinguishint; the sjseeies of
Polvciirus.
In the jiresent work the sublamilies I'richobranchinae, Polyeirrinae, 1 helepinae
and Tercbcllinac arc recognised. I agree with Fauvel in reducing Hessle's Tricho-
branchidac to the status of a subfamily and I agree with Hessle that Artacama with
its very striking proboscis is no more than a specialised member of the Terebellinae
olim Amphitritinae. The main diftcrencc from earlier works is that all genera
which lack gills are included in the sub-family Polyeirrinae. Fauvel following Hessle
has grouped the abranchiate genera Leaena, Laiiassa, Laphania, Phisidia, Pioclea and
Spinoiphaera in the Terebellinae and thus separates them from the abranchiate
genera Polvciniis, Amaeann, Lysilla and Hauchiella. Detailed discussions concerning
all these genera will be found in Hessle but I do not find his arguments sufficiently
convincing. The inclusion of all abranchiate genera in one subfamily certainly makes
the task of identification much simpler and I have broadened Malmgren's definition
of the Polyeirrinae to allow for this.
Kkv to Subfamilies
1 Tiioracic neurosclac as lont^-shafted liooks (fig. 36.1.CI). Branchiae always present
'1'richobranchinae (p. 710)
- Thoracic neiirosetae as untini (fig. 36.5. c, d). Branchiae sometimes absent ... 2
2 Branchiae absent. Tentacular lobe often large and frilled Polvcirrin.\e (p. 713)
- Branchiae present as simple filaments. Tentacular lobe usually small and collar-like
Thelepin.\e (p. 722)
- Branchiae present and usually blanched. Tentactilar lobe small and collar-like
TEREIiELI.INAE (p. 73 I )
Subfamily TRICHOBRANCHINAE Malmgien, 1866
(Trichobranehidae, Hessle k)];, ini hides (.AM-.PHORIDEA and
7 RICIIOLBRANCHIDEA iMal mgren )
Tentacular lobe expanded with the frilly margin on the dorsal surface of which
is a ridge bearing grooved tentacles. Branchiae either filamentous or united into
a single branchial trunk with four lamellate lobes. (Jlandular ventral pads not
developed. Thoracid neurosetae as long ai icular hooks with dentate crests ; abdom-
inal neurosetae as a\ieular uneini. Large nephridia present in front of the
diaphragm.
Records from southern Africa
Indiellides strnemi Sais .... 32PS, ;54C:d, 48C;d,
5iC;sd, 53C;a, - Ms
7 liihiihiaiichin ^^latiiili^ .\Ialmgren . . 44C;i, .jBCls, 5iC;sd
TEREBELLIDAE 711
Key to Genera
I A single branchial trunk distally divided into four lamellate lobes (fig. 36. r.g)
TEREBELLIDES
- Two to three pairs of simple filamentous gills (fig. 36.1.3) . TRICHOBRANCHUS
- Four pairs of simple gills OCTOBRANCHUS*
TRICHOBRANCHUS Malmgren, 1866
Tentacular lobe large and frilly with numerous grooved tentacles arising from
the dorsal surface. First segment with a pair of lateral lobes. Two to three pairs
of gills on segments 2-4, each a cylindrical tapered filament. Notosetac as smooth-
tipped capillaries on about 15 segments starting from the sixth. Neurosetae start
on setiger i, the thoracic ones being long acicular hooks with toothed crests and the
abdominal ones being avicular uncini. Five pairs of nephridia of which the first
three in segments 3-5 are the largest.
Type SPECIES : Trichnbranchus glacialis Wdlmgrcn, 1866.
Trichobranchus glacialis Malmgrcn, 1866
(fig. 36.i.a-e)
Trichobranchus glacialis Malmgren, 1866: 395, pi. 24 fig. 65; Hessle, 191 7: 131 ; Fauvel, 1927:
288, fig. too a-h.
Length up to 40 mm. Tentacular lobe (fig. 36.1.3) large with a frilly margin
overhanging the true mouth and numerous tentacles arising from its outer surface.
Prostomium with eye spots and fused to the buccal segment, which bears a pair of
horizontal wing-Uke lateral lobes and a ventral keel. Six gill filaments on segments
2, 3 and 4 arranged as three pairs with the dorsal gap betweeti right and left filaments
decreasing from the first to the diird. Notosetae on 15 segments starting from
segment 6 and long-shafted neurosetae on the same segments. Notosetae (fig. 36.i.e)
with very narrow wings and smooth tips. Shafts of neurosetae (fig. 36.i.d) con-
stricted below die head which bears an arc of about 12 long denticles above the
rostrum or main fang. Abdominal uncini (fig. 36.i.b, c) avicular and appear to have
three to four denticles above the main fang when seen in lateral view but one to
three arcs of denticles may be distinguished in face view giving the formula MF : ca
10 : 0-20. Nephridia in segments 3-7.
Type locality : Spitzbergen.
Records: Cape (34/18/s and 34/20/i).
Distribution : Arctic (s, d) ; Atlantic (from North Sea (s) to Senegal (s) and
N. Carolina (d) ; Magellan and Antarctic (d) ; Mediterranean ; N.\V. Pacific.
TEREBELLIDES Sars, 1835
Tentacular lobe large and deeply groo\cd with a frilly margin bearing numerous
tentacles on its dorsal surface but no eye-spots. Mouth iiidden in the groove of the
tentacular lobe. A prominent lower lip. A single gill with a stout trunk bearing
rolA CHAKIA OK SOL rur.RX Al RICA
Fic. 36.1. TiiclKihraihhin gliidiilts. (a) Anlcrinr cud. (b, c) EdKP-on view and profile nf
abdominal Lini_imis. (Dj Tfioracic hook, (ej W'intjcd (ajullary. 1 iieb<Uiih\ sliot'iui. (f)
Entire worm (three times life size), (o) .\ntcrior end. (H) 'I'liorac ic hook. (i. j) Edge-on
\iew and profile of abdftminal imcinus.
fiitir lamellate lobes. About 18 thoracic .setigcrs bearing winged capillary notosctae,
the first of which starts on segment 3. Xeuroselae from segment 8 (setiger 6), the
thoracic (Jiies being long-shalted hooks and the abdominal ones axiciilar tnicini.
Type species: 'TrnheUidts strncini Sars, 1835.
TEREBELLIDAE
7'3
Terebellides stroemi Sars, 1 835
(fig. 36.i.f-j)
Terebellides slroemi Sars, 1835: 48; Hessle, 1917: 137; Fauvel, 1927: 291, fig. 100 i-q.
Length up to 70 mm. Body (fig. 36.i.f) uniformly tapered. Tentacular lobe
erect and frilly bearing numerous short tentacles on its upper surface. No eyes. A
large shelf-like lower lip. The gill arises from segments 2-4 as a stout trunk
bearing four lamellate lobes which are partially fused (fig. 36.i.g). The two outer
lobes are larger than the inner pair. Anterior ventral margins of the third and succeed-
ing segments prominent. Eighteen segments with smooth-tipped notopodial capill-
aries starting on segment 3. Neurosetae start on setiger 6 (segment 8) ; the first
row of uncini are stout, unidcntate and acicular with a right angle bend below the
rostrum; uncini of subsequent rows (fig. 36.i.h) are more slender with a small
rostrum surmounted by a series of minute denticles. Abdominal uncini are borne
on long projecting pinnules and each is avicular with one to two arcs of denticles
above the main fang; formula MF : 0-3: ca 10. (fig. 36.1.1, j).
Type locality : Norway.
Records: Cape (28/14/d to 34/17/a and 33/27/s) ; Natal (29/3 1 /d) ; Mada-
gascar (s).
Distribution: Cosmopohtan from the Arctic to the subantarctic (s, d, vd, a).
Subfamily POLYCraRINAE Malmgren, 1865 (emend.)
Tentacular lobe typically enlarged and frilly with numerous tentacles but occasion-
ally short and collar-like. Gills entirely absent. Glandular ventral pads small and
restricted to the first few segments but the area around the thoracic neuropodia is
often swollen and glandular. Neurosetae are avicular uncini which may appear on
segment 5 but often further back or may even be entirely absent. Notosetae from
segment 3 or 4 but occasionally absent. Nephridia often enlarged.
Records front southern Africa
Amaeana accraensis (Augener)
as Amaea accraensis Augener
Amaeana triluhata (Sars)
Hauchiella tribullata (Mcintosh)
Lanassa capensis Day .
Leaena sp. ....
Ljsilla ubianensis Caullery
Polycirrus aurantiacus Grube .
Polycirrus cf haematndes (Claparcdc)
Polycirrus ptumosus (W'oUebaek)
Polycirrus tenuisetosus Langerhans .
Polycirrus swakopianus Augener
?48Cd
5 1 Cs, - Ns
44Ci, 51CS
5iCs
45PiNi
3iAi
^iCis
5iCd
51CS
•26\Vis
714 POLVCHAETA OF SOUTHERN AFRICA
Key to Genera
I Tentacular lobe expanded with a frilly margin (fig. 36.3.6) (Poi.ycirrinae sensu slriclo)
Tentacular lobe small and collar-like (fig. 36.4.3)
HAUCHIELLA (p. 714)
3
POLYCIRRUS I p. 715)
. AMAEANA (p. 718)
LYSILLA (p. 720)
5
8
2 Both notosetae and neurosetae entirely absent
- Notosetae present ; neurosetae entirely absent .
- Notosetae present ; neurosetae present from setiger 7 or later
3 .'Kbdornen with acicular notosetae (fig. 36. 3. g) .
- ,\bdornen without notosetae ......
4 Neurosetae start on setiger 2 .
- Neurosetae start on setiger 3-7 .....
5 Notosetae with smooth tips. Lateral lobes on segment 3 united by a dorsal ridge
LEAENA (p. 721)
- Notosetae with denticulate tips. Lateral lobes if present, not united dorsally . . 6
6 Longer notosetae with a hispid swelling before the spinulose tip. Ncphridia behind the
diaphragm large and united SPINOSPHAERA*
- Longer notosetae not swollen nor hispid. Nephridia not united ..... 7
7 Both long and short notosetae with similar spinulose tips. Lateral lobes sometimes
present on segments 2-4 ........ LANASSA (p. 721)
- Only the long notosetae with spinulose tips, the short forms with comb-like teeth. No
lateral lobes PHISIDIA*
8 Neurosetae start on setiger 3 ........ . PROCLEA*
- Neurosetae start on setiger 7 ........ . LAPHANIA*
HAUCHIELLA Lcvinsen, 1893
Tentacular lobe laige and expanded with a frilly margin. Xo eye spots. Branchiae
absent. Both notosetae and neurosetae entirely absent though notopodial papillae
are visible. Body of about 70 segments. Thorax of approximately 10 segments,
then a constriction followed by the swollen abdomen with annulated segments.
Thorax diffusely glandular. Glandular ventral pads small but distinct in both
thorax and abdomen.
Type species: Pohcirriis tiibullata Mcintosh, 1869.
Hauchiella tribullata (Mcintosh, 1869)
Pohcintis tribullata Mcintosh, 1869: 351.
Hauchiella tribullata : Hessle, 191 7: 233; Monro, 1930: 197; Monro, 1936: 184.
Body up to 46 mm. long, soft, diffusely glandular and swollen but often con-
stricted between thorax and abdomen. Tentacular lobe large and frilly, roughly
trefoil-shaped widi rather numerous grooved tentacles. Moudi ventral with two
posterior lips. No eye spots. Thorax not well-defined but appears to consist of 10
segments witlt notopodial papillae conspicuous after tlie first three. A mid-ventral
groove with small rectangular ventral pads. The groove and ventral pads are con-
tinued on the swollen abdomen which consists of about 60-70 biannulate segments.
No setae at all. Hessle reported four pairs of nephridia on the tliird, fourth, fifth
and seventh segments.
Records: C:ape (34/18/s).
Distribution: Norway (d) ; British Isles (d) ; subantarctic (s) ; Antarctic (s).
TEREBELLIDAE 715
POLYCIRRUS Gruhc, 1850
Tentacular lobe large and expanded with a frilly margin bearing numerous
tentacles but no eye-spots. Branchiae absent. Notosetae start from segment 3 and
continue for a variable number of segments. Notosetae are capillaries with smooth,
serrated or feathered wings. Neurosetae avicular with a crest of long denticles above
the main fang ; base often produced forwards. Neurosetae appear on the yth-iSth
segment and continue to the end of the abdomen. Only two to four well developed
ventral pads behind the lower Up followed by a glandular streak in a mid-ventral
groove. Glandular swellings on the vcntro-lateral sides of the thoracic segments.
Large nephridia in the first three to six setigers.
Type species: Polycirrus medusa Grube, 1850.
Key to Species
I Three pairs of nephridia. Uncini start on setigers 7-12
2
3
P. tenuiseiis
P. aurantiacus
P, cf. haematodes
P. swakopianus
- Six pairs of nephridia (fig. 36. 2. h). Uncini start on setigers 13-18
2 Twelve to twenty segments with smooth-bladed notosetae .
— Thirty to forty segments with smooth or hispid-bladed notosetae
3 Notosetae with smooth wings (fig. 36. 2.1) .....
- Notosetae with narrow, denticulate wings (fig. 36. 2. 1)
— Notosetae include some with smooth wings and others with broad plumose blades (fig.
36.3.d) ............ P. plumosus
Polycirrus tenuisetis Langerhans, 1880
(fig. 36.2.a-c)
Polycirrus tenuisetis Langerhans, 1880: no, pi. 5 fig. 25; Fauvel, 1927: 283, fig. 98 m-n ; Day,
■961 : 535-
Body evenly tapered and about 15 mm. long. Tentacular lobe longer than broad
with a frilly margin bearing rather short tentacles. Twelve to nineteen segments
bearing notosetae. Glandular swellings on the sides of nine thoracic setigers. Three
pairs of very large nephridia in setigers 1-3. Uncini commence between setigers 7
and 10. Notosetae (fig. 36. 2. c) with smooth wings. Uncini (fig. 36.2.3, b) with
one to three large teeth above the main fang and then an arc of 10-12 slender
denticles giving the formula MF : 1-3; 10-12.
Type locality : Madeira Island.
Records: Cape (34/22/s).
Distribution : English Channel (s) ; Madeira.
Polycirrus aurantiacus Grube, i860
(fig. 36.2.d-f)
Polycirrus aurantiacus Gmhe, i860: no; Fauvel, 1927: 280, fig. 97 e-k.
Body up to 100 mm. long with 120 segments. Tentacular lobe (fig. 36. 2. d) large
and divided into basal and distal parts with two types of tentacles. No eye-spots.
Thirty to forty segments with notosetae. Glandular swellings on the sides of eight to
716
^OI.^■c:^A^.•^A oi' soi-iukrx africia
Fig. 3G.2. Polrcimii Icmiu-li^. (.\. lil Profile and cds^c-on view of uncinus. (c) Xotosela.
Pohcirrus aurimilintm (after Kauvel, 19271. (d) \'cnlral view of anterior end. (1:) Juvenile
uncinus. (f) Notoseta. Pohcirrus cf lutemalode^. (g) Entire worm (12 times life size).
i'H) Dissection to show ncphridia. (ij Notoseta. (J. k) Edt;e-on view and profile of uncinus.
Polycirrui swakojiianm (after .\ugener). (i.) Notoseta. (M) Prolile of unciiuis.
TEREBELLIDAE 717
eleven thoracic setigers. Three pairs of nephridia of which the tliird in segment 6
is the largest. Notosetae (fig. 36.2.f) with smooth or hispid wings. Uncini start on
setiger 8 to 12 and the posterior ones are borne on pinnules. Each uncinus (fig.
36.2.6) with numerous denticles above the main fang in the adult.
Type locality : France.
Records : Not recorded from southern Africa.
Distribution : English Channel (i) ; Madeira (i) ; Mediterranean (i) ; Angola
Polycirrus cf. haematodes (Claparede, 1864)
(fig. 36.2.g-k)
? Aphlebina haemalodes Claparede, 1864 : 483, pi. 2 fig. i.
Polycirrus cf. haenwtodes : Day, 1961 : 535.
Body (fig. 36. 2. g) about 15 mm. long. Tentacular lobe large and trefoil-shaped
with pleated margins. Fourteen to twenty-seven segments bearing smooth-winged
notopodial capillaries (fig. 36.2.!). Uncini (fig. 36.2.J, k) from setiger 12-13 onwards
each with one to three large teeth above the main fang and the crest encircled by
about 12 long denticles giving the formula MF : 1-3: ca 12. .Six pairs of nephridia
(fig. 36. 2. h) decreasing in size from the first to the sixth. Paired ventro-lateral
glandular swelfings corresponding to the first 10-16 setigers ; they are separated by
the mid-ventral groove.
Records : Cape (from 33/1 7/s and 34/18/i, s to 35/2 i/d and 34/25/s).
Distribution : (of S. haematodes) Scotland (s) ; EngUsh Clhannel (i, s) ; Medi-
terranean.
Polycirrus swakopianus Augener, 191 8
(fig. 36.2. 1-m)
Polycirrus swakopianus .■\ugener, 1918 : 563, pi. 7 fig. 228, text-fig. 99.
Length about 16 mm. for 75 segments. Tentacular lobe trefoil-shaped. Sixteen to
twenty-five segments with notosetae. Notosetae (fig. 36.2.I) with markedly denti-
culate wings. Fourteen to fifteen segments with paired ventro-lateral cushions.
Uncini start about setiger 14 or 15 and the posterior ones are on swollen pinnules.
Six pairs of nephridia with small ncphridial papillae opening below the notopodia
of setigers 1-6. Uncini (fig. 36. 2. m) with two denticles above the main fang when
seen in profile but probably more when seen in face view ; dental formula MF : 3 : ?.
Type loc.\litv : Swakopmund, South West Africa.
Records : South West Africa (22, 14 i and 26 15 s).
Distribution : No other records.
7i8 POLYCHAEIA OF SOUTHERN AIRICIA
Polycirrus plumosus (WoUcbaek, 1912)
(fig. :56.3.a-d)
EreutJio plumosaW'oWchzLok, 1912: 82, pi. 21, figs, 1-4.
Polycirrus plumosus : Hessle, 1917: 224; Day, 1961 : 535, fig. 13 a-d.
A large species reaching 36 mm. for about 60 segments. Tentacular lobe large
and pleated. Seventeen segments with notosetae of two types : (a) with smooth
narrow wings (fig. 36. 3. c) and (b) broad with "plumose" blades formed by a scries
of transparent cones along a slender shaft (fig. 36. 3. d). Uncini start on setigcr 18
(the first abdominal) ; they are minute with few teeth, the formula being MF ; i : 1-3
(fig. 36. 3. a, b). Behind the ventral lip the ventral surface is covered with minute
glandular papillae up to setigcr 15.
Type locality : Norway
Records: Cape (32/1 7/d and 34/18/d).
Distribution: Norway, (s).
AMAEANA Hartman, 1959
( = .li\/.l£'/l Malmgren, 1866, pre-occupied)
Tentacular lobe expanded and trefoil-shaped with an anterior tonguc-hkc lobe
and a posterior frilled portion with numerous short tentacles. No eye-spots. No
branchiae. Ten to thirteen thoracic segments with notosetae starting on segment 3.
No thoracic neuropodia or neurosetae. About five achaetous segments at the end
of the thorax. Abdomen of numerous uniramous segments with conical notopodial
pinnules bearing acicular setae. No neuropodia or neurosetae. Anterior ncphridia
rather larger than posterior ones and all open on small jsapillac just below the
notopodia. Sides of body swollen and glandular below tlie thoracic iiotopodia.
Small ventral pads in a groo\e.
Type species: Polycirrus trilobata Sars, 1863.
Key to Species
I The first ten setigers bearing smooth-winged notosetae .... A. trilobata
- The first i 1-13 setigers bearing barbed notosetae ..... A. accraensis
Amaeana trilobata (Sars, 1863)
(fig. 36.3.e-hj
Polycirrus trilohala Sars, 1863 : 53.
Amaea trilobata : Fauvel, 1927 : 285, fig. 99 a-e.
Amaeana trilobata: Day, ig6i : 533.
Body (fig. 36. 3. h) about 20 mm. long for about 40 segments. It is swollen
anteriorly and cylindrical posteriorly. The first 10 setigers with long, slender
notopodia bearing fine, smooth-winged capillaries (fig. 36. 3. f) which are completely
retractile. Thorax arched dorsally, swollen and glandular laterally and with a
TEREBELLIDAE
719
Fig. 36.3. Polycirrus plumosus. (a, b) Edge-on view and profile of uncinus. (c, d) Smooth and
plumose notosetae. Amaeana trilobala. (e) \'entral view of head, (f) Thoracic winged
capillary, (g) Abdominal acicular seta, (h) Entire worm (five times life size). Lysilla
ubianensls. (i) Dorsal view of anterior end. (j) Barbed notoseta. Amaeana accraensis (after
Augener). (k) Barbed notoseta. (l) Abdominal acicular seta.
-■->o I'OLVCHAETA OF SOUTHERN AFRICA
median groo\'e ventially in which a scries of narrow ventral pads arc concealed.
A small but distinct nephridial papilla at the base of each notopodium. Five to
eight elongate ventral pads. Five to six achaetous segments between thorax and
abdomen. Abdomen with about 30 uniramous segments which are arched dorsally,
ridged ventro-laterally and grooved in the mid-ventral line. Each bears a small
conical notopodium from which projects five to six straight acicular setae with blunt
tips (fig. 36. 3. g). Colour purple in life.
Type locality : .\or\\ay.
Records: C'.ape (33,i7,d and 34 i8's) ; Natal (31/29/s, 2g/3i,s and 30/30/s).
Distribution : Arctic ; Norway ; North C^arolina (d) ; Mediterranean (i, s) ;
Japan.
Atnaeana accraensis (.^ugener, K)i8)
(fig. 36.3.k-l) ■"
Amaea accraemis Augener, 1918 : 561, pf 7 fig. 246. text-fig. 9R ; Kirkegaard, 1959 : 89, fig. 22.
Body 1 0-20 mm. long. Tentacular lobe trefoil-shaped with short broad tentacles.
Thorax with 11-13 segments bearing coarsely barbed notosetae. Notopodia long
and slender each with a nephridial papilla at its base. Thorax swollen and glandu-
lar laterally and grooved ventrally with se\en long ventral pads concelaed in the
groove. Fi\e achaetous segments foUow the thorax. Abdomen of about 26 segments
each with a conical notopodium bearing fi\'e acicular setae with straight shafts and
bent tips. (See also Polvcirrus plumosiis.)
Type LOf.ALiTY : .\ccra, Ghana.
Records: ? Cape (32'17'd, 34 i8,'s, d).
DiSTRiBUTio.N : Ghana (i; ; North C^arolina (d).
LYSILLA Malmgren, 1866
Tentacular lobe expanded and frilly with numerous tentacles restricted to the
basal margins. No gills. No lateral lobes on anterior segments. A small number
of thoracic segments (about six to twcKe) with notosetae starting on segment 3.
Neuropodia and neurosetae entirely absent both from the thorax and the abdomen.
\'entral pads narrow and restricted tn a groo\r. Nephridia of decre.ising size
present in all segments with notosetae.
Type sPEf;iES : Lv^iUn loieni M.ilmgrcn, 1866.
Key to Species
r Six thoracic segments with notosetae ...,,... L, loz'eni*
- Ten to twelve thoracic segments witli notosetae ..... Z-. ubianensis
TEREBELLIDAE 721
Lysilla ubianensis ClauUery, 1944
(fig. 36.3.H)
Lysiila ubianemh CauWery , 1944: 197, fig. 156; Day, 1957: 114.
Body swollen, about 30 mm. long. Head (fig. 36.3.1) with an anterior tongue-like
upper lip above which is a broad frilly tentacular lobe bearing numerous grooved
tentacles. Thorax with 10-12 segments bearing notosetae. Notosetae (fig. 36.3.J)
with barbed wings. Nephridial papillae on all segments bearing notosetae and
sometimes swollen from the fourth onwards. Thorax with a mid-ventral groove and
broad glandular ventro-latcral ridges. \'entral pads in the groove narrow and not
segmentally defined. Abdomen with about 30 segments. It is grooved laterally and
ventrally and segmental boundaries are indistinct. No notopodia or neuropodia
and no setae at all.
Type locality : Malay Seas.
Records: Natal (29/31/i) ; Mocambique (26/32/i).
Distribution : East Indies.
LEAENA Malmgren, 1866
Tentacular lobe small and collar-shaped. Branchiae absent. Anterior segments
with lateral lobes, those on segment 3 being united by a transverse ridge across the
dorsum. Notosetae from segment 4. Notosetae are all smooth-winged capillaries.
Avicular uncini from segment 5 (second sedger). Nephridia fairly large, not united
and present on segments 3, 6, 7, 8 and 9.
Type species: Leaena abranchiata Malmgren, 1866.
Leaena sp.
Leaena sp. Day, 1961 : 537.
Only a juvenile of 7 mm. is known. Lower lip swollen. Lateral lobe on segment
2 continuous across the ventrum. Lateral lobe on segment 3 continuous across the
dorsum. Seventeen segments with smooth, broad-winged notosetae starting on
segment 4. Uncini from segment 5 (setiger 2) and arranged in an alternating row
towards the end of the thorax. Uncini avicular with a close-set crest of denticles
above the main fang.
LAN ASS A Malmgren, 1866
Tentacular lobe small and collar-shaped. Anterior segments with or without
lateral lobes. Notosetae from segment 4. All notosetae with small wings and
denticulate tips. Neurosetae avicular and present from segment 5 (setiger 2) on-
wards ; they are arranged in double rows on some segments. Large nephridia in
segment 3, none in 4 and 5 but present again in segments 6, 7 and 8.
Type species : Lanassa nordenskioldi Malmgren, 1866.
722
POI.VCHAr, TA OF SOUTHERN ATRICA
Lanassa capensis Day, 1955
(fig. 36.4.a-c)
Lanassa capejisis Day, 1955 : 441, fig- 7 c-f.
Body about 20 mm. long, slender, and pale in colour. Tentacular lobe (fig. 36. 4. a)
small and has fi^'w tentacles. A few eye-spots. No gills. No lateral lobes on anterior
segments. Thirteen well marked ventral pads followed by a glandular streak
reaching setiger 20. Notosetac present on 27 segments starting from segment 4.
Both long and short forms of notosetae with a short winged portion followed by a
well developed denticulate tip. Uncini from setiger 2 to the end of the body. No
uncigerous pinnules. Each imcinus (fig. 36. 4. b, c) is avicular with a short base and
four to five denticles above the main fang when seen in lateral view but an edge-on
view shows four arcs of denticles giving the formula MF : 2-3 : 4-4 : ca 6 : ca 16.
Type locality : Still Bay, South Africa.
Records: Cape (34/18/i, s and 34/21/i).
Distribution : No other records.
Subfiimily THELEPINAE Malmgren, 1866
Tentacular lobe either short and collar-shaped or prolonged and frilled. Numerous
grooved tentacles and sometimes eye-spots. Branchiae usually present as simple
cylindrical filaments on segments 2, 3 and often 4. Lateral lobes sometimes present.
Glandular ventral pads present or absent. Smooth tipped notosetae* start on segment
2 or 3 and sometimes extend to within a few segments of the pygidium. Avicular
uncini with forwardly projecting bases bearing an attachment button, are present in
single or occasionally double rows ; they start between segments 4 and g but may
be entirclv absent.
Records from southern Africa
Euthelepus kinsfmhoensis Augener*
Streblosoma abranchiata Day .
Streblosoma chilensis (Mcintosh)
Streblosoma hesslei Day
Streblosoma persiea (Fauvel) .
as Pseudothelepus nyanganus Augener
Telolheleptis capensis Day
Thelepus comatus (Grube)
Thelepus pequenianus Augener
as Thelepus sp.
Thelepus plagiostoma Schmarda
as Thelepus setosus var. africanus Day
Thelepus setosus (Quatrefagcs)
Thelepus triserialis (Grube) .
26Ai
55Ca
55C"a
44C;i, ? - Ms
45Pi, - Nd
26Ai
44C;i, 51CS
44C;i
iGW'is, 44Gi, 51GS
i3Ci
27Mi, 35Gi, 360,
44Ca, 43PiNi, 51CS
4oPiNi
45 Pi, - Ps
4oNi, 44Ci, 51CS - Ms
* Eutkelfpwi kinseniboensis Augener, 1918 has denticulate notosetae,
genu.s AjuphitnU.
possibly it should be referred to the
TEREBELLIDAE 723
Key to Genera
Notosetae start on segment 2 (first branchiferous)
Notosetae start on segment 3 (second branchiferous) .
Lateral lobes present on segments 2-4. (fig. 36.5.6) .
No lateral lobes ........
Uncini from setiger 3. Tentacular lobe short and collar-like
Uncini from setiger 9. Tentacular lobe short and collar-like
Uncini absent from the thorax but present on the abdomen. Tentacular lobe elongated
with a frilled margin (fig. 36.6. b) TELOTHELEPUS (p. yji
STREBLOSOMA (p. 723)
2
EUTHELEPUS (p. 726)
3
. THELEPUS (p. 727)
PARATHELEPUS*
STREBLOSOMA Sais, 1872
Tentacular lobe small and collar-like bearing numerous tentacles. No lateral
lobes on anterior segments. Nil to three pairs of filiform gills on segments 2-4.
Smooth-tipped notopodial capillaries start on segment 2 (first branchiferous) . Uncini
start on segment 5 (setiger 4) and extend over a variable number of segments.
Ncphridia increase in size from segment 3 to 8 and are present on segment 5.
Type species : Grymaea bairdi Malmgren, 1 866.
Key to Species
1 Gills entirely absent (fig. 36.4.d) ........ S. abranchiata
- Two pairs of gills, each as a single filament (fig. 36. 4. h) .... 5. chilensis
- Three pairs of gills, each consisting of many filaments ...... 2
2 Posterior uncini arranged in loops. Thirty-three segments with notosetae . . S. hesslei
- Posterior uncini in single rows. Thirty-eight or more segments with notosetae . 5. persica
Streblosoma abranchiata Day, 1963
(fig. 36.4.d-g)
Streblosoma abranchiata Day, 1963 : 369, fig. 3 c-f.
Tubes fragile and covered with foraminiferan shells. Body uniformly slender,
about 30 mm. long by 1.5 mm. wide for 60-70 segments. Tentacular lobe (fig. 36.4.
d) short and collar-like with si.x to ten long tentacles. No eye-spots. Upper lip
overhanging the ventral mouth. Lower lip small. No lateral lobes on anterior
segments. No gills. Notosetae from segment 2 and continue for at least 19 segments,
the first three being smaller than the rest. All notosetae are smooth-tipped capill-
aries (fig. 36. 4. g). Uncini appear on segment 5 (setiger 4), are arranged in single
rows throughout and are borne on low uncigerous ridges on the abdomen. Each
uncinus (fig. 36.4. e, f) has a close-set cap of long denticles above the main fang.
They are irregularly arranged but approximate to the dental formula MF : 4-5 :
ca 8 : ca 12. Basal prow well marked and has a dorsal button.
Type locality : In 2269 metres off Cape Town.
Records: Cape (33/16/a, 34/16/a, 34/17/a).
Distribution : No other records.
724
l'()I.\ CIIAKTA OF SOL rUl.RX AIRUIA
lid. ']fi.\. Ijti/'i\ui idfurt'^'n. (a) Anterior end. (ii. (;} Proliic and cdt^c-on view of um inns.
Slri f/!i>\nr/i(i ufnanchiahi. (d) Anterior end. f i ,. F > Profile and cdt^e-on \ iew c_il uru iruis.
((;/ .\olosrla. SlTfhbiuima chilctisis fmodified from Mcintosh, i8f')-jj. (m) Anterior end.
(l) Xntoseta. (]i l.'nt inns. Slrehlnwma hrwlci. (k) Anterior end. (l, m) Proiile antl
edge-on view of unt inus. (n, oj Long and short types of notosetae.
TEREliELLIDAE 7-'5
Streblosoma chilensis (Mcintosh, 1885)
(fig. 36.4.h-j)
Eulhehpus chilensis Mcintosh, 1885 : 467, pi. 51 figs. 4-5 ; pi. 28.^ figs. 14-15.
Streblosoma clxilcnsis : Day, 1963: 370.
Body about 50 mm. long by 2.5 mm. broad. Tentacular lobe (fig. 36.4.11) short
and collar-Ukc with 12 or more long tentacles. A well developed upper lip and a
small lower lip. No eye-spots. Two pairs of gills on segments 2 and 3, each con-
sisting of a single long filament. No lateral lobes on anterior segments. Notosetae
from segment 2 (first branchiferous) and continue for at least 20 segments. Noto-
setae (fig. 36.4.1) have slender wings and smooth tips. Uncini from segment 5
(setiger 4) and ate arranged in single rows throughout. In lateral view (fig. 36.4.J)
each uncinus appears to have two teeth above the main fang, the first large and the
second very small; dental formula MF : i : 1-3. The prow of the uncinus is well
developed and has a dorsal button. About nine ventral pads. Tube massive and
made of mud.
Type locality: In 2610 fathoms, off Valparaiso.
Records: Cape (34/18/a).
Distribution: Chile (a).
Streblosoma hesslei Day, 1955
(fig. 36.4.k-o)
Streblosoma hesslei Day, 1955 : 439, fig. g-1.
Body about 25 mm. long with 73 segments. Tentacular lobe (fig. 36. 4. k) short
with numerous eye-spots. Three branchiferous segments each bearing several
filaments. Thirty-three segments with notosetae. Notosetae include long and short
smooth-winged capillaries (fig. 36.4.11, o). No lateral lobes on anterior segments.
Fifteen ventral pads. The first seven rows of uncini are in single rows but thereafter
the rows beome looped and eventually double rows are formed. Abdominal uncini
on well marked pinnules. Uncini with two rows of denticles above the main fang
and a large base with a broad attachment button at its end (fig. 36. 4. 1, m). Dental
formula of uncinus MF : 2-5 : 5-9.
Type locality : Robberg, South Africa.
Records: Cape (34/23/i) ; ? Madagascar (s).
Distribution : ? Endemic.
Streblosoma persica (Fau\el, 1908)
(fig- 36-5-a-d)
Grjmaea persica Fauvel, igo8 : 386; Fauvel, 191 1 : 419, pi. 20 figs. 35-43.
Streblosoma persica : Fauvel, 1953 : 432, fig. 229 and 230 c-m.
Body (fig. 36. 5. a) slender, evenly tapered, up to 60 mm. long. Upper lip hood-
shaped, lower hp well defined. Tentacular lobe small and collar-like with short
tentacles. No eye-spots. Three pairs of branchiae in the form of about six slender
726 POI.VCHAETA OF SOLTHERX AFRICA
filaments on segments ^-4 with a median gap between lateral groups. Notosctae
from the first branehiferous segment to the end of the abdomen. Notosetae are
short, broad-winged capillaries with smooth tips. Uncini from setiger 4, arranged
in single rows throughout (fig. 36. 4. b), and are borne on low pinnules in the abdo-
men. Dental formula of uncinus MF : 2-3: 1-5 (fig. 36. 5. d). Prow of uncinus
(fig. 36. 5. c) well developed and rounded with the dorsal button slanting forwards.
Thirteen well marked ventral pads which are not separated from the uncigerous
tori. Nephridial papillae not seen.
Type loiiality : Persian Gulf
Records: Natal (29/32/d) ; Mocambique (26/32/i).
Distribution: Persian Gulf; Gulf of Manaar ; Krusadai Is.; tropical western
Africa from Senegal (s) and Sao Thome (i) to Angola (i).
EUTHELEPUS Mcintosh, 1885
Tentacular lobe small and collar-like. Anterior segments with lateral lobes.
Filamentous gills on segments 2-4 sometimes reduced to a single pair of filaments
per segment. Smooth tipped notopodial capillaries start on segment 3 (second
branehiferous) and extend over about 20 segments. Uncini which have short bases
start on segment 5 (setiger 3). Distinct ventral pads present.
Type species: Euthelepus sctahulemis Mcintosh, 1885.
Euthelepus kinsemboensis Augcner, 191 8*
(fig- 36.5-c-i)
Eulhelepus kmemboensis Augcner, igi8 : 548, pi. 6 fig. 161, pi. 7 fig. 250, text-fig. 93.
Tentacular lobe (fig. 36. 5. c) broad and hood-shaped. Eyes not seen. \>ntral lip
stout and well defined. Small lateral lobes on segments 2-4. Simple filamentous
gills in right and left groups as follows : segment 2 with 3 and 3 ; segment 3 with 2
and 2 ; segment 4 with i and i. Notosetae start on segment 3 (second branehiferous)
and continue for 20 segments (posterior segments missing). \'entral pads not clearly
defined after the first few. Notosetae of two types - broad-winged capillaries (fig.
36. 5. h) and others with fine tapered blades with the blade serrate (fig. 36. 5. i).
Uncini from segment 5 (third setiger) onwards and are arranged in single rows. In
profile (fig. 36.5.f) the base is short with an anterior jjrow and button and three
to four teeth above the main fang; face view (fig. 36. 5. g) shows the formula as
MF : 2 : 3-4.
Type locality : Tropical western Africa.
Records : Not recorded from southern Africa.
DisrRiBUTiON : Angola (i) ; New Claledonia (i).
*PossibIy this species should be referred to AmfihitTite though notosetae are said to start on segment 3.
TEREBELLIDAE 727
THELEPUS Lcuckart, 1849
Tentacular lobe short and collar-like, usually with numerous eye-spots. Branchiae
as numerous simple filaments forming transverse rows across segments 2, 3 and
sometimes 4. No lateral lobes on anterior segments. Notosetae as smooth-tipped
capillaries starting on segment 3 (second branchiferous) and extending over at least
half the body. Uncini with the base prolonged forwards like the prow of a ship
on the upper surface of which is an attachment button. Uncini start on setiger 3
and continue to the posterior end. Ventral pads present. Nephridia in segments 4-7.
Type species : Amphitrite cincinnata Fabricius, 1 780.
Key to Species
Two branchiferous segments ......... T, cincinnatus*
Three branchiferous segments .......... 2
Uncini always in a single row .......... 3
Uncini of middle and later segments in incomplete double rows (fig. 36. 5. 1, m). (Base of
uncinus with both prow and button well developed) .... T, comatus
Prow of uncinus poorly developed .......... 4
Prow of uncinus well developed with a smaller button above ..... 5
Button of uncinus superior and separated by a notch from the small prow (fig. 36.6.3).
Notosetae stop halfway along abdomen . . . . . . . T. setosus
Button of uncinus terminal and in line with the tapered prow (fig. 36.6.0). Notosetae to
near end of abdomen ......... 7*. plagiostoma
Prow of uncinus broadly rounded and the button directed upward (fig. 36. 5. p). Notosetae
stop halfway along abdomen ........ 7". triserialis
Prow of uncinus and button equally developed with a deep notch between them (fig. 36. 5. q).
Notosetae continue to near pygidium . . . . . . . T. pequenianus
Thelepus cincinnatus (Fabricius, 1780)
Amphilrite cincinnala Fabricius, I 780 : 286.
Thelepus cincinnatm : Fauvel, 1927 : 271, fig. 95 i-m.
Body up to 200 mm. long with 100 segments. Tentacular lobe with eye-spots.
Two branchiferous segments bearing numerous branchial filaments. Glandular
ventral pads indistinct. Notosetae on 30-40 segments. Uncini always in a single
row and in the abdomen they are borne on rectangular pinnules. Prow of uncinus
broad, and much larger than the button which is directed upwards. Dental formula
of uncinus MF : 2 : 1-3.
Type locality : Greenland.
Records : No valid record from southern Africa.
Distribution: Arctic; North Atlantic ; Mediterranean (s).
POLVCllAKTA OK SOUTHKRN AFRICA
Fio. 36.5. Strfblowma perska. (ai Entire worm (three times life size), (b) Parapndia of
midcilc seE^ments. (f:, D) Profile and edge-on view of uncinus. EuthtUpm kimembormis
(modified from Augener, 1918). (h) Anterior end. (f, g) Profile and edge-on view of
uncinus. (h, i) Two types of notosetac. Thdepus cornatus. (j) Entire worm (three times
life sizei. (k, l, m) Uncigerous tori from the 3th, 7th and 12th setigers. showing the develop-
ment of double rows of Lincini. (n) unrinus. Thdepus plagiustoma. (o) Uncinus. Theh:pm
tnstnulis. (p) Uncinus. Thelepus pequenianus. (q) Uncinus.
TEREBELLIDAE 7^9
Thelepus contatus (Grube, 1859)
(fig. 36.5-J-n)
Terebella comala Grube, 1 859 : 1 09.
Thelepus comalus : Day, 1955 : 439, fig. 6 d.
Body (fig. 36.5.J) up to 60 mm. long with 100 segments. Tentacular lobe without
eye-spots. Three branchiferous segments each bearing numerous gill filaments.
Notosetae start on segment 3 and continue almost to the pygidium. Uncini of the first
five to seven sctigers in single rows (fig. 36. 5. k) ; thereafter the rows become loops
(fig. 36. 5. 1) and finally double rows are formed (fig. 36. 5. m). Abdominal uncini
borne on short ventro-lateral ridges. Prow and button of uncinus equally developed
with a deep notch between them (fig. 36. 5. n). Dental formula of uncinus MF : 2 :
11-15.
Type locality : Chile.
Records: Cape (32/18/i).
Distribution: Chile (i) ; Tristan da Cunha (i) ; S. Arabia (s).
Thelepus plagiostoma (Schmarda, 1861)
(fig. 36.5.0)
Terebella plagiostoma Schmarda, 1861 : 41, pi. 24 fig. 196.
Thelepus plagiostoma : Fauvel, 1953 : 430, fig. 228 a-f; Day, 1955 : 437, fig. 6 c.
Body up to 100 mm. long with 200 segments. Tentacular lobe with eye-spots.
Three branchiferous segments with numerous gill filaments. Notosetae continue
almost to die pygidium. Uncini always in single rows ; uncigerous tori poorly
developed and abdominal segments short and crowded. Uncini with the prow absent
and the button terminal and directed forwards (fig. 36.5.0). Dental formula of
uncinus MF : 2 : 1-3.
Type locality : New Zealand.
Records : Cape (from 34/18/i and 34/22/i, s to 32/38/i) ; Natal (from 31/29/i to
27/32/i) ; Mocambique (26/32/i and 23/35/s) ; Madagascar (s).
DiSTRiBUTio.M : Chile; subantarctic (Falkland Is. (s), Magellan Area (d), New
Zealand (d) ) ; Indo-west-Pacific (Red Sea (i) and Madagascar (i) to Japan and
Australia).
Theleptts setosus (Quatrefages, 1865)
(fig. 36.6.a)
Phenacia setosa Quatrefages, 1865: 376.
Thelepus setosus : Fauvel, 1927: 273, fig. 95 a-h.
Body up to 150 mm. long with 80-120 segments. Tentacular lobe with eye-
spots. Three branchiferous segments bearing numerous gill filaments. About 15
ventral pads. Notosetae continue two-thirds the way along the abdomen. Uncini
730 I'OLVCHAETA OF SOLTIIKRN AI-RICA
always in a single row. Uncigcrous tori well developed and form conspicuous
pinnules along the abdomen. Uncini (fig. 36. 6. a) with the prow smaller than the
button above it and a sligiil notch between the two. Dental formula of uncinus
MF : 2-3 : 0-2.
Type loc,\lity : France.
Records: Mocambiquc Lsland (i), (24/34/s).
Distribution: Nortii Atlantic from Scotland (s) and the English Channel (i, s),
IrelaTid to Senegal (i, s) ; Argentina (s), Falklands (i, s, d), iVorth Clarolina (i) and
South Georgia (i, s) ; Mediterranean (s) ; Pacific (W. Canada and Japan to S.
California and New C^alcdonia) ; Red Sea and Indian Ocean.
Theleptis triserialis (Grube, 1855)
(fig. 36.5.P)
Terebella Irisnialis Grube, 1855 : 1 18, pi. 4 fig. 16.
Tlielejnts triserialis : Fauvcl, 1927 : 274, fig. 95 n-r ; Day, 1955 : 439, fig. 6 a.
Body up to 100 mm. long, swollen anteriorly, with 100-150 segments. Tentacular
lobe with eye-spots. Three branchiferous segments bearing numerous gill filaments.
Notosctae stop halfway along abdomen. Uncini always in a single row. Abdominal
uncini borne on projecting pinnules. Prow of uncinus (fig. 36. 5. p) broad and
rounded and much larger than the dorsal button which is directed upwards. Dental
formula of uncinus MF : 1-3 : 0-5.
Type locality : Sicily.
Records: C^ape (34/20/i) ; Xatal (30/30/i) ; Madagascar (s).
Distribution: Mediterranean (i, s).
Theleptis peqtienianus Augener, 1918
(fig. 36.5.CJ)
Thelejius pequcnianin Augener, 1918 : 54-, ; Day, 1955 : 439, fig. 6 b.
Body up to 70 mm. long with 150 segments. Tentacular lobe with eye-spots.
Three branchiferous segments. Notosctae continue almost to pygidium. Uncini
always in a single row. Uncigcrous pinnules well developed on abdomen. Uncini
(fig. 36. 5. q) with the prow and button subcqual and with a deep notch between
them. Dental formula of uncinus MF : 2 : 0-5.
Type loc.m.ity : Ludcritzbuclu, South West Africa.
Records: South West Africa (26/15/i, s) ; C;ape (from 29/1 6/i and 34/18/i, s
to 33/26/s).
Distribution : Endemic.
TEREBELLIDAE 731
TELOTHELEPUS Day, 1955
Tentacular lobe large and elongate bearing numerous fine tentacles and a large
and expanded frilly margin. Branchiae as numerous simple filaments arising from
prominent bosses on two or three segments starting from segment 2. No lateral lobes
on anterior segments. Notosetae as smooth-winged capillaries starting on segment
3 (second branchiferous) and restricted to the thorax. No neurosetae on the thorax
but present as avicular uncini on the abdomen.
Type SPECIES : Telot/ielepus capensis Day, 1955.
Telothelepus capensis Day, 1955
(fig. 36.6.b-c)
Telothelepus capensis Day, 1 955 : 440, fig. 6 e-f.
Body about 50 mm. long with 100 segments. Tentacular lobe (fig. 36. 6. b) long
and bent back dorsally. Numerous short fine tentacles and an expanded frilly
margin. No eye-spots. Numerous simple gill filaments borne on a pair of bosses
on segments 2 and 3. No lateral lobes on anterior segments. Notosetae are smooth-
winged capillaries and start on the second branchiferous segment and total 15
bundles. Conspicuous nephridial papilla on segments 5-7. Ventral pads absent.
No uncini on the thorax but piesent on the abdomen and borne on square pinnules.
Each uncinus (fig. 36. 6. c) with a short base, a deep prow with a dorsal button and
a dental formula of MF : 5-6 : 8-10. No permanent tube.
Type locality : Langebaan Lagoon, South Africa.
Records: Cape (33/18/i, s, 34/23/e, 33/27/e) - locally common on sheltered
sandbanks.
Distribution : Endemic.
Subfamily TEREBELLINAE Grubc, 1850
(including AMPHITRICACEA and ARTACAMACEA Malmgren, 1865)
Tentacular lobe short and collar-like with numerous grooved tentacles. Buccal
segment usually smooth, rarely with a papillose proboscis. Gills usually present
on segments 2-4, often branched, rarely filamentous. Lateral lobes sometimes
present on segments 2-4. Glandular ventral pads present. Notosetae start on
segment 3 or 4 and are smooth-winged capillaries sometimes with denticulate tips.
Neurosetae are always present as avicular or pcctiniform uncini and start on
segment 5 ; in later thoracic segments they are in double or alternating rows.
Records from southern Africa
Amphitrite pauciseta Day .... 56Cd
Amphitrite cirrata (Miillcr) .... 55Ca
Artacama proboscidea 'SlaXragrcn . . . 56Cd
? Colymmatops granulatus Peters . . . i Pi
732
POI.YCHAE TA OF SOUTHERN ATRICA
Eupolymnia nehulosa (Montagu)
as Polymriia aebtilosa (Montagu)
as Polymriia capensis Mcintosh .
Lanice conchdega (Pallas)
as Lanice wollebaeki (Caullcry) .
Loimia medusa (Savigny)
as Terebella medusa Savigny
as Lanice fauveli Day
Nicolea macrobranchia (Schmarda) .
as Kicolea claparedii (Grubc)
as Terebella macrobranchia Schmarda
Nicolea lenustula (Montagu) .
jXicolea venustula var. africana Augcncr
Pista brevibranchia Caullery .
Pista cristala (Miillcr) .
? Pista cristala var. capensis Mcintosh
Pista fasciata (Grube) .
as Terebella fasciata Grubc
Pista folliigera Caullcry
Pista grubei Augcncr
Pista macrolobata Hessle
Pista quadrilobata (Augener)
as Nicolea quadrilobata Augcncr .
as Pista qulora Day
Pista unibranchia Day .
Terebella ehrenbergi Grube
Terebella pterochaeta Schmarda (partim)
as Leprea plerochaeta (Schmarda)
as Schmardanella pterochaeta (Schmarda)
Terebella schmardai Day
as Terebella pterochaeta Schmarda (partim)
as Terebella lapidaria Augcncr (non Kahlcr
Terebellobranchia natalensis Day
5iCsd, - Ms
27Mi, 44Ci, 45Pi
32Nd
26Ai, 48WsCs, - NsPs
4oNiPi, 45Pi, 5iCs
2GAi, 27Mi, 4oNiPi,
45PiNi, 5iCs, - NdMs
I Pi
36C.S
26\Vis, 33CS, 35Ci,
44Ci, 51CS
2iCa
4Ci
i6\Vi, 2iCi, 51 C^sd,
- NsMs
26Ais
4oPi, 45Pi
48Csd, - Ns
32Pi
15CS
27Mi, 4oNi, 44Ci,
51CS, - Ms
34\Vsd
45Pi, - PsdMs
5i^Vi
26Ai\Vi
44Ci, 51CS
56CS
45Pi, - Pd
4Ci, 3601, 40.\'i,
44Ci, 51CS
I iWi, 15CS, i6\\'i,
26\Vis
loCi, i3Ci
36C"i, 44Ci, 51CS
4Ci
26\Vis
4oNi
Key to Genera
1 A papillose proboscis below ihe mouth (fig. 36. 6. d). (Notosetae with smooth lips)
ARTACAMA (p. 733)
- No papillose proboscis ........... 2
2 Notosetae with smooth tips ........... 3
TEREBELLIDAE 733
- Notosetae with denticulate tips (fig. 36. 10. e) ........ 8
3 No lateral lobes on segments 2-4. (Two pairs of gills) . . . NICOLEA (p. 735)
- Lateral lobes present ............ 4
4 Uncini of the first row or first few rows differ from those of later segments, having the base
produced backwards as a long shaft (fig. 36.7.1). Not more than two pairs of gills . 5
- Uncini of the first row short and similar to those of later segments. Three pairs of gills 6
5 Lateral lobes present on segments 2 and 4 at least (fig. 36. 7. a). Posterior nephridia
separate PISTA (p. 736)
- Lateral lobes present on segment 3 only. Posterior nephridia united . . LANICIDES*
6 Uncini normal and avicular with close-set arcs of denticles above the main fang . . 7
- Uncini pectiniform with a single vertical series on teeth (fig. 36. 9. c, d) LOIMIA (p. 742)
7 Mouth of tube fringed and flattened (fig. 36. 8. n). Uncini set back to back on posterior
thorax LANICE (p. 743)
- Mouth of tube not fringed or flattened. Uncini not set back to back EUPOLYMNIA (p. 744)
8 Lateral lobes present on segments 2, 3 and often 4 . . . AMPHITRITE (p. 746)
- Lateral lobes absent ............ 9
9 Notosetae start on segment 3 (second branchiferous) .... NEOLEPREA*
- Notosetae start on segment 4 .......... 10
10 Two to three pairs of gills on segments 2-4 ..... TEREBELLA (p. 747)
— Three pairs of gills at intervals along thorax (fig. 36. lo.o) TEREBELLOBRANCHIA (p. 750)
ARTACAMA Malmgren, 1866
Tentacular lobe short and folded. A swollen papillose proboscis arising from the
buccal segment below the mouth. Three pairs of filamentous gills, each gill con-
sisting of several filaments arising from a common base. No lateral lobes on segments
2-4. Nephridia separate, the anterior one on segment 3 large and the posterior ones
on segments 6, 7, 8 and g smaller. Smooth-tipped notosetae start on segment 4 and
continue for 17 segments. Uncini start on segment 5 and are avicular with numerous
denticles above the main fang. Glandular ventral pads present on the thorax.
Uncigerous pinnules in abdomen expanded dorsally.
Type SPECIES : Artacama proboscidea 'hiia\mg'[i:n, 1866.
Artacama proboscidea Malmgren, 1 866
(fig. 36.6.d-g)
Artacama proboscidea Malmgren, 1866: 394; Hessle, 1917: 194, pi 2 fig. 13; Day, 1963a: 437.
Tentacular lobe (fig. 36. 6. d) small and horseshoe-shaped with a dorsal indentation.
Tentacles very short. No eye-spots. A large conical proboscis formed by the pro-
jecting lower lip and covered with numerous conical papillae. Segments 2 and 3
flanged but without real lateral lobes. Three pairs of short filiform branchiae on
segments 2-4, each in the form of a tuft of simple filaments arising from a basal
stump. .\ well developed nephridial papilla on segment 3 below the gill and smaller
ones on segments 6, 7, 8 and g postero-ventral to the notopodia. Seventeen bundles
of notosetae starting on segment 4. Uncini start on segment 5 and are arranged in
double rows on the posterior thorax. Eleven glandular ventral pads. Abdomen with
numerous segments bearing expanded and pedunculate uncigerous pinnules with
uncini restricted to part of the ventral margin.
Notosetae (fig. 36. 6. g) are winged capillaries of two lengths and end in fine.
734
POI.YCHAETA OF SOUTHERN AFRICA
Fig. 36.6. Tlielefim selosiis. (a) Uncinus. Telollulepus capemis. (b) Anterior end. (c)
Uncinus. Artacnma proboscidea. (d) Anlcrior end. (e, f) Edge-on and profile of uncinus.
(g) Short type of notoscta. Nicolea vmuiliila. (i, j) Edge-on and profile of uncinus. JVicolea
macTobranchia. (k) Entire worm (twice life size), (l) Anterior end. (m) Notoseta. (n, o)
Edge-on and profile of uncinus.
TEREBELLIDAE 735
smooth tops. Uncini (fig. 36.6.6, f) are small and avicular with a short base and a
cap of about 20 denticles above the main fang arranged in four alternating rows.
The attachment button is relatively large.
Type locality : Spitzbergen.
Records: Cape (32/17/d).
Distribution : Arctic (s, d) ; boreal North Atlantic (s) ; Bering Sea (s) ; N.W.
Japan (s) ; subantarctic (s, d).
NICOLE A Malmgren, 1866
Tentacular lobe short and collar-shaped with numerous long tentacles and many
eye-spots. Two pairs of branched gills on segments 2 and 3. No lateral lobes on
segments 2-4. Notosetae are winged capillaries with smooth tips which start on
segment 4 and extend over 15 to 40 segments. Uncini avicular with a close-set
cap of denticles above the main fang. They start on segment 5 and are set in
alternate rows back to back on a number of thoracic segments. Nephridia separate
and subequal on either side of diaphragm.
Type species: Terebella venmtula Montagu, 1818.
Key to Species
I Seventeen segments with notosetae ........ N. venustula
— Twenty-five to forty segments with notosetae . . . . . N. macrobranchia
Nicolea venustula venustula (Montagu, 1818)
(fig. 36.6 i-j)
Terebella vcmistiila Montagu, 1818: 344, pi. 13 fig. 2.
Nicolea venustula : Fauvel, 1927 : 260, fig. 90 a-f.
Body up to 50 mm. long. Tentacular lobe with numerous ocelli. Seventeen
segments with notosetae. Two pairs of long, branched gills. Thirteen to seventeen
ventral pads. Nephridial papillae on segments 3, 6 and 7. Uncini (fig. 36.6.1, j)
with two irregular arcs of denticles above the main fang according to the formula
MF : 2-4 : 3-5. Rows of uncini on the abdomen abruptly shorter than those
on the thorax and are borne on pinnules which have small superior papillae. Colour
brick red spotted with white.
Type locality : South coast of Devon, England.
Records: South West Africa (26/15/i) ; Cape (from 31/16/d and 34/18/s, d to
33/2 7/s) ; Natal (30/30/s) ; Mocambique (24/34/s) ; Madagascar (s).
Distribution : Atlantic from Greenland (s, d) and Scotland (s) south to the
Canary Is. (i) and Senegal (s, d) ; Mediterranean ; Red Sea ; Behring Sea to
north Japan.
736 POI.VCIHAF lA OF SOUTHERN AFRICA
Nicolea venustula africana Augcncr, 1918
J\"uolea venustula \ar. ajticann Augcncr, k)i8: 3J4. p\. 7 fig. ■2y-i.
Generally similar to the stem form but the imcigerous pinnules on the abdomen
ha\e a ciriiform dorsal appendage replacing the superior papillae.
Type LOc:Ai.rrv : I\ory Coast, western Africa.
DisTRiBi'Tio.N : Ivory Cloast, Angola (s).
Nicolea macrobranchia (Srhmarda, iH(3i)
(fig. 36.6. k-o)
Tert'bcllti nuicjobrmuhia Scliniarda, 1861 : 4-2. pi. 24 i'lg. 108.
NUotea macrobranchia: Augencr, 1918: -,27, pi 7 fig. 232-3, tcxt-fig. 89.
.'\ large species reaching 100 mm. (fig. 36. 6. k). Tentacular lobe (fig. 36. 6.1) with
numerous long tentacles and many eye-spots. The first gill much larger than the
second. Twenty-five to forty segments with smooth-tipped notopodial i.ipillaries
(fig. 36. 6. m). Eighteen ventral pads to setiger 17, some with transverse incisions.
Long rows of uncini on the thorax and short ones on the abdomen with an abrupt
change at setiger 18. Individual luicini (fig. 36. 6. n, o) with few teeth according to
the formula MF : 2-4 : 3-5.
Type locality : Clapc of Good Hope.
Records: Souili West Africa (22/i4;i and 26/15/i, s) ; C^ape (fiom 29/16,1 to
34/i8/i, s, 34;'23;c, i and 32,28 i) ; Natal (30/30/i).
Distribution- : Endemic.
PISTA .\Ialmgren, 1866
Tentacular lobe short, swollen and collar-like with numerous long tentacles. Eye-
spot present or absent. Lateral lobes on segments 2-4. One to two pairs of branched
gills on segments 2-3. Notosetae start on segment 4 and continue for 15 to 24
segments. They have well developed wings and, in most cases, smooth tips. LIncini
start on segment 5 and the first one or two rows usually have posteriorly elongated
bases but subsequent ones are avicular and arranged in two rows face to face.
Distinct ventral pads. Nephridium on segment 3 small or absent but large separate
ones are ]5resent in segments 6 and 7 beliiiid the diaphragm.
Type SPECIES : Amjthiliile cri\lala Muller, 1776.
Key to Species
1 Branchiae with filaments in closely packed whorls at the end of a naked trunk (fig. 36. 7. c) 2
- Branchiae with filaments which are dicliotoniously branched (fig. 36. 8. b). (Two pairs of
branchiae) ............. 4
2 Uncini of the first row without basal shafts ffig. 3G.7.C). \ single pair of branchiae
P, imibranchia (p. 737)
- Uncini of tiie first row with basal shafts (fig. 3(3. 7. tl). Two pairs of braiuhiac . 3
TEREBELLIDAE 737
3 Branchial trunks shorter than the terminal bunch of filaments (fig. 36.7.6)
P. brevibranchia (p. 737)
- Branchial trunks longer than the terminal bunch of filaments (fig. 36. 7. h) P. cristata (p. 738)
4 Uncini of the first row without basal shafts (fig. 36. 7. n) . . P. ttiacrolobata (p. 738)
- Uncini of the first row with basal shafts ......... 5
5 Seventeen segments with notosetae. Neck of uncinus bears no striated lobe ... 6
- Eighteen to twenty-four segments with notosetae. Neck of uncinus short and bears a
striated lobe .......... P.foliigera (p. 740)
6 Neck of uncinus from first row obviously elongated and shaft tapered (fig. 36. 8. c)
P. quadrilobata (p. 740)
- Neck of uncinus from first row not elongated and shaft slender not tapered (fig. 36. 8. k)
P.fasciata (p. 742)
N.B. A doubtful species has been omitted - P. crislaia var. capensis with two pairs of gills and a
single tooth above the main fang.
Pista unibranchia Day, 1963
(fig. 36.7.3-0)
Pista unibranchia Day, 1963a: 438, fig. 11 f-h.
A small species about 10 mm. long. Tentacular lobe (fig. 36.7.3) collar-shaped
with few eye-spots or none. Segments 2 and 3 with swollen, glandular lateral lobes
which almost encircle the anterior end. Small lateral lobes on segment 4. A single
median dorsal gill on segment 2 with a naked trunk and a pom-pom of short filaments
arranged in six to eight whorls. Seventeen segments with notosetae starting on
segment 4. Fifteen segments with ventral pads. Uncini start on segment 5 and
are arranged in alternating rows on the posterior thorax. About 26 long abdominal
segments with small uncigcrous pinnules. Notosetae are broad-winged capillaries
with smooth tips. Uncini (fig. 36. 7. b, c) with rounded bases which lack shafts even
in the first row. Main fang stout and surmounted with four irregular arcs of close
set denticles giving the approximate formula MF : 4 : 5 : 7 : 10.
Type locality : False Bay, South Africa.
Records: Cape (34/18/s).
Distribution : Endemic.
Pista brevibranchia Caullery, 191 5*
(fig. 36.7.d-g)
Pisla brevibranchia CauMery, 191 5: 76. Caullery, 1944: 152, fig. 121.
Body about 30 mm. long. Buccal segment (fig. 36. 7. e) with fairly small lateral
ridges and a stout curved ventral lip. Segment 4 with small dorso-lateral lobes,
segment 3 with large lateral lobes which cover most of segment 2, so that only small
ventro-lateral lobes of segment 2 are visible. Two pairs of branchiae of which one
is often missing and the other enlarged. Each gill consists of a stout trunk and a
slightly longer bunch of spirally arranged filaments. Seventeen segments with
♦Close to p. crislaia and P. typha.
738 POLVCHAETA OF SOUTHERN' AFRICA
notosctac. First row of uncini with long shafts (fig. 36.7.0!) ; succeeding uncini
with shorter shafts (fig. 36. 7. g) and four to five denticles above the main fang when
seen in profile. An edge-on \iew (fig. 36. 7. f) shows four to five arcs of denticles above
the main fang giving a dental formula of MF : 3-4 ; 4-5 : 4-5 : ca. 12.
Type loc.\lity : Malay seas.
Records: Mocambique (26/32,1).
Distribution : Indonesia (d, vd, a).
Pista cristata (Miillcr, 1776)
(fig. 36.7-lH)
Amphilrite crislata Muller, 1776: 216.
Pisia cristata : Fauvel, 1-27 : 266, fig. 93 a-g. Day, 1963 : 369.
Body up to 60 mm. long. Tentacular lobe (fig. 36. 7. h) short and collar-shaped
\vith many tentacles but no eye-spots. A hood-shaped upper lip. Buccal segment
with a straight lower lip notched in the middle and a pair of distinct, rectangular,
lateral lobes. .Segment 4 with rudimentary dorso-lateral lobes. Segment 3 with
large wing-hke lateral lobes which cover the sides of segment 2 ; below this segment
2 gives rise to a small but prominent pair of ventro-lateral lobes. Two pairs of
branchiae each with a long naked trunk and a shorter terminal bunch of filaments
arranged in whorls. Body of 70-100 segments of which 17 bear smooth-tipped
notosctac. Xehpridial papillae on segments 6 and 7. Seventeen to twenty ventral
pads. Uncini (fig. 36.7.J) avicular with rounded bases and numerous denticles
above the main fang (according to the formula MF : 5-6 : 4-5 : 12-14). Uncini of
the firstsix to ten rows with long shafts (fig. 36.7.!). Abdominal uncini without shafts
and borne on well marked pinnules.
Type locality : Norway.
Records: Cape (32,1 7,d, 33,16 a, 3416, a, 34/1 8/s) ; Xatal (29'31/s).
Distribution: Arctic; Atlantic from Sweden (d), North Carolina (s) to Gulf of
Mexico (s), English C^hannel (i, s), Senegal (s), Angola (s), Magellan area (i) ;
Mediterranean ; North Pacific from Japan to Behring Sea.
Pista macrolobata Hcsslc, 191 7
(fig. 36.7.k-n)
Piila macrolobata Hessle, 1917: 157. pi- 2 fig. 4, text-fig. 36; Fauvel, 1932: 229, text-fig. 39.
Length about 25 mm. Tentacular lobe without eye-spots. Buccal segment with
a pair of very large square lateral lobes extending forwards as a sheath to the
tentacles (fig. 36. 7. k). Second segment with a small pair of ventro-lateral lobes
which are absent in juveniles ; third segment with a pair of thick latcrall obes often
reflected back ; fourth segment with a pair of rudimentary lateral lobes or none at
all. Two pairs of dichotomously branched gills. Small ncphridia on segment 3
TEREBELLIDAE
739
Fig. 36.7. Pisla unibranchia. (a) Dorso-lateral \iew of anterior end. (b, c) Edge-on and
profile of uncinus of first row. Pista brevibranchia. (d) Profile of uncinus of first row.
(e) Anterior end. (f, g) Edge-on and profile of uncinus from 12th setiger. Pisla crisiata.
(h) Anterior end. (i, j) Profile and edge-on view of uncinus from sixth setiger. Piita
macrolobata. (k) Anterior end. (l) Winged capillary seta, (m, n) Edge-on and profile of
uncinus from first row.
740 I'Ol.VCHAETA OK SOUTHi.KX AFRICA
and large ones on segments 6 and 7. About i'2 ventral jjads. Seventeen segments
bear smooth-tipped notopodial eapillaries (fig. 36. 7. 1). Uncini (fig. 36. 7. m) with
a close-set cap of long denticles abo\c the main fang according to the formula
MF : 4-6: 4-() : ca 5: ea 12. Uncini of the first row (fig. 36.7.11) lack shafts and
are similar to those of succeeding segments. Abdominal uneini on square pinnules.
Type LocAi.rrv : Ronin Is., Pacific.
Records: Moeambique (a6 32:i, s, d) ; Madagascar (s).
DiSTRiBUTio.x : Japan ; Red Sea.
Pista foliigera Caullcrv, 1915
(fig. 36.8.f-j)
Piiliiftiliigera Caullcrv. 1915: 72; Fauvel, 1919 ; 451. pi- 17 tiR. 80, lexl-fit:!. 9.
Bod\' about 51.1 mm. long for 120 segments. Tentacular lobe (fig. 36. 8. f) with
reddish-brown tentacles and numerous eye-spots. Buccal segment with ventro-
lateral lobes and a well-de\eloped lower lip. Second segment telescoped and with
small lobes or none at all. Third segment with large, wing-like lateral lobes. Fourth
segment without obvious lateral lobes. Two pairs of diehotomously branched gills
with relatively few, stout, terminal branches. Fourteen ventral pads. Nephridial
papillae on segments 3-8. Eighteen to twenty-four segments with notosctae of two
types. The longer forms are normal winged capillaries with smooth tips and the
shorter forms (fig. 36.8.J) have finelv' spinulose tips; these arc better marked on
posterior segments. Uneini of the first two rows (fig. 36. 8. g) large with short broad
heads, a fibrillar attachment lobe below the main fang which disappears in KOH
and three arcs of teeth above the m.iin fang giving the formula MF : 2 : 5 : 6-8 ; shafts
very long and broad. Subsequent imcini avicular (fig. 36.8. h, i) ; abdominal uncini
on short ventro-lateral pinmiles.
Type locality : Uelebes.
Rec:ords : C^ape (from 3318 i and 34'18'i, s to 34'23's, d and 32/281) ; Natal
(31 2C) i) ; Madagascar (s).
Distribution: Madagascar (i) ; East Indies; tropical v\estern /\friea (i, s).
Pista quadrilobata (.\ugener, 1918)
(fig. 36.8.a-c)
yicolea quarhilobata .Vugener, 1918: 532, pi. 6 tig. 183, pi. 7 figs. 226-227. lext-fig. 90.
Pi\ta iplora Day, 1955 : 436, fig. 5 g-k.
Ptsta quadrilubata: Day. 1961 : 532, fig. 13 e.
Body (fig. 36.8.aj slender, gently tapering and about 33 mm. long. Tentacular
loljc collar-shaped with short orange tentacles and eye-spots. Buccal segment
(fig. 36. 8. b) with large lateral lobes united ventrally and extending forward as a
sheath for the tentacles. Second segment (first branchiferous) telescoped and with
small ventro-Lueral loljcs or none at all. Third segmi-nt with large, wing-like later.d
TEREBELLIDAE
74'
{f^rrm^
Fig. 36.8. Piila quadrilobala. (a) Entire worm (twice life size), (b) .\nterior end. (c) Uncinus
from first row. (d, e) Edge-on view and profile of uncinus from loth seliger. Pista foliigera.
(f) .\nterior end. (g) Uncinus from first row. (h, i) Profile and edge-on view of uncinus
from i6th setiger. (j) Short form of notoseta with spinulose tip from i6th setiger. Pisia
fasciala. (k) Uncinus from second row (after Fauvel, 1932). (L, .m) Edge-on and profile of
uncinus from abdomen (after Marenzeller, 1885). Lanke conchilega. (n) Top of tube.
(o) .-Vnterior end. (p) .\rrangement of uncini on posterior thoracic segment, (q, r) Edge-on
view and profile of thoracic uncinus.
742 POLVCHAETA OF SOUTHERN AFRICA
lobes. Two pairs of dichotomously branched gills. Seventeen segments with
smooth-tipped notopodial capillaries. Eighteen ventral pads starting from the first
branchiferous. Nepiiridial papillae not evident. Uncini avicular, those of the first
two ro\NS (fig. 36. 8. c) having long straight necks, relatively few teeth according to
tlic formula MF : 2-3 : 4-7, and long, markedly tapered shafts. Later luicini (fig.
36. 8. d, e) with more teeth (formula MF : 2-3 : 10-12) but no shafts. Abdominal
uncini borne on square pinnules.
Type locality : Swakopmund, South West .Africa.
Records: South West Afiica (22/14/i, 26/15/s and 28/16/s) ; Cape (from32/i8/s
and 34/i8;s to 32/28/i, s).
Distribution : Endemic.
Pista fasciata (Grube, 1869)
(fig. 36.8.k-m)
Terehellti {Phr-fHa) fnsclala Gruhe, 1869: 513; Ehlers, 1908: 148.
Pislafasciala: Marenzeller, 1884: 6, pi. i fig. 4; Fauvcl, 1932: 228, text-fig. 38.
Length 58 mm. for 131 segments. Tentacles short and stout. Buccal segment
with large lateral lobes ; second segment with small ventro-lateral lobes or none at
all ; third segment with lateral lobes one-third the size of those on the buccal
segment. Two pairs of stout, dichotomously branched gills one of which is often lost.
Seventeen segments with smooth-tipped winged capillaries. Fifteen ventral pads
starting on the first branchiferous segment. LIncini of all thoracic segments (fig.
36. 8. k) have slender but straight and well marked shafts ; abdominal uncini (fig.
36.8.1, m) without shafts. Crest of uncinus with a close set cap of denticles giving
the approximate formula MF : 4 : 7 ; ca 12.
Type locality : Red Sea.
Records: Cape (33/25/s).
Distribution : Indo-Pacific (from Red Sea and Zanzibar (d) ; Bay of Bengal (d),
Japan (s) to Alaska and S. California.
LOIMIA Malmgren, 1866
Tentacular lobe short and collar-like with eye-spots. Three pairs of branched
gills on segments 2-4. First few segments with lateral lobes. Notosetae with smooth
tips start on segment 4 and extend over 17 segments. LIncini start on segment 5
and are in double rows back to back on the posterior part of the thorax. Each
uncinus is pectinate with a single vertical series of teeth above the main fang. Ventral
pads present. Ncphridial papillae on segments 6-8. Nephridia small anterior to
diaphragm, large and united behind it.
Type species: Terebiiln medusa Savigny, 1820.
TEREBELLIDAE 743
Lointia medusa (Savigny, 1820)
(fig- 36-9- a-e)
Terebella medusa Savigny, 1820: 95. 8
Loimia medusa : Fauvel, 1953 : 416, fig. 218 a-f.
Length up to 250 mm. for 200 segments. Tentacular lobe short and collar-like
(fig. 36.9.3). Eye-spots present. Tentacles long, often banded with purple. First
branchia often larger than the other two. Buccal segment with a large membraneous
lower lip. Segments 2 and 3 fused with a horizontal membraneous lateral lobe and
a large united ventral pad below. \V'cll marked ventral pads up to setigcr 15.
Seventeen segments bear narrow-winged capillaries with smooth tips (fig. 36.9.6).
Uncini pectiniform with a single vertical series of about five to six teeth (fig. 36.9.
c, d). The uncini are in double rows back to back on the posterior thorax (fig.
36.9. b) and die rows are abruptly shortened at the end of the thorax and on the
posterior abdomen they are borne on square pinnules.
Type LOCALITY : Gulf of Suez.
Records: Cape (34/18/s, 34/22/s, 34/23/d) ; Natal (30/30/i and 29/32/d to
27/32/i) ; Mocambique (26/32/i and 23/35/e, i, s) ; Madagascar (s).
Distribution : English Channel (e, i, s) ; North Carohna, U.S.A. (s) ; West
Indies (s) ; tropical Indian Ocean (i, s, d) ; Red Sea (i) ; Pacific (Japan and S.
California).
LANICE Malmgren, 1866
Tentacular lobe short and collar-shaped with numerous long tentacles and eye-
spots. Three pairs of branched gills on segments 2-4. Lateral lobes on the first
three segments. Smooth-tipped notopodial capillaries start on segment 4 and are
present on a total of 17 segments. Uncini start on segment 5. They are avicular
with irregular transverse arcs of denticles above the main fang, and are set in two
rows back to back on the posterior thorax (fig. 36. 8. p). Ventral pads more or less
continuous. Nephridia small in front of diaphragm, large and united behind it.
Tube with the mouth flattened and fringed with stringy projections (fig. 36. 8. n).
Type species : Nereis conchilega Pallas, 1 766.
Lattice conchilega (Pallas, 1766)*
(fig. 36.8.n-r)
Nereis conchilega Pallas, 1766 : 131, pi. 9 figs. 14-22.
Lanice conchilega : Fauvcl, 1927 : 255, fig. 88 a-h.
Lattice tvollebaeki Caullery, 1944 : 125, fig. 99 ; Day, 1951 : 59.
Body large and soft with a swollen thorax and slender abdomen. Length up to
30 cm. Tentacular lobe with eyes sometimes present. Upper hp pronounced.
Buccal segment (fig. 36.8.0) with very large ventro-lateral lobes sometimes con-
tinuous vcntrally forming a sheath for the tentacles. No lobes on segment 2. Large
•Although the bodies and the setae of specimens from southern .\frica agree in detail ^\^lh those from
Europe, the tube is not muddy but is composed of large shell fragments and is attached to rocks.
7+4 POLVCHAETA OF SOUTHERN AFKIOA
square lateral lobes on segment 3 which cover segment 2. Three pairs of equal gills
with short trunks on segments 2-4. Fourteen to twenty \entral pads which arc not
distinct from one another and taper to a glandular streak. Ncphridial papillae on
segments 3 and 6-9. Uncini (fig. 36. 8. c], r) avicular with three teeth when seen in
profile but with three arcs of teeth when seen in face \icvv according to the formula
MF : 2 : 3-7. Abdominal uncini borne on long pinnules. Tube composed of mud
and sand or shell fragments and has a fringed mouth (fig. 36. 8. n).
Type locality: Huliand.
Records: Cape (from 34/18/5 to 3328's); Natal (31/29/i and 30/301, s to
2931 i,s); Mocambique (26'32/i and 23'35s).
Distribution : .\tLintic (from Sweden (d) to the English Clhannel (i, s) and
tropical west Africa (i, s, d) ) ; Mediterranean ; Persian Gulf (s) ; Scuithcrn
California.
EUPOLYMNIA \'crrill, 1900
Tentacular lobe short and collar-shaped with numerous tentacles. Eye-spots
present. Three pairs of branched gills on segments 2-4. Lateral lobes on segments
2-4. Smooth-tipped notopodial capillaries start on segment 4 and extend over 17
segments. Uncini with anteriorly produced bases start on segment 5 and are set
in alternating or double rows on the posterior thorax. Well marked ventral pads.
Nephridia separate and not missing from segment 5.
Type species: Amphilrite ncudi-iisii Delle Chiaje, 1825.
Etipolymiiia tiebulosa (Montagu, 181 8)
(fig. 36.9.f-h)
Terehella nebitlosa Montagu, 1818 : 343. pi. 12 fig. i.
Polymnia nebidosa : Fauvel, 1927 : 237, fig. 8g a-g ; Fauvel, 1953 : 419. fig. 219 a-g.
Tentacular lobe (fig. 36. 9. h) with numemus ocelli. .Small lateral lobes on seg-
ments 2, 3 and often 4. Fourteen to fifteen ventral pads extending to setiger 13 and
a narrow glandular streak after that. \Vinged notosetac with smooth tips start on
segment 4 and extend over i 7 segments. Uncini from segment 5, each with two large
teeth and one to five denticles abo\e the main fang in a close-set group according
to the formula MF : 2: 1-5 (fig. 36.g.f, g). Abdominal imcini borne on pinnules,
which are abru|itlv shorter than the thoracic ones. Obvious ncphridial papillae on
segments 3, 4 and 5 posterior and dorsal to the notopodia. Body soft, skin lliin
with numc'ous white dots which disappear in alcohol.
Type locality : Biitish seas.
Records: South West Africa (28,16 s; Clape (from 31 16 d and 34 i 8 i, s to
32 28 ij ; Xatal 130 30 s and 30 31 vd) ; Mocambique (26 32 i) ; Madagascar (si.
Distribution : Atlantic (from Scotland I'si and the English Channel (i, s) to
troyjical western Africa (s. A] and the Falkland Is. (d) ) ; .Mediterranean (i, s) ;
Red .Sea i ; Persian Gulf (si ; tropical Indian Ocean ^i) ; Pacific ( Jap.m).
TEREBELLIDAE
745
Fig. 36.9. Loimia medma. (a) interior end. (b) Uncigerous double row from setiger 12.
(c, d) Edge-on view and profile of thoracic uncinus. (e) Winged capillar\' seta. Eupolymnia
nebulosa. (f, g) Edge-on view and profile of thoracic uncinus. (h) Anterior end. Amphitrite
pauciseta. (i) Winged capillar^' seta and side view of tip. (j) Anterior end. (k, l) Edge-on
view and profile of uncinus. Amphitrite cinala. (m) -Anterior end. (n, o) Profile and edge-on
view of thoracic uncinus. (p) Tip of notoseta. (q) Notoseta.
746 POI.VC;HAKTA OF SOUTHERN AFRICA
AMPHITRITE Mullcr, 1771
Tentacular lobe short and collar-like witli numerous tentacles. Eye-spots seldom
present. Two to three pairs of gills starting on segment 2. Gills either branched or
as simple filaments arising from a basal stump. Lateral lobes present on segments
2-4. Notosetae are winged capillaries with finely serrated tips. They start on
segment 4 (third branchiferous) and extend over 13-25 thoracic segments. A\icular
uncini appear on segment 5 (setiger 2) and are arranged in alternating or double
rows on posterior thoracic segments. Ventral pads well developed. Abdominal
segments numerous and bear uncigerous pinnules.
Type SPECIES : Amphitrite cirrata 'WuWcv, 1771.
Key to Species
I Thirteen segments witli notosetae. Branchial filaments arise from a main stem A» pauciseta
— Seventeen segments with notosetae. Branchial filaments arise from a basal stump A, cirrata
Amphitrite pauciseta Day, 1963
(fig. 36.9.1-1)
Amphilrile pauciseta Day, 19633: 439, fig. 11 j-m.
Body up to 20 mm. long for 70 segments. Tube fragile. Tentacular lobe (fig.
36.9.J) with long tentacles. Eye-spots minute or absent. Obscure lateral lobes on
segments 2, 3 and 4. Two pairs of gills on segments 2 and 3. Each gill as a tuft
of digitiform filaments arising from a basal stump. A group of subdermal reddish
spots visible in fresh specimens at the base of each gill. Thirteen segments with noto-
setae starting on segment 4. Uncini from segment 5 (setiger 2) and arranged in
double rows on the posterior thorax. A large nephridial papilla on segment 3 lateral
to the gill base. Ten glandular ventral pads.
Abdomen smoothly tapered with 40 or more segments bearing uncigerous tori
anterir>rly and small uncigerous pinnules near the end. Notosetae (fig. 36.9.1) with
narrow blades and minutely denticulated tips. Uncini with short bases ; in profile
(fig. 36. 9. 1) they appear to have five teeth above the main fiing but an edge-on view
(fig. 36.9.!;) shows irregular arcs approximating to the formula MP : ca 5 : ca 7 :
ca 9 : ca 12.
Type locality: 160 metres ofl' Saldanha Bay, South Africa.
Records: Cape (33/1 7/d).
Distribution : Endemic.
Amphitrite cirrata Mullcr, 1771
(fig. 36.9. m-q)
Amphilrile citrala 'SiuWcr, 1771 in 1776: 216; Fauvcl, 1927: 251, fig. 86 i-o ; Day, 1963: 368.
Body up to 100 mm. long for 85 segments. Tentacular lobe (fig. 36.9.111) without
eye-spots. A prominent shelf-like lower lip. Three pairs of gills, each composed
TEREBELLIDAE 747
of numerous simple filaments arising from a basal stump. Small lateral lobes on
segments 2-4. Ten to twelve ventral pads. Seven pairs of nephridial papillae on
segments 3, and 6-1 1. Notopodial capillaries on 17 segments from segment 4
onwards. Thoracic uncini from segment 5 and arranged in two rows from setiger
7 to 16. Abdominal uncini borne on projecting pinnules. Thoracic notosetae
(fig. 36.9.P, q) arc winged capillaries with minutely denticulate tips. Thoracic
uncini (fig. 36.g.n, o) avicular with irregularly arranged cap of denticles above the
main fang approximating to the formula MF : 4-5 : 5-6 : 8-10 10-15.
Type localit\- : Iceland.
Records: Cape (34/16/a).
Distribution : Arctic ; North Atlantic from Greenland (i, s, d, vd, a) and
Sweden (d) ; Azores (a), Senegal (s) ; Mediterranean ; Behring Sea and North
Pacific to Japan ; Western Canada to central California.
TEREBELLA Linnaeus, 1 767
Tentacular lobe short and collar-shaped. Eye-spots present. Two or three pairs
of branched gills on segments 2-4. No lateral lobes on the first few segments.
Notosetae start on segment 4 (third branchiferous) and continue for a variable
number of segments. Notosetae have serrated dps. Uncini are avicular ; they start
on segment 5 (setiger 2) and are set in double or alternate rows face to face on the
posterior thorax. The uncigerous tori are poorly marked on the abdomen and are
ventral in position. Ventral pads well marked on tlie thorax. Posterior nephridia
united by a pair of lateral canals.
Type species : Terebella lapidaria Linnaeus, 1767.
Key to Species
1 Two pairs of gills ........... T. pterochaeta
- Three pairs of gills. (Xo spur at the base of the denticulate blade of the notosetae) 2
2 Notosetae stop 20-40 segments fron^ the pygidium ..... T". ehrenbergi
— Notosetae continue almost to pygidium ....... 7". schmardai
Terebella pterochaeta Schmarda, 1861
(fig. 36.10.a-f)
Terebella pterochaeta Schmarda, 1861 : 43, text-figs. a-d.
Schmardanella pterochaeta : Mcintosh, 1885: 449, pi. 53 fig. i. pi. 27.-^.
Body (fig. 36. 10. a) slender, evenly tapered, up to 100 mm. long. Head (fig.
36.io.b) with well marked upper and lower lips. Two pairs of gills with short,
close-set branches. Twenty-eight to thirty-three segments with notosetae. About
16 ventral pads followed by a narrow streak of glandular tissue in a ventral groove
along the abdomen. Uncini on low- tori which decrease evenly in size after the
first three and originate from ventral ridges on the abdomen. Notosetae of two
lengths ; anterior ones (fig. 36.io.e) with characteristic winged shafts and denticulate
748 POLVCIHAETA OF SOUTHERN AFRICA
tips which become proportionately larger on posterior segments until they form
most of the blade (lig. 36. 10. f). Xo spur at the base of the denticulate blade.
Uncini seen in profile (fig. 36.io.d) appear to ha\e three to four teeth above the
main fang but in face view (fig. 36.10.C) they show three to four irregular arcs of
teeth; dental formula : MF : 2-4 : 5-7 : 8-10. Attachment button well developed.
Type locality : Cape of Good Hope.
Records: South West Africa (22/14/i and 26/15/i, s) ; Cape (from 29/16/i and
34,18,1, s to 32;28/i) ; Natal (31/29/i to 29/31/i).
Distribution : Senegal ( ?d) ; tropical Indo-west-Pacific from the Red Sea (i)
to Indo-China and New Caledonia.
Terebella ehrenbergi Grube, 1870
(fig. 36.10.g-i)
Terebella Ekrenbfrgi Gruhe, i8yo : iii; Gravier, 1905b : 213, pi. 4 figs. 2'24-225 ; Hessle, 1917: 188.
Three pairs of branched gills. Notosetae from segment 4 to within 20-40 segments
from the pygidium. A large nephridial papilla on segment 3 between the bases of
the first and second branchiae and small nephridioporcs on segments 6-12. Thirteen
ventral pads followed bv a narrow mid-ventral glandular streak. Anterior notosetae
(fig. 36.10.1) have long, uniformly narrow wings and denticulate tips. Posterior
ones lack the wings but the denticulate tip is then enlarged to form a spiral blade
without a basal spur. Shorter notosetae with coarse teeth at the base of the blade.
Uncini (fig. 36.10. g, h) with two to three arcs of denticles above the main fang
giving the formula MF : 2-3 : 5-7. Attachment button obscure.
Type locality : Red Sea.
Records : Mocambique (26/32/1 and 26/33/d).
Distribution : Tropical Indo-vvest-Pacific from the Red Sea (s) to Burma, Japan
and New Caledonia (i).
Terebella schmardai Day, 1934
(fig. 3G.10.j-n)
Terebella schmardai Day, 1934: 6g, fig. 13 a-e.
Body up to 50 mm. in length. Tentacular lobe (fig. 36.10.I) with well developed
Hps and eye-spots. Three pairs of short, branched gills. Notosetae from segment 4
almost to the pygidium. Inconspicuous nephridial papillae on segments 3, 6, 7, 8, 9.
Ventral pads distinct on ten segments and then a narrow streak. Notosetae have
well marked wings and serrated blades (fig. 36.10.J, k) which become fairly broad
and markedly spiral but never have a spur at the base. Uncini (fig. 36.10.111, n)
TEREBELLIDAE
749
Fig. 36.10. Terebella p/erochaela. (a) Entire worm (three times life size), (b) Anterior end.
(c, d) Edge-on view and profile of thoracic uncinus. (e) .interior notoseta. (f) Posterior
notoseta. Terebella ehrenbergi. (g, h) Edge on view and profile of thoracic uncinus. (i)
Anterior notoseta. Terebella schmardai. (j) Blade of posterior notoseta. (k) Anterior noto-
seta. (l) .interior end. (.M, n) Edge-on view and profile of thoracic uncinus. Terebello-
branchia nalalerisis. (o) .interior end. (p) Thoracic uncinus. (q) Tip of posterior notoseta.
(r) Tip of anterior notoseta.
750 POLVCHAKTA OF SOUTHERN AFRICA
with about 12 denticles above the main fang arransrcd according to the formula:
MF : 2-3 : 8-10. Attachment button poorly developed.
Type locality : False Bay, South Africa.
Records: South West Africa (22/14/i to 28/16/i, s) ; Cape (from 29/16/i to
34/18, i, s and 34/24/d to 33/28/s).
Distribution : Endemic.
TEREBELLOBRANCHIA Day, 1951
Generally siinilar to Tenhella except for the arrangement of the branchiae ; there
arc three pairs of branched gills at intervals along the thorax.
Type species: Tercbdlohranchia natalaisis Day, 1951.
Terebellobranchia natalensis Day, 195 1
(fig- 36-io.o-r)
Terebellobranchia natalensis Day, 1951 : 58, fig. 8 b-e.
A small species less than 20 mm. long. No eye-spots. No lateral lobes on anterior
segments (fig. 36.10.0). Thirteen ventral pads on segments 2-15, followed by a
glandular ventral streak. Three pairs of branched gills on seginents 3, 7 and 13.
Notosetae from segment 4 and total 19 or more. Each has a denticulate tip (fig.
36.io.r) and in posterior segments the denticulate portion becomes long and spirally
twisted (fig. 36.l0.q). Uncini from segment 5, at first in a single row and later in
two rows. Each uncinus (fig. 36.10.P) with a series of three denticles above the main
fang when seen in profile but three to four arcs when seen in edge-on view. Attach-
ment button not developed.
Type locality : Durban, South Africa.
Records: Natal (30/30/i and 29/31/i).
Distribution : No other records.
SABELLIDAE
75'
Family SABELLIDAE Malmgren, 1867
Tubicolous, filter-feeding worms living in tough tubes reinforced with mud or
sand. Prostomium indistinct. The food-gathering apparatus consists of two terminal
branchial lobes which project forward from either side of the mouth and bear
numerous bipinnate radicles which are sometimes united by a web. No operculum.
Two grooved palps and often a pair of membraneous lips. The peristome often
develops a collar which enshcaths the base of the branchial lobes and in addition
there may be both eyes and otocysts internally. Body sharply divided into an
anterior thorax of about eight setigers bearing winged capillaries dorsally plus
avicular or long-shafted hooks ventrally and a posterior abdomen of few or many
segments with the setal types inverted.
Records from southern Africa
Amphiglena mediterranea (Leydig) .
Branchiomma capensis (Mcintosh) .
as Dasychone capensis Mcintosh .
as Dasychone argus var. capensis Mcintosh
Branchiomma natalensis (Kinberg) .
as Sahella natalensis Kinberg
as Dasychone natalensis (Kinberg)
as Dasychone violacea Mcintosh (non Schmarda)
as Dystilia violacea Quatrefages (non Schmarda)
Branchiomma nigromaculata (Baird) .
as Dasychone nigromaculata Baird
as Dasychone corollifera Ehlers
as Dasychone cingulata Grube
as Dasychone argus var. chefinae Mcintosh
as Dasychone bairdi Mcintosh
Branchionnna nigromaculata loandensis Treadwell
as Dasychone loandensis Treadwell
Branchiomma serratibranchus (Grube)
as Dasychone serralibranchis Grube
as Dasychone near orientalis Mcintosh .
Branchiymma violacea (Schmarda)
as Sahella violacea Schmarda
as Sabella foliifera Kinberg
as Dasychone violacea (Schmarda)
as Dasychone violacea var. capensis Day (non
Mcintosh) .....
Chone collaris Langerhans ....
Chone filicaudata Southern ....
51CS
loCd, 5iCsd
32CS
3?Ni
33Ci, 44Ci, 51CS
loCi
6Ci
-Ms
36Ci, 4oNi, 440,
45NiPi, 51CS
2iCi
syMi, 28Mi
3qPs
29Ai
39-^
4oNi, 45Ni
3 2 Pis
-Ms
4Ci
7Ci
loCi (pp.), I iW'i,
i6\Vi, 2oCi, 33CS, 26\Vis,
35Ci, 36Ci, 44Ci, 5iCs
36Ci
45Pi
48CS, 51CS, -Ns
752
rOLVCHAETA OF SOUTHERN AFRICA
Chone Icttentedti (Kinberg) .
as Parachonia letterstedti Kinberg
Desdemona ornata Banse
as Oridia parvida Day (non Ehlcrs)
Euchone capcnsis Day .
Etichone rosea Langcrhans
Fabrkia bansei Day
Fabricia capensis (Monro)
as Oridia capensis Monro .
Fabricia filamentosa Day
Fabriciola nwssambica (Day) .
as Fabricia mossambica Day
Hypsicomus capensis Day
Hjpsicomus phaeotaenia (Schmarda)
Jasmineira caudata Langerhans
Jasmineira elegans St. Joseph
Jasmineira ( ? = Fabricia) analis Ehlcrs
Megalomma quadrioculatum (W'illey)
as Branchiomma quadrioculatum Willcy
?as Brancliiomma resicnlosiim Ehlcrs (non Men
tagu)
as Branchiomma mushaensis Gravier
Megalomma bioculatum (Ehlcrs)
as Branchiomma bioculatum Ehlcrs
Megalomma vesiculosum (Montagu) .
as Branchiomma resiculosum (Montagu)
Myxicola infundibulum (Rcnicr)
as Myxicola michaelseni Augcncr
Oriopsis bansei Day ....
as Oria parvula Augcncr (non Ehlcrs)
as Oriopsis parvula Bansc (non Ehlcrs)
Oriopsis ehlersi Day
Oriopsis eimeri Langcrhans .
Oriopsis neglecta Bansc
Oriopsis parvula (Ehlcrs)
as Oria parvula Ehlcrs
Potamilla renijurmis (Lcuckart)
as Pntamilla ehlersi Monro ( ? Gr
Potamilla linguicollaris Day .
Potamilla torelli Malmgrcn .
Sabella Jusca Grubc
? as Sabella mossambica Peters
3Ci
47Ci
44Ci, 45Ne
5 1 Csd
48CS
5iCis
44Ci, 51CS
38Ci
56Cd
56Cd
45Pi
5iCd
28Mi
—Ms
48CS, 5iCsd,— Ns
i6\Vi
— Ps
35Ci, 36Gi, 4oPiNi,
44Ci, 45PiNi, 51CS
2iCi
2 8 Mi
2 6 As
45Pi
36Ci, 44Ci, 51CS
26\Vi
5i\Vi
26\Vis
47\Vis
5 ' t'S
51 CI
47\Vis, 5iCis
51CS
2iCi
'SCi, 32CS, 36Ci,
4oNi, 44Ci, 51CS, -NsPd
38Ni
5 1 Csd
15CS, 44Ci, 45PiNi,
51CS
45Pi
I Pi
SABELLIDAE 753
Sabella penicillus hinnacus .... 5iCsd
as Sabella pavonina Savigny . . . 33CS
Sabellaslarte longa (Kinberg) . . . 35Ci, 36NiCi,
4oXiPi, 44 Ci, 51CS
as Sabella longa Kinberg .... yNi
as Bispira lolutacornis Mcintosh (non Mon-
tagu) i3Ci
as Dasychone odhneri Fauvel . . . 28Mi
Sabellastarte sanctijosephi (Graxier) .
as Sabella indka Savigny (non Linnaeus) . 32 Pi
as Sabellastarte indica (Savigny) . . . 4oNiPi
BIOLOGICAL NOTE.S
The most primitive group of sabeUids is the subfamily Fabriciinae, the members of
which make temporary mucus tubes and creep about quite actively. They move
either backwards or forwards and it is not surprising that many of them have two
pairs of eyes, one on the head and the other on the pygidium. When feeding they
either spread out their branchial crowns to trap suspended pardclcs or sweep the
surface with long mobile palps to pick up deposited material.
Typical sabeUids arc, however, tubicolous and rely entirely on suspended food
particles. They are common on the sides of pylons and on rock ledges where there is
considerable water movement. They make tough tubes Uned with muco-protein
and covered with mud, sand or general debris. The base of the tube is usually
lodged in a crevice and some actually bore holes in soft coral. The mouth of the tube
often collapses when the worm retracts and Potamilla is even better protected from
predators for the end of the tube rolls up like a scroll.
When feeding, a sabellid pushes its head out of the mouth of its tube and distends
its trumpet-shaped branchial crown. The whole crown is formed of two branchial
lobes each of which bears a semi-circle of bipinnate radioles. These are ciliated and
the food current is drawn down into the funnel where the food particles are trapped
in mucus on the pinnules. From the pinnules the food strings pass down grooves on
the inner sides of the radioles and collect in basal gutters which lead them to the
palps where the particles are sorted. The lighter particles are carried to the mouth
while the heavier silt and sand particles are either ejected or used to build the tube.
The posterior end of the tube is closed and there is a special method for the removal
of faecal pellets. From the anus they are carried forward along a midventral groove
called the copragogue. At the juncdon of the thorax and abdomen the pellets move
onto the dorsal surface and eventually fall out of the mouth of the tube.
THE NLM.\ DIAGNOSTIC CHAR.ACTERS
General discussions of the family SabeUidae are given by Rioja (1923) and Fauvel
(1927). A more detailed account of the subfamily SabeUinae is given by Johansson
(1927) and recent discussions of the subfamily Fabriciinae will be found in Hartman
(1951a) and Banse (1956 and 1957).
The family SabeUidae is well defined and easily distinguished from the SerpuUdae
754 POLYCHAETA OF SOUTHERX AFRICA
by the lack of a calcareous tube and an operculum. The most important taxonomic
characters include structures associated with the branchial crown (Siebapparat),
the collar and the nature of the setae.
The branchial crown. The Sabellids are suspension feeders. The suspended particles
are collected by a pair of branchial lobes which bear a number of bipinnate radioles.
The particles entangled in mucus are conveyed down an inner groove of each
radiole to a ciliated sorting groox'e from which the food particles are led via the
ciUated palps to the mouth which is guarded by a dorsal lip and two vcntro-lateral
ones.
In certain genera (e.g. Fabricia, Manaviinkia and Oriopsis) there are only three or
four pairs of radioles but in the great majority of Sabellids the radioles are numerous
and the exact number of radioles varies so much with the size of the worm that
it loses systematic value. The colour too is of little importance. The radioles are
usually arranged in a semicircle on each branchial lobe but in some species the
radioles are arranged in a series of whorls aroimd a central axis provided with a
spiral groove to convey the mucus strings to the palps and mouth. With rare
exceptions (Manaviinkia) each radiole has a central "cartilagenous' "axis of clear
cyhndrical cells and a series of paired side branches or pinnules so that the whole
radiole is said to be bipinnate with the naked tip projecting distally. In Chone some
of the smaller ventral radioles lack pinnules and appear to act as filamentous palps.
A pair of filamentous palps is well defined in Fabriciola but normally all the radioles
possess lateral pinnules and the palps are flattened tapering organs on either side
of the mouth.
In Mvxicola the radioles of each branchial lobe are united to one another by a
membraneous web so that when the whole branchial crown is expanded, it forms a
funnel leading down to the mouth. In other genera however, the web between the
radioles is spht to varying degrees and when the splitting is complete the radioles
arc said to he free or imited only at the base. The remains of the web form a pair of
lateral fanges on the side of the individual radioles. These flanges may be incised at
intcr\als to form a series of lappets or external stylodes and in the genus Brunchiornma
(olim Dasychone) the shape and distribution of the stylodes is a uselul specific
character. In many genera the lateral flanges are lost entirely.
In several genera some of the radioles bear eve spots or ocelli. These may be
regularly arranged in pairs on the side of the radioles or irregularly scattered over
the outer surface of the dorsal radioles. In Megalomma (olim Branchiomma), well
developed sub-terminal eyes occur near the tips of the radioles. The number of
radioles which bear such eyes seems to be a function of the size of the worm.
The radioles have a respiratory as well as a nutritive function and in a few genera
there is a pulsatile branchial heart in each branchial lobe. This is not easy to see
unless it is dilated with blood. In Manayunkia however, the branchial hearts are
contained in jiromincnt palp-like organs.
The Collar. The base of each branchial lobe is usually surrounded by an elevated
fold or half fn//rtr. The two halves of the collar always remain separate dorsally but
mav be united ventrallv. Further, each half collar is often incised or notched back
SABELLIDAE 755
dorso-latcrally so that the whole collar then consists of two small dorsal lobes and
two large ventro-lateral lobes. These main differences are important but variations
in the shape of the lobes are of doubtful systematic value. In some genera (e.g.
Oriopsis) the collar segment may lack flanges and a true collar is thus absent ; in
this case the ventral lips may be enlarged and thickened to form a triangular pro-
jecdon below the mouth. This has sometimes been mistaken for a triangular
ventral collar.
Olocysts or statocjsts. These are embedded in the collar segment and are open or
closed. They are seldom easy to see and are thus of little value in classification.
Eye-spots or ocelli. Apart from the radiolar eyes mentioned earlier, ocelli may occur
in the tissues of the collar segment, on the surface of the setigerous segments between
the notosetae and neurosetae or on the pygidium. They are not of great systematic
value as those of the first segment are hidden deep inside the collar and those on
the parapodia tend to fade in alcohol.
The setae. Thoracic notosetae include ivinged capillaries, paleae with very short
broad blades and sometimes intermediate forms with hastate or spatulate blades.
The winged capillaries vary in detail and their exact structure is of little specific
value but the shape of the paleae is more important. Thoracic uncini may be in
two rows or only one. When there are two rows the first row always consists of
pick-axe setae with broad pointed blades set at right angles on top of a slender shaft.
The second row always consists of Z-shaped avicular uncini with a pointed rostrum, a
dentate crest, a broad neck, an angular prow and a short tail. \Vhcn there is only
one row of thoracic uncini these may be avicular uncini or long-shajted hooks with a
stout rostrum and dentate crest mounted on a long curved shaft.
In the abdomen the setal types are inverted. The notosetae are uncini usually
similar to those in the thoracic neuropodia, but where long-shafted hooks occur in
the thorax the abdominal uncini are cither square uncini or long-handled hooks
with numerous teeth. These abdominal uncini are arranged in a close-set row and
in order to obtain a clear profile view of a single uncinus it is necessary to excise the
parapodium and masserate it with 5°o KOH for a few minutes before making a
temporary mount in glycerine.
Subfamilies
The family Sabellidae was divided by Rioja (1923) into three subfamilies:
Sabellinae, Fabriciinae and Myxicolinae. This subdivision is followed by Fauvel (1927)
but Johansson (1927) has included Myxicola in the Fabriciinae so reducing the
number of subfamilies to two. This subdivision is adopted here and the two sub-
families arc defined below :
Subfamily SABELLINAE Rioja, 1923
Sabellidae with avicular uncini in the thorax and abdomen. Longitudinal body
muscles not nematoid. Branchial crown innervated by a single pair of nerves.
Sub-fimily FABRICIINAE Rioja, 1923 (including Myxicolinae Rioja, 1923).
756 POI.YCHAETA OF SOUT?IF.R\ AI'RKIA
Sabellidae with long-shafted hooks in the thorax. IjOnsitiidinal muscles nematoid.
Branchial crown innervated by two pairs of nerves.
Key to Genera
1 Thoracic neurosetae are a row of avicular uncini (fig. 37.5.<") and sometimes a row of
pick-axe setae as well (subfamily Sabellinae) ....... 2
- Thoracic neurosetae are a single row of long-shafted hooks (fig. 37. 5. in) (subfamily
F.abriciin.^e) ............. I I
2 Thoracic neurosetae include a row of a\icular uncini and a row of pick-axe setae (tig.
37--;'ii. vi 3
- Thoracic neurosetae are a single row of avicular uncini only ..... 9
3 C'oUar well developed, at least venlrally ......... 4
- Collar replaced by a triangular ventral projection (fig. 37.i.b) . AMPHIGLENA (p. 737)
4 Large sub-terminal eyes on some branchial radioles (fig. 37.i.h) MEGALOMMA (p. 757)
- Xo terminal eyes but lateral eye-spots sometimes present on the dorsal radioles (fig.
374-b) 5
5 Notosetae of first row very short and set in a row (fig. 37. 2. a). Base of branchial lobes
elongated. Lateral eye-spots always present on radioles . HYPSICOMUS (p. 7G0)
- Notosetae of first row in a compact tuft. Base of branchial lobes not elc^ngated. Radioles
with or without eye-spots ........... 6
6 Thoracic notosetae are all winged capillaries of varying length ..... 7
- Thoracic notosetae include winged capillaries and either paleae cjr spatulate setae
(fig. 37-i-cl) 8
7 Branchial lobes symmetrical, semicircular (fig. 37.2.0) . . SABELLA (p. 7()3)
- Branchial lobes asymmetrical, spirally coiled SPIROGRAPHIS*
8 Branchial lobes spiral with three to four whorls of radioles. Posterior thoracic notosetae
include striated sabre-setae ......... BISPIRA*
- Branchial lobes flat, semicircular. Thoracic notosetae include winged capillaries
plus paleae POTAMILLA (p. 764)
9 Thoracic notosetae include winged capillaries and subspathulate setae with abruptly
pointed tips LAONOME*
- Only winged capillaries in thorax .......... to
10 Branchial radioles with external lappets or stylodes (fig. 37.4.b) BRANCHIOMMA (p. 767)
— Branchial radioles without stylodes (fig. 37. 5. b) . . . SABELLASTARTE (p. 770)
11 .\bdominal uncini minute and arranged in long rows almost encir(!ing the segments.
Radioles completely united by a web (fig. 37. 5. j) . . MYXICOLA (p. 773)
— Abdominal uncini in short rows. Radioles either separate or united by a web . . 12
12 Radioles united for most of their length ........ 13
— Radioles either free or united only at the base ....... 14
13 The last few abdominal segments flanged to form a \entral s]3oon-shapeil hollow (lig.
37.6.a, d) . . . " EUCHONE (p. 774)
— No spoon-shaped hollow ......... CHONE (p. 776)
14 .'\bdominal uncini avicular (fig. 37. 7. p, q). Thoracic notosetae include both winged
capillaries and spathulate forms ..... JASMINEIRA (p. 779)
— .\bdominal uncini either long-shafted hooks or serj>uliform uncini. Thoracic notC)setae
are all winged ca[jillaries . . . . . . . . . . 15
15 Xo branchial radioles with lateral pinnules ..... MANAYUNKIA*
— Most branchial radioles with lateral pinnules . . . . . . . 16
16 Abdominal uncini with long shafts (fig. 37. 8. k). Three abdominal segments. Brani hial
hearts present ............. 17
— .Abdominal uncini with short bases (fig. 37.io.f). More than three abdominal segments.
.\o branchial hearts ........... 18
SABELLIDAE 757
17 A pair of ventral, elongated palps which do not contain blood vessels {fig. 37.8.3)
FABRICIOLA (p. 781)
— No elongated, filamentous palps ....... FABRICIA (p. 782)
18 Branchial radicles with marginal flanges (fig. 37.10.3) . . . ORIOPSIS (p. 785)
— Branchial radicles without flanges cr only minute ones (fig. 37.10.1) DESDEMONA (p. 788)
AMPHIGLENA Claparede, 1864
Body small, 5-15 mm. long. Branchial lobes not spiral and radicles without eyes
or external stylodes and not united by a web. A pair of long tapered palps. Mouth
between the branchial lobes which arise from a triangular ventral projection of the
peristomial segment. No collar. A pair of otocysts embedded in the peristome.
First setiger with winged capillaries only. Notosetae of setigers 2-8 include both
winged capillaries and paleae. Thoracic neurosetac are avicular uncini plus pick-
axe setae. Abdominal notosetae are avicular uncini and the neurosetae are broad-
Vkdnged capillaries.
Type species: Amphicora meditcrranea Leydig, 1851.
Antphiglena mediterranea (Leydig, 1851)
(fig. 37.i.a-g)
Amphicora mediterranea Leydig, 1951 : 328, pi. 9 figs. 6-7.
Amphiglena mediterranea: Fauvel, 1927: 324, fig. 112 k-r.
Body (fig. 37. 1. a) 6-15 mm. long with 35-40 segments. Branchial lobes each
with se\-en radioles not united by a web. Distal pinnules short and beyond these there
is a long naked tip. No collar but the peristomial segment forms a triangular ventral
lobe fused to the base of the branchial lobes (fig. 37.i.b). Eight thoracic and 24-30
abdominal segments. Setiger i with winged capillaries only. Notosetae of setigers
2-8 are winged capillaries (fig. 37.1.C) and paleae with long tips (fig. 37.i.d).
Thoracic neurosetae include pick-axe setae with heart-shaped blades ending in long
tips (fig. 37. 1, e, f) and avicular uncini with striated crests (fig. 37.i.g). Abdominal
notosetae are avicular uncini similar to those on the thorax and the neurosetae are
broad-bladed capillaries. Animal hermaphiodite.
Type locality : Mediterranean Sea.
Records: Cape (34/18/3).
Distribution : North Atlantic from the English Channel (i, s) to Madeira and
Morocco (i) ; ? AntarcUca (d) ; Mediterranean (i, s) ; Persian Gulf.
iW£G.i4LOMM.4 Johansson, 1927
(= BRANCHIOMMA Claparede, 1870, non KolUker, 1858)
Branchial lobes equal ; radioles without external stylodes but with subterminal
eyes at least on a few dorsal radioles. A pair of grooved palps. A two- or four-lobed
collar. Collar setae in a compact bundle. Thoracic notosetae are winged capillaries
758 P0I>V(;HAETA of southern AFRICA
(plus paleac in B. qundnociilaluin). Tlioracic ncurosctac arc avicular iincini plus
pick-axe setae. Abdomiual uuciui are avicular uuciui.
Type SPECIES : Amphilrile vesiculosiim Montagu, 1815.
Key to Species
1 Only winged cai^iUarics in thoracic notopcuiia ........ 2
- Winged capillaries plus paleac in thoracic notopodia (fig. 37.i.k, 1) . AT, quadrioculatuin
2 Ciollar widely gaping dorsally with very small dorsal lobes. Lateral lobes slant forward
ventrally. Two terminal eyes ........ M. bioculatum*
- Collar narrowly divided dorsally with large dorsal lobes (fig. 37.1.S). Numerous sub-
terminal eyes .......... M. vesiculosiim
Megalomma quadrioculatunt (Willcy, 1905)
(lig. 37.i.ii-o)
Branrhiormna quadrioculatum Willey, 1905 : 307.
Bramhiomnta Mushaeiisis Gravier, 1908 : 94, pi. 7 figs. 267-270, text-figs. 447-453.
Body (fig. 37.1.111) slightly tapered. Length up to 50 mm. Two or more radioles
of each branchial lobe bear large subterniinal eyes (fig. 37.1.11). Collar notched
back dorsally to form two small dorsal and two large lateral lobes. Palps short.
Thoracic notosetae of setigers 2-8 include two types of setae : (a) long slender-
winged capillaries (fig. 37. 1. 1) and (b) paleac with short tips (fig. 37.i.k). Thoracic
neurosetae include a row of pick-axe setae with symmetrical blades (fig. 37.i.i, j)
and a row of avicular uncini with fairly long shafts (fig. 37.i.n, o).
Type loc.m.ity : CV'ylon.
Records : South West Africa (26/15/i and 28/16/s) ; Cape (from 33/1 7/s, d and
34/18/i, s to 32/28/i) ; Natal (31/29/i to 29/31 /i( ; Mocambique (26/32/i, 24/34/s).
DisTRiBUTio.N : Red Sea; S. Arabia (s) ; Madagascar (i); Ceylon.
Megaloinnia vesiculosum (Montagu, 1815)
(fig- 37-i-P-u)
Amphitrite vesiculosa Montagu, 1815 : 19, pi. 5 fig. i.
Branchiomma vesiculosum: Fauvel. 1927 : 315, fig. 109 a-q.
Body up to 100 mm. or more, encased in a horny tube reinforced with sand.
Twenty to thirty branchial radioles with the tips coiled inwards, each bearing a
subterniinal eye. The dorsal radiole of each lobe is stouter than the rest. Palps
stout and triangular. Two acuminate ventral lips with pockets. Collar (fig. 37.1.S)
divided dtnsally and notched back on each side to form large rectangular dorsal
lobes and large lateral lobes which broaden to form large lappets on either side of
the mid-ventral line. Tlioracic notosetae are winged capillaries of two lengths ; the
shorter forms (fig. 37.1 .r) h,i\e slightly broader wings than the longer ones. Thoracic
ncurosctac include a\icular uncini with long tails and striated crests (g. 37.i.t, u).
SABELLIDAE
759
Fig. 37.1. Amphigtena medilenanea. (a) Entire worm (10 times life size), (b) \>ntral view of
head end. (c) Thoracic winged capillar)', (d) Palea. (e, f) Plan and profile of pick-axe
seta, (g) Thoracic uncinus. Megalomma quadrioculalum. (h) Tipof radiole withsublenninal
eye. (i, j) Pick-axe seta in plan and profile, (k) Palea. (l) Thoracic wing capillary.
(m) Entire worm (2.5 times life size), {n, o) Edge-on and profile of thoracic uncinus.
(p, q) Pick-axe seta in plan and profile. Megalomma vesiatlosum. (r) Shorter form of thoracic
notoseta. (s) Lateral view of collar, (t, v) Edge-on view and profile of thoracic uncinus.
76o POLVCIHAE lA OF SOUTHERN AFRlflA
plus a row of pick-axe setae with asymmetrical blades and tapering tips (fig. 37.1
p, cp. Abdominal uncini are essentially similar but have shorter shafts than those
on the thorax while the capillaries arc similar to the thoracic ones.
Type locality : Devon, En^dand.
REt:oRDS : Mocambique (26;32;i and 23/35;'$).
Distribution : North Atlantic from the English Ghanncl (e, i, s) to Cape Verde
Is. (s) and Senegal (i, s) ; Mediterranean (s) ; West Indies (s) ; Indian Ocean.
Megalontma bioculatunt (Ehlers, 1887)
Bjanchiommn biiniilahim Elilcrs, 1887 : 260, pi. 53 iigs. 1-9.
A small species up to 20 mm. long. Branchial lobes with 10-20 radioles. The
dorsal radiole on each side has a subterminal eye. Palps broadly flanged. Collar
narrowly di\ided dorsally and each half notched back to form a small dorsal lobe
with a rounded end and a large lateral lobe which slants forward ventrally to end in
a pointed triangular lappet. (Ehlers saw no dorsal lobes at all.) Thoracic notosctae
with wings broad at the base. Uncini include a row of pick-axe setae with sym-
metrical blades tapering to long points and avicular uncini with long shafts.
Type locality: Southern Florida, U.S.A.
Records : Not recorded from sonthern Africa.
Distribution : North Carolina (s), Florida and tropical western Africa (i, s).
HYPSICOMUS Gruhe, 1870
Branchial lobes borne on a long stalk, each lobe bearing nimierous radioles without
external stylodcs but with eye-spots in rows. A collar present. First setiger with the
setae short and set in a slanting row, not a compact bundle. Subsequent thoracic
notosctae include both winged capillaries and paleac. Thoracic neurosetae include
both pick-axe setae and avicular uncini. Abdominal notosctae arc avicular uncini
and the notosctae are winged capillaries and paleae.
Type spe<::ies : Sahella phaeolacnia Schmarda, 1861.
Key to Species
Collar high and slanting. Collar setae are stout smooth-winged capillaries (fig. 37.*::.d).
Thoracic paleae with short tips ........ H. capensis
Collar I(jw and straight. C'ollar setae with \ery short broad blades (fig. 37. 2. i, j).
Thoracic paleae without tips ........ H. phaeotaenia
SABELLIDAE 761
Hypsicomus capensis Day, 1961
(fig. 37.2.a-h)
Hypsicomus capensis Hay , 1961: 537, fig. I3.f.-ni.
A long slender worm with a tough brown translucent tube. Branchial lobes
(fig. 37. 2. a) borne on a long stalk ; each lobe with 12 radioles united by a web for
one quarter of their length. Each radiole with about 10 pairs of eye-spots about
half way along and a blunt tip. Collar divided dorsally and spht ventrally, each
half notched back dorsally to form a small rounded dorsal lobe and a large lateral
lobe which slants forward ventrally to form a triangular lappet Collar setae
(fig. 37. 2. d) are stout smooth-winged capillaries arranged in a line. Notosetae of
setigers 2-8 include two to three smooth-winged capillaries (fig. 37.2.6) and numer-
ous paleae with rounded blades ending in pointed tips (fig. 37. 2. b). Neurosetae of
setigers 2-8 include a row of pick-axe setae with transparent tapered blades (fig.
37. 2. c) and a row of avicular uncini with fairly broad tails and striated crests (fig.
37.2.f). Abdominal notosetae are avicular uncini similar to the thoracic ones and
the neurosetae are slender-winged capillaries (fig. 37.2.11) and paleae with much
longer blades than those on the thorax (fig. 37. 2. g).
Type locality : Agulhas Bank, South Africa.
Records: Cape (34/23/d).
Distribution : No other records.
Hypsicomus phaeotaenia (Schmarda, 1861)
(fig. 37.2.i-n)
Sabella phaeolaenia Schmarda, 1861 : 35, pi. 22 figs. 188.
Hypsicomus phaeotaenia: Fauvel, 1953: 447, fig. 236 a-1.
Hypsicomus pigmcniaius Gravier, 1908 : 81, pi. 6 figs. 252-254, text-figs. 424-431.
Body slender, up to 50 mm. long, commonly boring in old coral. Branchial
lobes borne on a common stalk. A pair of tapering palps. Each branchial lobe with
about 15-20 radioles which are united for a third their length. Distally the free
radioles have small irregularly arranged lateral eye-spots and fairly long naked tips.
Collar (fig. 37. 2. k) very low with a smooth straight margin which is quite con-
tinuous, there being only a shallow depression dorsally and a prominence ventrally.
Collar setae (fig. 37. 2. i, j) very short and stout and set in a slanting double row.
The two lines of setae are difficult to distinguish as they are closely packed but the
blades diiTer slightly, one row having rather longer blades about two times as long
as broad. Notosetae of setigers 2-8 include (a) winged capillaries with normal
tapered blades (b) paleae with rounded blades and no tip at all (fig. 37. 2. 1). Thor-
acic neurosetae include a row of pick-axe setae with smooth symmetrical blades
(fig. 37. 2. m) and a row of avicular uncini (fig. 37. 2. n) with finely striated crests
and very broad tails. Abdominal notosetae similar to the avicular uncini of the
thorax. Abdominal neurosetae of two types : (a) very slender winged capillaries
POLYCHAETA OF SOUTHERN AKRICA
Fig. 37.2. Hypsicomus capeiisis. (a) /Vnterior end. (b) Thoracic palea. (c) Pick-axe seta.
(d) Collar seta, (e) Notoscta of sctigcr 3. (f) Thoracic uncinus. (g) Abdominal palac.
(h) Abdominal capillary. Hypsicomus phaeotaenia. (i, j) Profile and face view of collar seta.
(k) Dorsal view of collar, (l) Thoracic palea. (m) Pick-axe seta, (n) Thoracic uncinus.
Sabella penicilhis. (o) Anterior end and part of mud tube, (p) Dorsal view of gill base and
collar, (q) Thoracic capillary, (r) Thoracic uncinus. (s) Pick-axe seta. Sabella fusca.
(t) Dorsal view of collar region, iv) Thoracic uncinus. (v) Pick-axe seta.
SABELLIDAE 763
with long narrow blades (b) stout paleae with oval blades from the rounded end
of which a slender tip arises very abruptly.
Type locality : Ceylon.
Records: Madagascar (i).
Distribution : Western Africa (Senegal (s) and Sao Thome (s) ) ; Mediterranean,
Red Sea (s) ; tropical Indo-wcst-Pacific from the Persian Gulf (s) and Madagascar
(i) to W. Australia, New Caledonia (i) and Japan.
SABELLA Linnaeus, 1 767
Body elongate, branchial lobes not spiral. Radioles not united by a web and
without external stylodcs or subterminal eyes but may have rows of eye-spots
externally. Collar widely separated dorsally, each half notched back to form a
dorsal lobe and a larger lateral one. Collar setae are winged capillaries in a compact
bundle. Notosetae of setigers 2-8 include winged capillaries of two lengths but no
spatulate setae or paleae. Thoracic neurosetae include a row of pick-axe setae
and a row of avicular uncini. Abdomen of numerous segments bearing avicular
uncini dorsally and winged capillaries ventrally.
Type SPECIES : Sabella penicillus hinnaeus, 1767.
Key to Specles
I Base of gills swollen but setigers 1-4 not glandular dorsally . . . S.penicillus
— Base of gills and setigers 1-4 swollen and glandular dorsally .... S./usca
Sabella penicillus Linnaeus, 1 767
(fig. 37.2.0-s)
Sabella penicillus hinnaeus, 1767: 1269.
Sabella pavonina Savigny, 1818: 79; Fauvel, 1927: 298, fig. 102 a-1.
Body encased in a slender mud tube, up to 250 mm. long and pale apart from
the gills wliich are banded widi purple (fig. 37.2.0). Brancliial lobes each with 10-40
radioles which are long and slender with short tips. Branchial base (fig. 37. 2. p)
without glandular cushions but with annular rings when contracted. Collar widely
separated dorsally and reflected back dorso-latcrally to form four lobes of which the
larger lateral pair end in tliickened lappets ventrally. Thoracic notosetae are
winged capillaries of two Icngdis, eidner longer and narrower or shorter and broader
(fig. 37. 2. q). Thoracic neurosetae include a row of pick-axe setae with broad
symmetrical blades ending in pointed dps (fig. 37.2.S) and a row of avicular uncini
(fig. 37. 2. r) each with a cap formed of 30 or more minutely denticulate ridges above
the rostrum and a short tail.
Type locality : Southern Europe.
Records: Cape (from 30/15/d to 34/18/s and 33/27/s).
Distribution : N. Atlantic from Scotland (i, s, d) to Morocco (s, d) and Senegal
(s) ; Mediterranean (s).
764 POLYCHAETA OF SOUTHERN AFRICA
Sabella ftisca Grube, 1870
(fig- 37-2-t-v)
Sabtilafuica Grube, 1870: 516; Gravicr, 1908; 71, figs. 243-245, text-fig. 410.
Sabella porifera Grube, 1878 : 252, pi. 14 fig. 3 ; Fauvel, 1930 : 260 ; F"auvel, 1953 : 439, fig. 323 a-f.
A large stout species with the gills often banded in yellow and brown. Radiolcs
united by a web for about a quarter of their length. Collar (fig. 37. 2. t) low, widely
separated dorsally and attached back to sedger 2, notched laterally with a pair of
stout lappets ventrally. A dorsal glandular area extends from the base of the gills
to the third or fourth setigcr ; it is often divided medially or trans\ersely and is con-
tinuous with bands on the inside of the collar which curves round to the \cntral
surface. This ventral area ma\' be separate according to some workers. Thoracic
notosetae are the usual winged capillaries. Thoracic neurosetae include a row of
pick-axe setae witli long symmetrical blades (fig. 37. 2. \) and a row of avicular
uncini with 20 obliciue rows of striations on the crest and long broad shafts
(fig- 37-'^-uj-
Type locality : Red .Sea.
Records: Mocambique (23'35/i).
Distribution' : Red Sea (s) ; Zanzibar (i) ; Andaman Is. ; C'.eylon ; \. Australia.
POTAMILLA Malmgrcn, 1866
Branchial lobes not spiral, dorsal radiolcs usually with rows of external eye-spots
but none of the radiolcs bear external stylodes. A pair of palps. Collar well
developed, each half notched dorsally and not fused ventrally. Notosetae of setigcr i
grouped in a compact bundle. Notosetae of setigers 2-8 of two kinds, (a) winged
capillaries and (b) palaea. Thoracic neurosetae include a row of pick-axe setae
and a row of a\icular uncini. Abdomen with a\icular uncini in the notopodia
and winged capillaries in the ncuropodia.
Type spec;ies : Sabella neglecla Sars, 1851.
Key to Species
1 Dorsal radicles with rows of eye-spots. fPick-axe setae with broad, tinstriatcd blades)
P, renifonnis
- Xo external eye-spots on branchial radioles ........ 2
2 C^ollar slanting forward like a scoop below the radiolcs. Pick-axe setae with plain blades
P. linguicoUaris
- Collar straight. Pick-axe setae with small striated blades . . . .P. torelli
Potamilla reniformis (Miiller, 1771)
(fig. 37-3-a-fj
Amjihilrile renifiirmii O. F. Mullcr, 1771: 194.
Sabella reni/oriais: Leuckart, 1849: 183, pi. 3 fig. 8.
Potamilla rfni/unnis : Fauvel, 1927; 309, fig. 107 a-1.
Tube (fig. 37.3.3) horny \\ith the free end flattened, sandy and partly rolled up.
Body (fig. 37.3.13) rather slender, up to 50 mm. long. Some dorsal radioles have one
SABELLIDAE 765
to four external eye-spots about half way along. A pair of large tapered palps.
Collar (fig. 37. 3. d) divided dorsally and each half notched back to form smaller
dorsal and larger lateral lobes with a straight edge. Setiger i with winged capill-
aries only ; setigers 2-8 with both winged capillaries and paleae (fig. 37.3.C) with
broad wings and short pointed tips. Thoracic neurosetae include a row of pick-axe
setae (fig. 37. 3. f) with smooth, asymmetrically pear-shaped blades having long
tapered tips and posteriorly, a row of avicular uncini (fig. 37. 3. e) with fairly broad
shafts and finely striated crests.
Type locality : Iceland.
Records: South West Africa (28/16/s) ; Cape (from 31/17/d and 34/18/i, s to
32/28/i, s) ; Natal (30/30/s and 29/3 i/i) ; Mocambique (26/32/i, 24/35/d).
Distribution : Arctic ; N. Atlantic from New England to North Carolina (s) ;
Scotland (s) and the English Channel (i) to the Cape Verde Islands (i) ; Morocco
(s, d) and Senegal (s) ; Mediterranean ; N. Pacific from Japan to Behring Sea.
Potamilla linguicollaris Day, 1961
(fig- 37-3-g-k)
Potamilla linguicollaris Day, 1961 : 539, fig. 14 d-1.
A slender species up to 15 mm. long with a fragile tube. Branchial lobes each
with six radioles which lack eye-spots and have fairly long tips. Palps slender.
Collar (fig. 37. 3. g) widely gaping dorsally ; each half is notched back to form a
small rounded dorso-lateral lobe and a large lateral lobe which slants forward below
the gills to meet its fellow, the whole collar resembling a long scoop supporting the
bases of the gills. Body with up to 12 thoracic and numerous abdominal segments.
Pygidium with a pair of small oval lobes without eye-spots. Collar setae are small
winged capillaries. Notosetae of setigers 2-8 include six winged capillaries and six
spear-shaped paleae (fig. 37. 3. k). Neurosetae include a row of pick-axe setae (fig.
37. 3. h, i) with plain, symmetrical pear-shaped blades ending in fine tips and a
row of long-shafted avicular imcini (fig. 37.3.J) with coarsely serrated crests. Ab-
dominal uncini are similar to those on the thorax and the neurosetae are three
broad-bladed paleae. All setae are symmetrical.
Type locality : Agulhas Bank, South Africa.
Records: Cape (33/17/s, 34;'2i;'s, 34/24/d).
Distribution : Endemic.
766
POLVCHAETA OF SOUTHERN AFRICA
Fig. 37.3. Potamilla rcniformls. (a) Tubes, (b) Entire worm (three times life size), (c)
Thoracic palea. (d) Dorsal view of collar, (e) Thoracic uncinus. (f) Pick-axe seta.
Potamilla linguicollaris. (g) Collar region, (h, i) Pick-axe seta, (j) Thoracic uncinus.
(k) Thoracic palea. Pulamilla torelli. (l) Hastate thoracic palea. (m) Thoracic uncinus.
(n, o) Pick-axe seta, (p) Collar region.
Potamilla torelli Malingren,
(fig. 37.3.1-p)
Sabdla {Polamilla) lorelli Malmgrcn, 1866: 402.
Potamilla torelli : Fauvcl, 1927: 310, fig. 107 m-s; Day, 1955
1866
447, fig. 8 d-f.
Tuby horny with tlie free end encrusted with sand, flattened and partly rolled up.
Body up to 60 mm. long with well marked glandular pads ventrally. Branchial
lobes each with 10-15 radioles which lack eye-spots but have long flanged tips. A
SABELLIDAE 767
pair of large tapered palps. Collar (fig. 37. 3. p) divided dorsally and each half
notched back to form a narrow dorsal lobe and a large lateral lobe which ends in
a rounded lappet overlapping its fellow in the mid-ventral Une. Collar setae arc
normal winged capillaries. Notosetae of setigers 2-8 include winged capillaries
and hastate paleae (fig. 37. 3. 1). Thoracic neurosetae include a row of pick-axe
setae (fig. 37. 3. n, o) with small striate blades on which a long blade-like tip is set
edgeways and a row of avicular uncini (fig. 37. 3. m) with coarsely serrated crests
and long shafts.
Type locality : Iceland.
Records: Cape (from 34/18/s and 34/19/i, s to 34/25/s) ; Natal (29/3 1 /i) ;
Mocambique (23/35/s).
Distribution : North Atlantic from Iceland and the English Channel (s) to
Madeira and Morocco (s, d) ; Mediterranean (s) ; Japan.
BRANCHIOMMA KoUiker, 1858
(= DASrCHONE Sars, 1862)
Body usually stout. Branchial lobes sometimes spiral. Radioles mainly free from
one another and possess external processes (stylodes) and double rows of eye-spots.
A well developed collar widely separated dorsally. Thoracic notosetae are winged
capillaries of two lengths but paleae are absent. Thoracic neurosetae are avicular
uncini and pick-axe setae are absent. Abdominal segments numerous, with avicular
uncini dorsally and winged capillaries ventraUy.
Type species : Amphitrite bombyx Dalyell, 1853.
Key to Species
1 Stylodes broad (fig. 37. 3. b, g). Uncini with very numerous (over 40) poorly marked
denticles surmounting the rostrum (fig. 37. 3. d, f) ...... . 2
- Stylodes slender or minute (fig. 37.30, p). Uncini with less than 40 denticles above the
rostrum (fig. 37. 3. k, n, q) 3
2 Branchial lobes spiral with about five whorls of radioles (fig. 37.3.3) . . B. natalensis
- Branchial lobes not spiral . . . . . . . . . . B. violacea
3 Stylodes separate and slender (fig. 37. 3. p) ........ 4
- Stylodes reduced to notches on the sides of the radiolar flange (37. 3. j) . B. serratibranchis
4 Uncini with few teeth arranged in two arcs above the rostrum. Abdomen flecked with
dark pigment (37. 3. m) ........ B. nigromaculata
- Uncini with numerous denticles arranged in five to si.x rows above the rostrum. Colour
uniform ............. B. capensis
Branchiomma natalensis (Kinberg, 1867)
(fig. 374-a-d)
Sabella natalensU Kinherg, 1867: 353.
Dasychone natalensis : Day, 1 955 : 444, fig. 7 g-j.
Body stout and brownish shading to purple anteriorly. Collar (fig. 37.4.3) widely
separated dorsally, notched laterally and ending in rounded lappets ventrally.
768 POLYCHAF, lA OF SOUTHERN AFRICA
Branchial lobes spiral with four to six whorls of radiolcs of decreasing length.
Radiolar eye-spots present. Stylodes (fig. 37. 4. b) broad and extend back as llanges
on tJic sides of the radiolcs ; they increase in size distally. Thoracic uncini (fig.
37. 4. c, d) have straight tails and 20 or more minute denticular ridges abo\e the
rostrum.
Type locality : C^apc of Good Hope.
Records: South West Africa (26/15/i) ; Clapc (from 29/16/i to 33/18/i, s and
34/i8;i).
Distribution : Endemic.
Branchionima violacea (Schmarda, 1861)
(fig. 37.4.c-i)
Salirllti riolinea Schmarda, 1861 : 34, pi. 22 fig. 187.
Da\n'lmne iiolacea : Day, 1955: 443. fig. 7 k-m.
Body (fig. 37. 4. h) up to 100 mm. long; colour brownish shading to purple
anteriorly. Branchial lobes not spiral and bear not more than one whorl of radioles.
Stylodes (fig. 37.4. g) broad and truncate and increase in size distally. Radiolar
eye-spots present. Collar widely separated dorsally, notched laterally and ending in
rounded lappets ventrally (fig. 37.4.1). Thoracic uncini (fig. 37. 4. e, f) have short
broad tails and 12-14 ''ows of indistinct denticles above the rostrum.
Type locality : Clape of Good Hope.
Records: South West .-Xfrica (22/14/1, s and 26/15/i, s) ; Cape (from 29/16/i to
34/18/i, s and 34/23/e, i to 32/28/i) ; Natal (30/30/i).
Distribution : Endemic.
Branchiomnia serratibranchis (Grube, 1878)
(lis- 37-4-J-l)
Sabelhi (DajTclione) smalibraiichis Griitje, 1878: 262, pi. 14 fig. 7.
Da\ychone serratibranchis : Day, 1951 : 64.
Body uniformly pale or brownish and up to 70 mm. long. Branchial lobes semi-
circular, not spiral. Xo radiolar eye-spots seen. Stylodes (fig. 37.4.J) short and
appear as small triangular serrations or steps on the sides of the radiolar liange.
Ciollar low, the two sides well separated dorsally and rertected back dorso-laterally
with smooth lateral lobes and small ventral lappets. Thoracic uncini (fig. 37.4. k, I)
\\\\\\ narrow nei ks, four to five arcs of rather coarse teeth above the rostrum and
broad tajjered tails.
Type locality: Philippine Islands.
Records: Xatal (29 31/i) ; Mocambique (26/32/i, s).
Distribuiion : Indo-west-Pacific (from India to the Philippine Islands, .Japan
■uid .\ustrali.i to .New Zealand).
SABELLIDAE
7C9
Fig. 37.4. Branchiomma natalemis. (a) Anterior end with the gil
show the spiral axis, (b) Stylodes on distal part of radiole.
Branchiomma vhlacea, (e, f) Thoracic uncinus. (o) Stylodes
(h) Entire worm (twice life size), (i) Ventral view of collar,
(j) Stylodes on distal part of radiole. (k, i.) Thoracic uncinus.
(m) Dorsal view of thorax showing collar and pigmentation,
(p) Stylodes on distal part of radiole. Bramhiumma capemis.
(s) Stylodes on distal part of radiole.
s cut short on one side to
(c, d) Thoracic tmcinus.
on distal part of radiole.
Branchiomma serratibranchis,
Branchiomma nigromaculala.
(n, o) Thoracic uncinus.
(q, r) Thoracic uncinus.
770 rOLVC.HAETA OF SOUTHERN AFRICA
Branchiomnta nigromaculata (Baird, 1865)
(fig. 37.4.m-p)
Sabella nigromaculula Tiiilrd, 1865a: 159.
Dasychone nigromaculala : Day, 1955 : 445, fig. 7 n-r.
Length up to 70 mm. Body irregularly flecked with dark pigment (fig. 37. 3. m).
Branchial lobes semi-circular, not spiral and radiolcs arranged in a single whorl.
A row of long stylodes at the base of the radicles, later stylodes (fig. 37. 4. p) slender
and as long as the radiolc is broad. Radiolcs without marginal flanges. Collar
broadly separated dorsally, reflected back dorso-laterally and with smooth lateral
lobes which end in well marked ventral lappets. Thoracic uncini (fig. 37.4. n, o)
avicular with short tails bent downwards and with two arcs of teeth above the
rostrum, the first with four to six and the second with six to eight teeth.
Type locality : St. \'incent, \Vest Indies.
Records: Cape (from 34/18/1, s to 33/25/s) ; Natal (30/30/i and 29/31/i);
Mocambicjue (26/3'2/i, s) ; Madagascar (s).
Distribution : Tropical and sub-tropical Atlantic (from North Carolina (s), tJie
Gulf of Mexico (i) and West Indies to Cape Verde Is.) ; Red Sea (i) ; tropical Indian
Ocean (i, s) ; Pacific (Gambier and Japan).
Branchiomma capensis (Mcintosh, 1885)
(fig. 37.4.q-s)
Dasychone cajiensh Mcintosh, 1883 : 506, pi. 54 fig. i, pi. 31A figs. 9-1 1, pi. 39 fig. 8 ; Day, 1961 :
538, fig- 14 a-c-
Body slender, up to 50 mm. long and uniformly pale apait from interramal
eye-spots. Branchial lobes semi-circular, not spiral and bear not more than one
flattened whorl of radiolcs. Collar widely separated dorsally, low and without
lateral notches. Stylodes (fig. 37.4.5) small, strap-like and one to two times as long
as the width of the radicle. Radicles without marginal flanges. Collar divided
dorsally, reflected back dorso-laterally, and forming small lappets ventrally. Thoracic
uncini (fig. 37.4.9, r) h.ave six to eight rows of denticles above the rostrum.
Type locality : 98 fathoms off Cape of Good Hope.
Records: Cape (from 31/16/d to 34/18/s and 35/18/d to 34/25/s).
Distribution : Endemic.
S ABELL AST ARTE Kioyer, 1856
Body stout, often large. Branchial lobes sometimes spiral, with more than one
whorl of radicles. Radicles with or without a double row of eyc-spcts but never
with stylodes. Cellar widely separated dorsally. Thoracic notosetac are winged
capillaries of two lengths but spatulate setae are absent. Thoracic neurosctae arc a
SABELLIDAE 771
single row of avicular uncini, there being no pick-axe setae. Abdominal segments
numerous with dorsal avicular uncini and ventral winged capillaries.
Type SPECIES : Euralo sanctijosephi Gravier, 1906 (= Sabella indica Savigny, 181 8,
non Linnaeus, 1788).
Key to Species
I Branchial lobes spiral with about five whorls of radicles (fig. 37. 4. a). Outer radioles with
eye-spots and marginal flanges. Body uniform in colour .... 5. longa
- Branchial lobes not spiral. Eye-spots absent. Body flecked with dark pigment S. sanctijosephi
Sabellastarte longa (Kinberg, 1867)
(fig. 37-5-a-e)
Sabella longa Kinberg, 1867 : 352.
Sabellastarte longa : Johansson, 1925 : 10, figs. 3, 5, 6, 7 ; Day, 1951 : 62.
Dasychone odhneri Fauvc\, 1919: 24, pi. i figs. 10-14.
A large species reaching 150 mm. by 15 mm. Body (fig. 37. 4. a) shading from
brown to purplish anteriorly, and the gills uniformly orange brown or barred.
Branchial lobes each with a central axis bearing four to six whorls of radioles. Outer
whorl of radioles (fig. 37.5.b) with double rows of eye-spots and lateral flanges
formed by a continuation of the basal web. No stylodes. Collar widely separated
dorsally, not notched laterally and forming a pair of stout triangular lappets vcntrally
(fig. 37. 5. c). Thoracic capillaries of two lengths, the shorter ones having broader
wings than the longer ones. Thoracic uncini (fig. 37. 5. d, e) with finely denticulate
crests, rather narrow necks and fairly long tails.
Type locality : Durban, South Africa.
Records: Cape (from 34/18/i to 34/22/i, s and 32/28/i) ; Natal (31/29/i to
27/32/i) ; Mocambique 26/32/1.
Distribution : Madagascar (i) .
Sabellastarte sanctijosephi (Gravier, 1906)
(fig- 37-5-f-')
Eurato sancti-Josephi Gravier, 1906 : 105, pi. 7 figs. 281-283.
Sabella indica Savigny, 1820: 77 (non .S. indica Linnaeus, 1788).
Sabellastarte indica : Day, 1951 : 63.
Length up to 80 mm. by 1 1 mm. Body irregularly flecked with purple and gills
banded. Branchial lobes not spiral. Individual radioles (fig. 37. 5. f) without any
trace of external stylodes or eye-spots and the lateral flanges are reduced to faint
ridges with a median groove between them. Long palps. Collar (fig. 37. 5. g)
divided dorsally, and reflected back dorso-laterally to form a pair of small fonvardly
projecting dorsal lobes and a pair of large lateral lobes which terminate as triangular
772
POI.VCIIIAK TA OK SOUTHKRX ATRKIA
Fig. 37.'). Sabeliaslarte longa. (a) Entire worm with radiolos removed from axis on the right
side (life size), (b) Part of radiole. (c) N'entral view of collar, (d, f.) Thoracic uncinus.
Sabellastarte sancfijosephi. (f) Part nf radiole. [c.) Dorsal view of collar, (h, i) Thoracic
uncinus. Myxicola infiimlibulum. i \) Dorsal view of anterior end. (k) Ventral view of collar
region, (i.) Abdominal unciiuis. (m. n) 'I horarir vnu inus. (o) Thoracic winged capillary.
SABELLIDAF, 773
lappets ventrally. Thoracic notosetae are winged capillaries of two lengths. Thor-
acic neurosetae are a single row of avicular uncini each with an arched neck and
the rostrum surmounted by a prominent striated crest (fig. 37. 5. h, i).
Type locality : Red Sea.
Records: Natal (29/31/i and 28/32/i) ; Mocambique (26/32/i).
Distribution: Western Africa (Mauritania (i), Senegal (i, s)) ; Red Sea (i) ;
tropical Indo-west -Pacific from India (i, s) to Japan and New Caledonia.
MYXICOLA Koch (in) Renicr, 1847
Branchial lobes equal, not spiral. Radioles lack external stylodes and eye-spots
and are united for most of their length by a web. Collar indistinct. Thoracic
notosetae are winged capillaries and the neurosetae are minute, long-shafted hooks.
Abdominal segments numerous with capillaries ventrally and an almost complete
semi-circle of avicular hooks dorsally. Tube mucilaginous.
Type species : Terebella infundibulum Renier, 1 804.
Myxicola infundibulum (Renicr, 1804)
(fig- 37-5-J-o)
Terebella infundibulum Kenier, 1804: p. xiii.
Myxicola infundibulum : Fauvel, 1927: 342, fig. 119 a-i.
Body markedly tapered with a pair of large branchial lobes each bearing 20-40
radioles which are united by a web almost to their bare tips (fig. 37.5.J). CoUar
without a flanged margin but forming two low, closely apposed dorsal lobes, a
stout lateral ridge with a notch in front of the notosetae and widening ventrally to
form a median triangular projection between the branchial lobes (fig. 37. 5. k). An
otocyst in the first segment. Thoracic notosetae are numerous winged capillaries
(fig. 37.5.0). Thoracic neurosetae are long-shafted hooks each with a minute
denticle above the rostrum (fig. 37. 5. m, n). On the abdomen the tiny uncini form
an almost continuous arc across the dorsum. They are avicular with a single tooth
above the rostrum (fig. 37. 4. 1). Abdominal neurosetae are winged capillaries like
the thoracic notosetae. Pygidium with ocular specks. Tube muchlaginous.
Type locality : Adriatic.
Records: South West Africa (22/14/i and 26/15/i) ; Cape (34/18/i, s and
33/25/e).
Distribution : .\rctic ; N. Atlantic from Greenland (s, d) to Scotland (s) and
the English Channel (i, s) to Mauretania (s) ; Mediterranean ; N. Pacific from
the Behring Sea to Japan and Alaska to California.
774 POI.YCIIAETA OF SOUTHERN AFRICA
EUCHONE MalniRrcn, 1866
Small SabcUids with eight thoracic and numerous abdominal segments. Each
branchial lobe has several radioles which arc united for most of their length by a
web. A few ventral filaments lack pinnules. A pair of palps. Otocysts present in the
collar segment. Collar well developed and usually continuous ventrally. Thoracic
notosetae include both winged capillaries and paleae. Thoracic ncurosetae as
long-shafted hooks. The last few abdominal segments are flanged laterally to form
a spoon-shaped cavity. Abdominal notosetae are avicular imcini without tails.
Abdominal ncurosetae are narrow-winged capillaries. Pygidial eye-spots sometimes
present.
Type SPECIES : Sabella analis Krbycr, 1856.
Key to Species
I Collar shallowly notched \'entraily. Anal concavity extending over about five setigers E. rosea
- Collar split ventrally. Anal concavity extending o\cr about eight seligers . . E. capensis
Etichone rosea Langerhans, 1884
(fig. 37.6.a-i)
Etichone rosea Langerhoms, 1884: 271, pi. 16 fig. 35; Fauvel, 1927: 340, fig. 1 18 a-1.
Body (fig. 37. 6. a) up to 12 mm. long, shghtly tapered, with 20-26 segments.
Branchial lobes each with five radioles united by a web for more than half their
length and ending in long naked tips. About two pairs of ventral filaments which
are small, rolled inwards and lack pinnules. Two long palps. Collar widely divided
dorsally, well developed laterally and shallowly notched ventrally where it is attached
to the lips (fig. 37. 6. b). A pair of otocysts and a pair of internal eyes in the collar
segment. Thoracic notosetae include winged capillaries (fig. 37.6.11) and spear-
shaped paleae (fig. 37. 6. g). Thoracic ncurosetae are long-shafted hooks (fig. 37. 6. c)
with numerous denticles above the main fang. Abdomen with 12-18 segments and
a ventral concavity at the posterior end (fig. 37.6. d) with lateral flanges for about
five to six setigers. Abdominal notosetae are quadrangular uncini with several
rows of denticles above the main fang (fig. 37.6. e, f). Abdominal ncurosetae are
narrow-winged capillaries (fig. 37.6.1). Pygidium with two eyes but without a
posterior prolongation.
Type locality: Madeira Island.
Records: Cape (from 32/17/d to 34/18/s).
Distribution: S.\V. Ireland (s) ; Madeira (i) ; Ivory Coast (s).
SABELLIDAE
775
Fig. 37.6. Euchone rosea,
(c) Thoracic hook,
(c) Thoracic palea.
capensis. (j) Ventral
(m) Thoracic hook.
(a) Entire worm (ten times Ufe size), (b) Ventral view of collar.
(d) Ventral view of posterior end. (e, f) Abdominal uncinus.
(h) Thoracic winged capillary, (i) Abdominal capillary. Euchone
view of collar, (k) Thoracic palea. (l) Minute thoracic seta.
(n, o) Abdominal uncinus. Chone jilicaiidala. (p) Thoracic palea.
((i) Dorsal view of posterior end. (r, s) .■\bdominal uncinus. (t) Anterior end. (u) Minute
thoracic seta, (v) Head of thoracic hook, (w) Thoracic winged capillary.
776 P(T|At;iI.\I-. lA OV SOU THF.RX AFRICA
Euchone capensis Day, ii)6i
(fig. 37-*^-J-o)
Euchom cajunsi.s Day, 1961 : 540. fig. 14 m-t.
Length up to 20 nun. for 40 segments. Branchial lobes each with nine radioles
bearing pinnules and two to three elongated pinnules ventrally. Radioles all
imited by a web for two-thirds their length. C'ollar divided dorsally and notched to
form a pair of small dorsal lobes and a pair of large lateral lobes ; the latter are slit
ventrally and soldered onto the ventral lips (fig. 37.6.J). A pair of eyes and a pair of
otocysts inside the collar segment. Thorax with eight setigers. Notosetae of sctigers
two to eight of three types : (a) four winged capillaries ; (b) five paleac with broad
blades and long tips (fig. 37.6.!;) and (c) three setae with minute blades which just
project from tlie surface (fig. 37. 6.1). Thoracic neurosetae are long-shafted hooks
(fig. 37. 6. m) with arcs of numerous small denticles above the main fang. Abdomen
vvith 32 setigers. Abdominal neurosetae are slender capillaries. The notosetae are
quadrangular uncini (fig. 37. 6. n, o) with five to six close-set rows of teeth above the
main fang. The last eight segments are short and ILmged to form a spoon-shaped
conca\ity. Pygidium conical and without eyes.
Type loc:ality : Agulhas Bank, South Africa.
Records: Clape (3'2/'i6/d and 34/2 i/s).
Distribution' : Only two records.
CHONE Krbycr, 1856
Rather small Sabellids with eight thoracic and numerous al^dominal segments.
Branchial lobes each with several radioles of which a few \entral ones lack pinnules.
The rest are united by a web for most of their length. Two palps. A well developed
collar. Thoracic notosetae include winged capillaries and paleac. Thoracic neuro-
setae arc long-shafted hooks. Pygidiuin sometimes with eye-spots or a terminal
appendage. Abdominal notosetae are quadrangular uncini with numerous denticles
while the neurosetae are winged capillaries.
Type species: Chonc iiijundibulifornns Kroyer, 1B56.
Key to Species
1 Pygidium uilli a terminal appendage (fig. ;;7.6.q) ..... C. filicaudata
— No pygidial appendage ........... 2
2 Collar with a scalloped margin (fig. 37.7.;!) ...... C. collaris
- Clollar with a -smootli margin ........ C letterstedti
Chone filicaudata Southern, n)i4
(lig. 37.G.P-W)
(.hour fihcau/lata .Southern, i')i4 : 141, pi. 14-13 tig. 32 ; l''au\'el, 1927 ; 337, fig. 1 17 a-k.
Body short and rather stout, 5-1 1 mm. long with 26-30 segments. Each branchial
lobe with six to ten radioles plus two elongated v'cntral filaments. All radioles imited
for half their length and thereafter Hanged. Two short p.il]5s ,ind a bifid \ential lip.
SABELLIDAE 777
Collar (fig. 37. 6. t) well developed with small apposed dorsal lobes, smooth lateral
lobes which are not notched ventrally. Eyes and otocysts doubtful. Thoracic
notosetae of three types : (a) winged capillaries (fig. 37. 6. w) (b) paleae with tapered
blades (fig. 37. 6. p) and (c) short fine setae with tapered tips but no blades (fig.
37. 6. u). Thoracic neurosetae arc long-shafted hooks with the main fang surmounted
by two teeth, numerous denticles and "une limbe etroite" (fig. 37.6.V). Abdominal
notosetae are quadrangular uncini (fig. 37. 6. r, s) with six close-set rows each with
five to six teeth above the main fang and basal prow. Neurosetae are fine-bladed
capillaries. Pygidium with a tapered terminal appendage (fig. 37. 5. q).
Type locality : Ireland.
Records: Cape (33/17/s, 34/18/s, 34/21/s) ; Natal (30/30/s).
Distribution: Ireland (s) ; North Carolina (s, d).
Chone collaris Langerhans, 1880
(fig- 37-7-a-f)
Chone collaris Langerhans, 1880 : 1 16, pi. 5 fig. 29 ; Fauvel, 1927 : 337, fig. 1 16 p-x.
Body small with about 30-40 segments. Branchial lobes each with four to eight
radiolcs united by a web for two-thirds their length, and thereafter flanged. A pair
of small ventral radioles lack pinnules. A pair of long slender palps. Collar (fig.
37. 7. a) narrowly divided dorsally, straight laterally and united ventraUy ; its
margin is scalloped. Eyes and otocysts inside the collar segment. First setiger with
winged capillaries only. Setigers 2-8 with winged capillaries (fig. 37. 7. b) and
paleae (fig. 37. 7. c) in the notopodia and long-shafted hooks in the neuropodia.
Individual hooks (fig. 37. 7. d) with numerous small teeth above the main fang when
seen in lateral view. Abdominal segments numerous. Abdominal uncini (fig.
37. 7. e, f ) are square with three to four arcs each of six teeth above the large main
fang. Neurosetae are winged capillaries. The pygidium may have eye-spots.
Type locality : Madeira Island.
Records: Mocambique (23/35/i).
Distribution: Madeira (i) ; Mediterranean (s) ; ? Persian Gulf (s).
Chone letterstedti (Kinberg, 1B67)
(fig. 37-7-g-k)
Parachonia letterstedti Kinberg, 1867 : 73, pi. 28 fig. 6 ; Johansson, 1925 : 27, figs. 9-1 1.
Body about 25 mm. long with numerous segments. Branchial lobes each with
eight long radioles united by a web for half their length, and with long slender
flattened tips. Number of ventral radioles lacking pinnules unknown. Collar low,
widely divided dorsally and reflected back dorso-laterally to form pocket-Like dorsal
lobes which reach back to setiger 3 ; on the sides there are straight lateral lobes
which are ( ?) united ventrally (fig. 37.6.g). First setiger with winged capillaries only.
Y
778
rOLVCHAETA OF SOUTHERN AFRICA
Fig. ?,"].'. Clione collaiiu (a) Anterior end. (b) Thoracic winged capillary, (c) Thoracic
hastate seta, (d) Thoracic hook, (e, f) Abdominal unciniis. Chom Ictlcrstedli. (a) Ventral
view of anterior end (from Kinberg). (h) Winged thoracic capillary, (i) Thoracic palca.
(j) Thoracic hook, (k) Abdominal uncinus (all from Johansson). Jasmmeira elegans.
(l) Entire worm (four times life size), (m) Ventral view of collar, (n) Ventral view of
posterior end. (o) Thoracic hooks, (p, q) Abdominal uncinus. (r) Thoracic palea.
(s) Abdominal capillary.
SABELLIDAE 779
Setigers 2-8 with (a) winged capillaries (fig. 37.7.11), (b) paleae (fig. 37. 7. i)
and (c) fine setae in the notopodia. Neuropodia with long-shafted hooks (fig. 37.7.J).
Abdominal uncini (fig. 37. 7. k) rhomboidal with six rows each with five to ten
teeth above the large main fang. Anal segments without special features.
Type locality : "Cape of Good Hope".
Records: Cape (33/18/i).
Distribution : A single record.
JASMINEIRA Langcrhans, 1880
Body small and elongate. Branchial lobes not spiral. Radioles without eyes or
external stylodes, and the web between them is not developed. A few ventral
radioles may lack pinnules. A pair of palps. A well developed collar. Otocysts and
eyes present in the first segment. Thoracic notosetae include winged capillaries and
paleae. Thoracic neurosetac are long-shafted hooks. Abdominal segments numerous
with avicular uncini dorsally and capillaries ventrally.
Type species: Jasmineira caudata Langcrhans, 1880.
Key to .Species
Three radioles to each branchial lobe. Pygidium wit
Eight or more radioles to each branchial lobe. Pygid
Radioles with flattened, lamellate tips
Radioles with normal tapering tips.
Pygidium with a slender caudal appendage
Pygidium without a caudal appendage
1 large glandular lappets . J. analis^
um without large glandular lappets 2
. J. candela*
J. caudata
J. elegans
Jasmineira ( ?) analis Ehlers, 1 908
Jasmineira analis Ehlers, igo8a: 48.
Length 3-5 mm. with eight thoracic and 7-15 abdominal segments. Branchial
lobes each with three free radioles which bear double rows of long pinnules and
end in long naked tips. Collar well developed dorsally, deeply notched laterally
and with ventral triangular lobes which are reflected back. A pair of eyes and a
pair of otocysts in the first segment. Thoracic notoseate of two lengths, the shorter
ones hastate but not quite paleae and the longer ones normal winged capillaries.
Thoracic neurosetae in a single row of about 12, probably avicular with serrated
crests and long, sharply bent shafts. No pickaxe setae. Abdominal notosetae are
uncini similar to those of the thorax but with shorter necks and tails. Abdominal
neurosetae are a few, narrow-winged capillaries. Pygidium with large rounded,
glandular lappets.
Type locality : Luderitzbucht, South West Africa.
Records : South West Africa (26/15/i).
Distribution : Only a single record.
tThe affinity oij. analis is doubtful, this species should probably be referred to a different genus.
yV.o roiA CIIIAI, I A Ol S()L"1HKR\ AIRICA
Jasniineira caudata Langcrhans, 1880
Jnsmimira ciwilata Langrrlians, 1880: ii.j, ]il. -, lig. y2\ I'.uivcl, 1927: 372, liR. 115 g-k.
Body about 5-7 mm. long witli L'f, -'8 segments. About seven or eight ratlioles
in each branchial lobe; each radiole free and without lateral flanges, hut with
long pinnules and a breaking plane near the base. A pair of long, slender palps.
C'ollar well dc\eloped, the two halves overlapping dorsally and united to the lips
ventrally ; margin straight, not obliciue. Two elongate red eyes visible in fresh
specimens through the base of the collar. Two otoliths. Thorax of eight setigers ;
first setiger with only winged capillaries, the remaining seven with both winged
capillaries and spathulate setae in the notopodium and four to five long-shafted
hooks in the neuropodiuni. Abdomen tapered with about 20 setigers and a pygidium
with a long, slender caudal appendage. The winged capillaries of the thorax have
blades of normal width but the spathulate setae have blades which widen distally
and then suddenly taper to slender tips ; they might almost be called paleae. The
thoracic hooks have a prominent rostrum surmounted by about four arcs of long
denticles. Abdominal capillaries have long slender blades ; the hooks are z-shaped
or avicular, each with a long rostrum surmounted by close set arcs of denticles
(about 13 in all) and a long manubrium which bends back abruptly to form a lightly
chitiniscd base.
Type locality : Madeira.
Records: Mocambicpic (24/35/d).
DlSTRiBrxiON' : Warm North Atlantic from Madeira (s) to Ireland (s).
Jasniineira elegans Saint- Joseph, 1894
(fis- 37-7-M '
JaMnim-ira rli'gam Saint-Joseph, 1894: 316, pi. 12 figs 337-348; Fauvel, 1927: 330, fig. 114 k-r ;
Day, 1961 : 542.
Body (fig. :^7.7.1) U]:) to 20 mm. long with 30-40 setigers. Colour greyish green
when fresh with a brilliant narrow line encircling the body between setigers 2-3.
Branchial lobes each witli 8-20 radioles. Web absent and the radioles have a
breaking plane at the base. Two conical dorsal lips between the gills, two slender
palps and two long radioles without pinnules ventrally (fig. 37. 7. m). Collar well
developed, inflected inwards doisally and overlapping in the mid-ventral line. Two
internal eyes and a pair of otocysls in the collar segment. Setigers 2-8 with large
winged capillaries and short-lipped jjaleae (fig. 37. 7. r) in the nntopodia and a row
of long-shafted hooks with a crest of long denticles and small winged capillaries in
the- neuropodia (fig. 37.7.0). Abdominal segments numerous with long-necked
horiks (fig. 37. 7. p, q) capped with 12-20 denticles in the n(Jli i|30(Jia and slender
SABELLIDAE 781
winged capillaries (fig. 37.7.5) in the iicuropodia. Pygidium pointed but no pygidial
eyes or caudal appendage.
Type locality : France.
Records: Cape )32/i7/d, 33/17/s, 34/18/s) ; Natal (30/30/s and 29/31/d).
Distribution: N. Atlantic from Scotland (s), Ireland and the English Channel
(s) to Senegal (i, s) ; Angola (s).
FABRICIOLA Friedrich, 1939
Small Sabcllids usually 2-3 mm. long with eight thoracic and three abdominal
segments. Each branchial lobe with a branchial heart, and three radioles with long
pinnules but without eyes or external stylodes. A membraneous Up and a long
filamentous palp without an internal blood-vessel on either side of the mouth.
Collar distinct dorsally and united ventrally. A pair of eyes in the collar-segment.
The first setiger bears winged capillaries only. Notosetae of setigers 2-8 are wunged
capillaries and sometimes a few spatulate setae in the middle thoracic segments.
Thoracic neurosetae are long-shafted hooks. Three abdominal segments with short-
shafted uncini in the notopodia and capillaries in the neuropdia. Pygidium with a
pair of eye-spots.
Type SPECIES : Manayunkia pacijica Anncnkova, 1934.
Fabriciola mossambica (Day, 1957)
(fig. 37-8.a-g)
Fabricia mossambica Day, 1957: 115, fig. 8 e-o; Day, 1963a; 440.
Body 2-5 mm. long with eight thoracic and three abdominal segments. Each
branchial lobe with a branchial heart and three radioles bearing about five to eight
pairs of pinnules (fig. 37.8. a). A pair of filamentous "palps". Collar rudimentary
except for a large triangular ventral lobe which is retractile. .\ pair of eyes in the
collar segment. Thoracic segments about as long as broad. Thoracic notosetae of
two lengths, the longer ones (fig. 37. 8. b) with normal blades, the shorter ones
haxing very broad, hastate blades (fig. 37. 8. c). Thoracic neurosetae are long-shafted
hooks (fig. 37. 8. d, e) each bearing a single large tooth and an arc of seven denticles
above the main fang. Abdominal notosetae are uncini (fig. 37. 8. f, g) with bent,
flattened shafts and two to three rows each with six to seven teeth. The neurosetae
are fine capillaries. Pygidium with eye-spots.
Type locality : Mocambique Island.
Records: Mocambique (23/35^1 and .Mocambique Is.); ? Cape (34/17;^,
32/17/d, 36/21/d).
Distribution : No other records.
782 POLVCHAETA OF SOUTHERN AFRICA
FABRICIA BlainviUc, i8j8
Small Sabcllids measuring about 3-4 mm. with eight thoracic and three abdom-
inal segments. Each branchial lobe with a branchial heart and three free radioles
bearing long pinnules but without eyes or external stylodes. Palps not filamentous.
Collar often reduced dorsally but united ventrally. A pair of eyes in the collar
segment. First setiger with winged capillaries only. Setigcrs 2-8 with winged
capillaries and sometimes a few spatulate setae in the notopodia ; long-shafted
hooks in the ncuropodia. Three abdominal segments with long-handled hooks in
the notopodia and fine capillaries in the ncuropodia. Pygidium with eye-spots.
Type SPECIES : Aniphkora sabella Ehrcnhcrg, 1837.
Key to Species
Collar well developed dorsally. Middle thoracic segments elongated .... 2
Collar rudimentarv' dorsally. Thoracic segments short, (.\bdominal hooks with one to
three longitudinal rows of teeth) ......... F, baiisei
.Abdominal hooks with eight longitudinal rows each with about lu teelh. .\ large species
(10 mm.) ........... .F.capensis
.•\bdominal hooks with three longitudinal rows each with si.K to seven teeth (tig. 37. 9. g).
.\ small species (3-4 mm.i ......... F.filameiitosa
Fabricia bansei Dav, 1961
(fig. 37.8.h-l)'
Fabricia bansei Day, igGi : 543, fig. 15 c-f.
Body ffig. 37-B.h) about 3 mm. long, dusky anteriorly. Branchial lobes each
with a branchial heart and three radioles. Xo elongated palps or free ventral
filaments. Collar small and indistinct dorsally but forms a triangular lobe ventrally
closely apposed to the mouth. A pair of eyes in the collar segment. Body with eight
thoracic and three abdominal segments. Thoracic notosetae of setigcrs 2-8 include
four to fi\e longer \vinged capillaries and two to three shorter paleae (fig. 37.8.])
with hastate blades and fine tips. Neurosetae are long-shafted hooks (fig. 37. 8.1)
with a single long tooth over the main fang and then an arc of several smaller teeth.
.Abdominal hooks 14 per ro\v, each with a long handle (fig. 37.8.J, k) and a small
rostrum surmounted by a single row of three to four larger teeth and a pair of
terminal teeth.
Type locvlh v : Cape Town.
Records: Cape (33/18 i and 34,18'i, s).
Distribution : Endemic.
SABELLIDAE
783
Fig. 37.8. Fahriciola mossambica. (a) Anterior end. (b) Thoracic winged capillary, (c)
Thoracic hastate seta, (d, e) Thoracic hook, (f, g) Abdominal uncinus. Fabrkia bamei.
(h) Entire worm (20 times life size), (i) Thoracic palea. (j, k) .Abdominal hook, (l)
Thoracic hook. Fabncia capensis. (m) Entire worm (10 times life size). (n) Dorso-lateral
view of collar region, (o, p) Abdominal hook. {q_) Thoracic hook, (r) Thoracic winged
capillary.
784 POI.VCHAETA OF SOUrili;K\ AFRKIA
Fabricia capensis (Monro, 11)37)
(fig. 37.8.m-r)
Oridia cnjiemis M<>nn>, 1937: 366, figs. 1-8.
Fabikia caficnsii : Day, I9i5: 447, fig. 8 g-k.
Body (fig. 37.8.111) .slender, 5-10 mm. long and encased in a long sandy tnbc.
Animals gregarious with jnvcniles in the tubes of the adults. Each branchial lobe
with a branchial heart and three radicles bearing about six pairs of long pinnules.
A pair of short dark flattened lips and a similar pair of palps. No free filamentous
processes \-entrally. Oillar (fig. 37. 8. n) well developed, rellected inwards on cither
side of the mid-dorsal groove to form a single small dorsal lobe and a pair of large
lateral lobes which are smoothly continuous ventrally. Eyes in the collar segment.
Middle thoracic segments greatly elongated, the sixth, seventh and eighth being
fi\e times longer than broad. Thoracic notosetac arc all long, slender-winged
ca]iillaries (fig. 37. 8. r). Thoracic nem-osctac are long-shafted hooks (fig. 37. 8. q)
with one to two large teeth and an arc of 12-14 denticles above the main fang.
Abdominal notosetac are long-handled hooks (fig. 37.8.0, p) with 18 rows with
eight teeth each. .Xbdominal capill.uies \ery slender. Pygidium slender witli
eye-spots.
TvPE lo(:ality : Port Elizabeth, South Africa.
Rec;ords : Clape (from 34/18/i and 34/'-!5/s to 32/28/i).
Distribution : Endemic.
Fabricia filamentosa Day, 1963
(fig. 37.9.a-g)
Fabricia tdanieulo^a Day, iqGja : 439, fig. 12 a-g.
A small thread-like species (fig. 37.9.3) about 4 mm. long by 0-2 mm. encased
in a fragile mud tube. Xo colour pattern. Each branchial lobe with three radiolcs
bearing five pairs of very long slender pinnules. .\o lilamenlous palps. C^oUar
(fig. 37. 9. b) well developed \vith a short dois.il lajipet and a smooth-edged, cup-
shaped flange which is continuous ventrally. A jjair of conspicuous eyes. Body
slender with eight thoracic and three abdominal segments. Setigers 5-8 greatly
elongated. Thoracic notosetac ffig. 37. 9. c) are four narrow-bladcd capillaries;
neurosetae are foui" long-shafted hooks (fig. 37. 9. d, e) each with an arc of [\\c to
eight teeth above the rostrum. Abdominal capillaries are three very slender capill-
aries. About If, abdominal uncini per segment, each with a long "handle" and
about six rows of three teeth (fig. 37.9.1', g). Pygidium conical with a pair of eye-
sjiots.
Type locai.iiy: Dredged off Lamberts Ha)-, South Africa.
Records: C^ape (32/17,^, 34/17/d, 36/21/d).
DisTRiBuiiox : Endemic.
SABELLIDAE 785
ORIOPSIS CauUcry and Mcsnil, 1896
Small Sabcllids with eight thoracic and four or more abdominal segments.
Branchial lobes without branchial hearts but each has three to five flanged radioles
with long pinnules but without eyes or external stylodes. Usually one to two free
ventral filaments simulating palps. Lips membraneous. Collar divided dorsally
and united ventrally. A pair of eyes and sometimes a pair of otocysts in the collar
segment. Thoracic notosetae are winged capillaries and the neurosetae are long-
shafted hooks. Abdominal uncini serpuliform without shafts. Pygidium often with
eye-spots.
Type species : Fahricia armandii Claparede, 1864.
Key to Species
1 Five to seven abdominal setigers . ......... 2
- Eiglit or more abdominal setigers .......... 3
2 Collar absent (fig. 37. 9. j). Abdominal imcini 15-20 per row ... O. neglecta
— Collar low laterally and with a tongue-shaped projection ventrally. Up to 10 abdominal
uncini per row ............ O. parvula
- Collar with a smooth, straight margin (fig. 37.9.0). .'Abdominal unrini 12 per row O. ehlersi
3 Collar with a scalloped margin (fig. 37.10.3). -Abdominal uncini with an enlarged basal
tooth ............ .0. eimeri
— Collar with a smooth, straight margin. .\bdominaI uncini without an enlarged basal
tooth ............. O. bansei
Oriopsis neglecta Banse, 1957
(fig- 37-9-h-n)
Oriopsis neglecta Banse, 1957: 85, fig. 5 d-e ; Day, 1 96 1 : 546, fig. 1 5 g-m.
Body (fig. 37. 9. n) fairly stout and 3-4 mm. long. Branchial lobes (fig. 37.9.J)
without branchial hearts but each has three to four free radioles with broad
flanges and long pinnules. Two triangular lips medially and a pair of slender "palps"
ventrally each half as long as the radioles. Collar represented by a thickened ridge
around the base of the branchial lobes with a triangular ventral expansion. No
peristomial eyes. Body of eight thoracic and five to six crowded abdominal setigers.
Pygidium without eye-spots. Thoracic neurosetae are long-shafted hooks (fig.
37. 9. k, 1) with one large tooth and an arc of several denticles above the main fang.
Abdomen with two slender capillaries per neuropodium and 15-20 very small,
roughly square uncini per notopodium. Each uncinus (fig. 37. 9. h, i) with about six
rows of six very long teeth above the large basal tooth and recurved prow.
Type locality : Luderitzbucht, South West Africa.
Records: South ^Vest Africa (22/14/i and 26/15/i, s) ; Cape (34/i8;'i, s).
Distribution : Endemic.
786
P(M,Yt:ii.\i: r.\ of southern atrica
l''Ki. 37.9. Faliricia Jilaiiicnlriui. (a) Entire worm (30 times life size). (11) Dorsal view of collar.
(f:^ Thorarir notoscta. (i), F.) Thoracic hook, (f, g) Abdominal iincinus. Orinpsis iieglirla.
(II, ij Abdominal uncinus. (j) Ventral view of collar region, (k, l) 'Ihoracic hook.
(m) Tliorat ic capillary, (n) Entire worm (30 times life size). Oiiojnis ehlrrsi. (o) Lateral
view of collar region, (p, Q) Thoracic hook, (r, s) Abdominal uncinus. (x, u) Longer and
shorter forms of thoracic capillary.
SABELLIDAE 787
Oriopsis parvula (Ehlers, 1913)*
Oria panuta Ehlers, 1 9 1 3 : 580.
Oriopsis parvula : Day, 1 96 1 : 545.
Body I "4 mm. long with eight thoracic and six abdominal setigers. Each branchial
lobe with three flanged radicles bearing long pinnules and a shorter, stouter filament
projecting between the dorsal pair. Collar widely gaping dorsally, low laterally and
forming a triangular projection ventrally. Eyes present. Thoracic notosetac of
setigers 2-8 with seven similar capillaries without wings (?). Neurosetae as four to
five long-shafted hooks. Abdominal uncini up to 10 per row, each with several rows
of several teeth above the recurved prow.
Type locality : Simonstown, South Africa.
Records : Cape (34/18/s) - known only from the original record.
Oriopsis ehlersi Day, 1961
(fig. 37.9.0-u)
Oriopsis ehlersi Day, 1 96 1 : 546, fig. 16 a-g.
Body 3 mm. long with eight thoracic and five to seven abdominal setigers. Each
branchial lobe (fig. 37.9.0) with three flanged radioles having long naked tips and
six to eight pairs of pinnules which aU reach the same level. Two long filaments
(? palps) between the ventral pair of radioles. Collar fairly well developed and
equal laterally and ventrally. The edge is smooth apart from a deep notch in the
mid-ventral Une. Eyes present. Thoracic notosetae of setigers 2-8 are longer and
shorter winged capillaries (fig. 37.9.1, u), the shorter forms having very narrow
blades. Neurosetae are seven to eight long-shafted hooks (fig. 37. 9. p, q) with two
arcs of smaU denticles above the main fang. Abdominal uncini about 12 per
notopodium, each uncinus (fig. 37. 9. r, s) having eight to eleven rows of teeth with
five teeth per row. Lowest tooth larger than the others.
Type locality : False Bay, South Africa.
Records: Cape (34/18/s).
Distribution : A single record.
Oriopsis eimeri (Langerhans, 1880)
(fig. 37.10.a-g)
Oria eimeri Langerhans, 1880 : 1 17, pi. 5 fig. 31 a-c.
Oriopsis eimeri : Day, 1961 : 547, fig. 16 h-o.
Body 1-2 mm. long with eight thoracic and eight to ten abdominal setigers.
Branchial lobes (fig. 37. 10. a) vsithout branchial hearts but each with three flanged
radioles with eight to ten pairs of long pinnules and long naked tips. A pair of
membraneous lips. CoUar widely divided dorsally, well developed laterally and
*This species was not figured by Ehlers and the exact characters are doubtful.
788 I'OIACllIAETA OF SOUIIll^RX AI-RK;.\
unilctl \('iUr<illy. The free edges are crenulatc with about lo scallops on each side.
K\(s and otoeysts present inlernaUy. Thoracie notosetae (fig. 37.io.b, c) are longer
anil shorter winged capillaries. The long-shafted neuropodial hooks (fig. 37.io.d, e)
ha\e the main fang siuiiKiunted by a pair of large teeth and then two arcs of smaller
denticles. Abdominal capillaries arc very fine with the blade only visible at the base.
.Abdominal uncini nine to eleven per notopodiiim, each roughly square with a rather
large basal tooth and a broad crest of about six rows each with six to eight teeth
(fig. SJ.io.f, g).
Type locality : Madeira Island.
Records: Cape (34/18 i).
Distribution : Madeira.
Oriopsis bansei Day, 1961
(fig. 37.io.h)
Oriopsis bansii Day, 1961 : ^,46.
Oriopsis paniila : (non Ehlcrs) Banse 1957: Bo, fig. 5 a-c.
Body about 2 mm. long with eight thoracic and nine aisdominal segments. Each
branchial lobe with three to four Hanged radioles and at least one long ventral
filament (? palp). C'oUar equally developed laterally and ventrally with a smooth
edge. Thoracic notosetae of setigcrs 2-8 are six long and three to four short
capillaries, the long ones having characteristically broad blades. Neurosetae have
several closely packed denticles above the main fang. Abdominal uncini 10-12 per
notopodium, each with about 10 rows of teeth and six teeth per row; basal tooth
not enlarged (fig. 37.10.11).
Type Loc:ALrrY : Luderitz, South West Africa.
Records: .South ^Vcst Africa (26/15,1).
DiSTRinuTicjN : A single record.
DESDEMONA Banse, 1957
Small Sabellids closely related to Oriopsis with eight thoracic and four to twelve
abdominal segments. Branchial lobes without branchial hearts but each with three
free radioles with poorly de\eloped lateral flanges or none at all. A pair of mem-
Ijraneous lips on either side of the mouth and a p.iir of filam<-ntous ]ialps \cnlrally.
Collar divided dorsally, but united ventrally to form a median \entral lobe. A pair
of eyes within the collar segment. Thoracic notosetae are winged capillaries and
the neurosetae arc long-shafted hooks. Abdominal notosetae are square serpuliform
uncini and the neurosetae are slender capillaries. Pygidium with or without eyes.
Type species: Diukiiwna oinata Banse, 1957.
SABELLIDAE
789
Fig. 37.10. Oriopsis eimeri. (.k) Dorsal view of anterior end. (b, c) Longer and shorter
types of thoracic capillary, (d, e) Thoracic hook, (f, g) Abdominal uncinus. Oriopsis
bansci (from Augener, 191 8). (h) Edge-on view of thoracic uncinus. Desdemona omata.
(i) Entire worm (30 times life size), (j, k) Thoracic hook, (l, m) Abdominal uncinus.
(n, o) Longer and shorter forms of thoracic capillary.
790 Pdl.VClIAETA OF SOUTHERN AFRICA
Desdemona ornata Bansc, 1957
(fig. 37.10.i-0)
Dt'sdttnona ornata Banse, 1957 : 90, fig. 7 a-c.
Body (fig. 37.io.i) about 2-4 mm. long with ciglit thoracic and seven to nine
abdominal segments. Branchial lobes without branchial hearts but each with three
free radiolcs with reduced lateral flanges and with four to five pairs of long stout pin-
nules. Each radiolc has two streaks of dark pigment at the base and there is a pair
of dark membraneous lips between the branchial lobes. The ventral lips are not
pigmented. Between the ventral pair of radiolcs there is a pair of long filaments
which simulate palps but have the same structure as adjacent pinnules. Collar
indistinct dorsally, notched laterally and forms a stout contractile lobe ventrally.
A pair of lateral eyes within the collar segment. Thoracic notosetae (fig. 37.10.11, o)
arc slcndcr-wingcd capillaries of two lengths, the shorter ones having very fine tips.
Thoracic neurosetae are two to three long-shafted hooks (fig. 37.10.J, k) with three
to four denticles above the main fang as seen in lateral view and a close-set cap of
about nine denticles in fice view. Abdominal uncini (fig. 37.10.I, m) are C[uad-
rangular plates with four to six rows each with about five teeth. Pygidium without
eye-spots.
Type locality : Hermanus Estuary, South Africa.
Records: Cape (from 33/18,'e to 33/27/c) ; Natal (30/30/e and 29/31/1).
Distribution : Endemic.
SERPULIDAE 791
Family SERPULIDAE Savigny, 1818
Tubicolous worms encased in calcareous tubes usually attached to hard objects.
Body divided into three regions namely a head region provided with a branchial
crown for respiration and filter-feeding, a thorax of three to eight segments and an
abdomen of numerous segments. Prostomium indistinct and fused to the buccal
segment which bears the branchial crown formed of two fans of bipinnate radioles.
One radiole usually modified and enlarged to form a stalked operculum. Palps
seldom present. A pair of laige nephridia open by a single dorsal pore at the base
of the branchial crown. Second segment (first setiger) expanded to form a mem-
braneous collar, the two halves of which extend back as thoracic membranes above
the notosetae. A tube-building glandular fillet associated with the collar. Para-
podia biramous but poorly developed. Thoracic notosetae are bundles of limbate
capillaries and the notosetae are rows of toothed plates (uncini). Abdominal setae
similar but inverted so that the neurosetae are capillaries and the notosetae are
uncini. A ventral groove or copragogue runs forward from the anus and bisects
the glandular ventral pads of the abdomen but curves round to the dorsal surface
at the junction of thorax and abdomen.
BIOLOGICAL NOTES
The serpulids are closely related to the sabellids and like them they are suspension
feeders. But they are more specialised than the sabelUds for they have lost their
palps, the tube is always calcareous and one of the radioles of the branchial crown
has been modified to form a stalked operculum which plugs the entrance of the tube.
When alarmed the worm retracts extremely rapidly for it has well developed giant
nerve fibres. The operculum serves not only for protection but also to reduce water
loss and it is noteworthy that while non-operculate forms such as Protula are infra-
tidal, forms with calcareous opercula such as Pomatoleios kraussii live above mid tide.
Large coralliform masses of Pomatoleios dominate many Cape and Natal shores but
the distribution stops sharply at Cape Point presumably because tlie water of the
Bcnguela current is too cold.
The tubes of most serpuHds are attached to rocks or other hard objects. In the
tropics, of course, many of them grow on coral. Hydroides norvegica seems to prefer
floating objects such as buoys and the hulls of ships, particularly the shaded area
under the stern. It is not surprising that H. norvegica has a cosmopolitan distribution.
Mercierella enigmatica is another traveller. About 1920 it suddenly appeared in the
estuaiy of the river Seine in France, possibly carried by ship from some estuary in
India. Since then it has been reported from warm estuaries all over the world and
is even found in the warm conduits from power stadons in England. It can tolerate
salinides both above and below that of sea water but it is never found in the sea,
always in estuaries.
The subfamily Spirorbinae includes some of the most specialised of the serpulids.
They are all small and asymmetrical and, as described later, the larvae settle down
wdth the dorsal surface against the substrate. Some species will settle on a variety
of substrates but others, as Professor Knight-Jones and his colleagues have shown.
792 i'OIA CHAK lA Ol' SOLlllKKX AIKIC.A
arc NLTV scli-iti\i'. In Suulli Alikan seas tlicrc is (ini- spi-cics tli.il li\cs mi call .ircinis
bryozoa, anotln-r ihat lives on siu-lls inhabilrd by ii<Tniit crabs ami aridthcr (in tia-
fronds of algae.
One of the \ery few serpulids that does not have an atlaehed tnbe is Di/rii/iii, a
deep water genus whicli lives on muddy bottoms in a tusk-shaped tube rather like
that of the mollusc Denlalium. The method whereby it prevents itself from being
bin'ied in the sih is imknuwn.
DI\ISI<)X IXIO SUB-FAMIMES
Useful re\icws of the family Serpulidae will be finuid in Morch (1863), Pixel]
(1912 and 1913a) and Fauvel (1927). The three subfamilies are distinguished by
the nature of the opcrciduiu and number iif thoracic segments.
Key to Scbf.xmilies
1 Thorax symmetrical with lu-e to twelve tlioracic setigers ...... 2
- Thorax asymmetrical witli three to four thoracic setigers (hg. 38.i.d). (Shell small and
spirally coiled) .Spirorbinae( p. 792)
2 Operculum always present and never has pinnules on its stalk. Six to seven thoracic
setigers Serpclin.xe (p. 798)
- Operculum either absent or poorly developed and retains pinnules on its stalk. Five to twelve
thoracic setigers Filogr.aninae (p. 816)
Subfamily SPIRORBINAE C:hamberlin, 1919
Small Serpulids with spirally coiled shells. Thorax of three to four spirally
twisted and asymmetrical segments, the last of whii li may lack setae on th<' convex
side. Operculum always present and lacks pinnules on its stalk. Developing eggs
retained either in the hollow operculum or inside the shell in a membraneous sac.
Records from sotithern Africa
Spirorhis (Dtwiospiia) foraminosiis Moore . . . 45?!, 51CS
Spirorhis {Laeospira) laevis Quatrci'a.gcs . . . aiGi
Spiiorbii (Paralaeoipira) adeonella Day . . . .'iCC^s
■AS Spirorhis (Laeospira) sp. ..... .'j'C'.s
Spirorbis (Paralaeospira) capouis Day . . . 5 i(;;sd
Spirorlns {Paralaeospira) patagonkus CLtullery & Mesnil S'^^'s
as Spirorbis borealis Day (non Daudin) . . . 44tli
THE MAIX DI.AGXO.STIC CHARACTERS
The main works on the Spiiorbinac are those of ClauUery and .Mesnil (1897) and
Pixell (1912).
Thf lube. The larva settles with its dorsal surface against the substratum and the
tube becomes spirally twisted. If it coils in a clockwise direction when \iewed from
above (i.e. frr)m the ini>rjihologically ventral surface) it is said to be sini\lral and if it
SERPULIDAE 793
coils anticlockwise it is said to be dextral. The surface of the shell may bear longi-
tudinal ridges or cross bars but only major differences are important.
The operculum. The stalk is expanded distally and the operculum is a calcareous
plate, cone or hollow cylinder embedded in the stalk by a basal talon. Both the shape
of the talon and the operculum are important, but it should be noted that the
operculum thickens with age.
Reproduction. The fertilised eggs are retained either in the hollow and perforated
operculum or in a membraneous sac (possibly corresponding to a false operculum)
which lies inside the tube next to the abdomen.
The number of thoracic segments. There are three to four thoracic segments of which
the first bears notopodial collar setae only and the rest both notopodial capillaries
and neuropodial uncini. Occasionally the capillaries are missing from the convex
side of the last thoracic segment.
Setae. Thoracic notosetae are Umbate capillaries. The collar setae are often
speciahsed with a separate toothed lobe or fin at the base of the blade which itself
may be either smooth or saw-edged. Subsequent notosetae may include "setae of
Apomatus" with minute comb-teeth on the distal part of the blade. Thoracic uncini
have two to five rows of numerous teeth and the basal gouge may be truncate,
emarginate or even be divided into five divergent teeth.
SPIRORBIS Daudin, 1800
Body minute and asymmetrical, encased in a spirally coiled tube. Operculum
calcareous and sometimes contains the eggs ; it may have a basal projection or
talon in the stalk which lacks wings. Thorax of three to four segments. First few-
abdominal segments greatly elongated. Collar setae often with a separate toothed
lobe at the base of the blade. Setiger 2 with smooth-bladed capillaries. Setiger 3
with smooth-bladed capillaries and often setae of^ Apomatus with the distal end of the
blade denticulate. Thoracic uncini are rectangular plates with two or more rows
of teeth and a basal gouge. Abdominal capillaries have toothed blades set at an
angle to the shaft.
Type SPECIES : .Spirorbis borealis Da.ud\n, 1800.
Key to Species
Three rows of uncini on the inner, concave side of the thorax. Tube coils clockwise when
seen from above (Paralaeospira) (fig. 38.1.3, d) ...... . 2
Two rows of uncini on the inner, concave side of the thorax. Tube coils clockwise or anti-
clockwise when seen from above .......... 4
Tube smooth or longitudinally ridged. Setae of /!/)omato present (fig. 38. i.h) . . 3
Tube transversely ridged (fig. 38.1. k). No setae of .^/lomadts . . S. [P.' adeonella
Talon smoothly conical (fig. 38.1. b, c) ...... S. (P.) patagonicus
Talon warty (fig. 38.1.5) . . . . . . . . . S. (P.) capensis
794 I'OL^CIHAKTA OF SOLrUl^R.N Ai RU:.\
4 'I'ubr coils clockwise wlien seen from above. Goui;e of uncinus wiih a smooth edge
S, (Z,.) laevis
— Tube coils anti-clockwise when seen from above. Gouge of unc iiiiis with three to h\e
prongs (fig. 38.:2.c. f) ........ S. {D.) foratninosus
Spirorbis [Paralaeospira] adeonella Day, 19G3
(fig. 38.1. k-p)
spirorbis {Paralaeospira) adeonella Day, 1963a : 44'2. fit2;. rj h-]3.
Spirorbis (Laeospira) sp. Day, 1961 : 557.
Length ;5 mm. Tube ((fig. 38.i.k) vitreous and coiled clockwise when viewed from
above (sinistral) with a well marked series of annular lidges. Operculum (fig. 38.1.
I, m). o\al with a concave surface and a knob-shaped talon. Incubation in the tube.
Seven branchial radicles. Collar separate dorsally. Four thoracic setigers, the first
with collar setae only, the second and third with both notosetae and uncini and the
foiuth represented by a short row of uncini on the concave side of the body. No
notosetae on setiger 4. Abdomen with about 13 setigers. Collar setae include a few
fine capillaries and several setae with a separate dentate lube .it tlie base of the blade
(fig. 38.1.11). Notosetae of setigers 2 and 3 are winged capillaries and a few finer
capillaries. No setae o^ Apomaliis. Thoracic uncini (fig. 38.1.0, p) with three to
four rows of teeth and 18 teeth per row. Gouge expanded .ind trunc.ite. Tvibe
growing on the calcareous polyzoan Adeonella sp.
Type locality ; Mossel Bay, South Afiica.
Records: Cape (34/22/s, 34/25/s).
Distribution : Endemic.
Spirorbis [Paralaeospira) patagonicus C'aullerv & Mesnil, 1897*
(fig. 38.1. a-j)
Spirorbis (Paralaeoipira) patagonicus Claullery & Mcsnil, 1897 : 20-,, pi. 8 fig. la ; Day, 1961 : 554.
Length 3-4 mm. Tube (fig. 38.1. a) coiled clockwise when seen from above
(sinistral), dense, chalky white (about 2 mm. in diameter), occasionally erect. Four
asymmetrical thoracic setigers (fig. 38.i.d) the first with collar setae only, the second
and third with both notopodial capillaries and neuropodial uncini, and the fourth
with uncini only on the concave side. Twenty to thirty abdominal segments.
Operculum (fig. 38.i.b, c) subcircular with a simple conical talon. Incubation in
the tube. Adults with eight to nine radioles. Thoracic membrane free to the end
of the thorax on the outer side. CVillar setae (fig. 38.i.e, f) include fine capillaries
and stouter firms with a separate toothed lobe at the base of the minutely serrated
or pilose lil.ide. Setiger 2 with limbate capillaries only (fig. 38.i.g). Setiger 3 with
both capillaries and setae of Apomalui with three-quarters of the blade dentate
*Vt.Ty close ttj .V. 7ii(iUitdi f laiillrry anti Mtsnil.
SERPULIDAE
795
Fig. 38.1. S/iirorbis fiatagonicus. (a) Tube, (b, c) Two forms of opercula. (d) Entire worm
(25 times life size), (e, f) Collar setae, (g) Normal capillary from setiger 2. (h) Seta of
Apomatus. (i, j) Thoracic uncinus. Spirorbis adeonella. (k) Tube, (l, m) Operculum.
(n) Collar seta, (o, p) Thoracic uncinus. Spirorbis capensis. (q) Collar seta, (r) Seta of
Apomalus. (s) Operculum, (t, u) Thoracic uncinus. (\') Tube.
796 POLVCHAETA OK SOUTHERX Al-RKIA
(fig. 38.1.11). Uncini (fig. 38.1.1, j) with a large cmarginale gouge and three to
four rows of teeth with 14 teeth per row.
Type locality : Orange Bay, Patagonia.
Records: South West Africa (22/14/1 and 26/15/i) ; Clapc (from 29/16/1 to
34/18 1, s and 32/28/i).
DiSTRiBLTiON : Subantarctlc (Kerguelcn (1, s), Patagonia, Falkland Is., Tristan
da Cunha (i), Marion Is. (i) ).
Spirorbis [Paralaeospira) capensis Day, 1961
(fig. 38.i.q-r)
Sfiirorbis (Pa^alaeopira) capensis Day, 1961 : 5=14, fig. 18 a-h.
Length 3 mm. Tube (fig. 38.1.V) coiled clockwise when seen from above (sinis-
tral). Operculum subcircular with a conical talon having tooth-like outgrowths
(fig. 38. 1. s). Incubation in the tube. Four thoracic setigcrs, the first having collar
setae only, the second and third with both notopodial capillaries and neuropodial
uncini and the fourth with uncini on the concave side only. Collar setae of two types :
(a) stout setae with coarsely serrate blades and a separate toothed boss at the base
(fig. 38. 1. q) and (b) slender setae with smooth blades. Setiger 3 with both smooth-
bladed capillaries and setae oi Apomatus with the distal third of the blade expanded
and finely toothed (fig. 38.i.r). Thoracic uncini (fig. 38.i.t, u) with two to four
rows of teeth and 20 teeth per row ; basal gouge expanded and truncate.
Type locality : False Bay, South Africa.
Records: Cape (34/18/s and 36/21/d).
Distribution : Only two records.
Spirorbis (Laeospira) laevis Qiiatrcfages, 186-,*
(fig. 38.2.a-b)
S/>irorbii Intvii Quatrefagcs, 1865 : 490.
Sptrorbis {Laeospira) laevis: Fauvel, 19J7 : 397, fig. 134 1-p.
Tube coiled clockwise when seen from above (sinistral). Body (fig. 38. 2. a) witii
three thoracic and nine abdominal setigers. Only two branchial radicles on eacii
side. Operculum (fig. 38. 2. b) vase-shaped, covered with a calcareous lid and
contains the developing eggs ; no talon. Collar setae are plain, winged capillaries.
Setae ni Ajwmatiis with minutely toothed blades are present in the third setiger.
Type locality : Gulf of Gascony, France.
Records: Cape (34/18/i).
Distriiu:tion' : Bay of Biscay ; Mediteirancan ; Senegal (d).
*.\ccorcling to Caullcry and Mcsnil (l8g7) the original description is vague and incomplete and pmbaljly
refers to a young form of a well-known European species. The South .\frican record is doubtful.
SERPULIDAE
797
Fig. 38.2. Spirorbis laevis. (a) Entire worm (60 times life size after Claparide). (b) Oper-
culum (after Quatrefages). Spirorbis foraminosus. (c) Tube, (d) Collar seta, (e) Oper-
culum, (f, g) Thoracic uncinus.
Spirorbis [Dexiospira) foraminosus Busch, 1904
(fig. 38.2.c-g)
Spirorbis (Dexiospira) formninosus Busch, 1904: 176, text-fig e; Day, 1961 : 556, fig. i8j-k.
Length 3-4 mm. Tube (fig. 38. 2. c) coiled anticlockwise when viewed from above
(dextral) with five longitudinal ridges and transverse slits in the grooves between
them. Body with three thoracic setigers or two rows of uncini on each side. Incuba-
tion in the operculum which is cylindrical with perforated calcareous walls, a lid
with a marked rim and a subcircular basal plate (fig. 38. 2. e). Collar setae (fig.
38. 2. d) with pilose blades and without basal lobes. No setae of Apomatus in setiger
3. Uncini (fig. 38. 2. f, g) with four to five rows of teeth and 20-25 teeth per row;
basal gouge with three to five divergent prongs. Tubes found on fronds of algae.
Type locality : Japan.
Records: Cape (34/1 8/s) ; Mocambique (26/32/i).
Distribution : West Indies ; Indian Ocean (i, s) ; Japan (s).
798
l'OI,> t:HAKTA OF SOUlllKRX AFRUIA
Subfaniilv SERPULINAE M.uLeay, iS-n
Body symnii'lrical. Pruslumium indislinguisluiblc from the buccal segment. A
well developed operculum without pinnules on the stalk. Thorax of seven segments
but the fust or collar segment may lack setae. Thoracic uncini with a single row of
teeth.
Records from southern Africa
Ficopomatus captrtsis Day
Hydroides liifiircalus (Pixell)
Hydroides dipoma (.Schmarda) .
as F.tipolamus dipoma Schmarda .
as Hydroides uncinalits \'ar. tnaironvx Ehlers
Hydroides heteroceros (Grube) .
Hydroides lumdifera (Claparede)
Hydroides monoceros (Gravier) .
Hydroides norvegica Gunncrus .
as Hydroides uneinatn \ar. niultispinosa McI utosh
Hydroides ralumianus Angener .
Hydroides uncinata (Philippi) .
Mercierella enigmatica Fauvcl .
Neovermilia eapcnsis Day
Pomatoleio\ kraussii (Baird)
as Placostegiis cariniferus var. krassii Baird
as Placostegus caendeus Schmarda (pp) .
as Pomatoleios crosslandi Pixell
as Pomatoceros caeruleus Fau\el (non .Schmarda
Scrpula verrriieiilaris Linnaeus
Serpiili! lertiiiciilaris var. echinata Linnaeus .
Spirohranchus giganteus (Pallas)
Spirobranchus tetraccros (Schmarda) .
Vernnliopsis babylonia Day
as Vermiliopsis pygidialis (non \V'illcy) .
Vermiliopsis glandigerus (Gravier)
5 ' C's
4oNi
26Ai, 44Gi, -, iCs
4Gi
2iCi
56MS
45Ni
45Pi
33CS, 4f,PsNs, 5iGsd
32C:sNd
45Pi, — Ps
45P1
4oNe Cc
48C:s
44Gi, 48C;i
5c;i
4C;i
35Ci, 36XiCi, 40X1
27Mi
2oC;i, 2iCi, 35Ci,
44Ci, 45NiPi, 5iC:sd
51GS
4oNi, 44Ci
45Pi, -Ms
Vema Sea Mount
51GS, 44Ci
2 7Mi, 36Gi,
44Ci, 5iCsd
IHE MAIN DI.-^GNOSTIC CHARACTERS
The operculum is cither well chitinised or bears a calcareous plate. Calcareous
opercula become thicker with age changing from flat to dume-shaped. The most
complex opercula are found in the genus Hydroides where a calyx-like /w««f'/ composed
of numerous radii supports a croivn of horny spines. The opercular stalk may be
cylindrical or flattened with a pair of wings at the distal end belo\v' the operculum.
Opercula are easily lost and a small reserve operculum often develops from the
other branchial lobe. It has been termed njiilse operculum.
SERPULIDAE 799
Thoracic membranes. The backward prolongations of the right and left sides of the
collar may continue as free membranes to the posterior end of the thorax and form
a ventral frilly membrane at the junction of the thorax and abdomen. In some
species however the thoracic membranes are much shorter and end behind the
second or fifth thoracic setiger. In rare cases the two sides of the collar fuse across
the dorsal surface (e.g. Meopomatus). In most species the collar is divided into three
main lobes - a ventral lobe and two lateral ones. On each side at the junction
of the ventral and paired lateral lobes there is a small glandular fillet responsible
for tube formation.
Thoracic notosetae. Although the second segment or collar lacks uncini it possesses
notosetae in all genera except Pomatoleios and Placostegus. These collar setae are often
different from those of later thoracic segments. There are two forms ; one is hair-
like with hardly a trace of a blade and the other form is usually much stouter with
a coarsely toothed blade and in some genera they form bayonet setae with stout bosses
at the base of a sword-like blade. Subsequent notosetae are mainly limbate capill-
aries but some genera develop a few setae of Apomatus in which the distal part of the
blade has minute comb-like teeth.
Thoracic uncini are quadrangular plates with curved teeth along one side usually
arranged in a single row. In some geneia the first and largest tooth is expanded to
form a gouge. Care is needed when examining the gouge for if the uncinus is twisted
the gouge may appear forked.
Abdominal setae. While the neuropodial uncini are essentially similar to those of
the thorax the capillaries are more variable. The posterior ones tend to be more
elongated than the anterior ones which have toothed blades. Geniculate setae have
gradually tapering blades set at a slight angle to the shaft. In others the base of the
blade is expanded and the tip shortened to form a trumpet-shaped structure or in
the extreme case of Serpula the toothed blade is expanded at right angles to the shaft
forming a T-shaped structure. The elongate capillaries at the end of the abdomen
may be quite smooth.
Key to Genera
1 Collar setae absent ............ 2
- Collar setae present ............ 4
2 Opercular stalk winged. (Operculum a flattened calcareous plate) POMATOLEIOS (p. Soo)
- Opercular stalk smooth ........... 3
3 Tube free, tusk shaped. No row of eyes on collar ..... DITRUPA*
- Tube attached. -^ row of eyes on the collar PLACOSTEGUS*
4 Opercular stalk winged (fig. 38.3. f) ......... 5
- Opercular stalk smooth ........... 8
5 Collar setae few, fine and capillary POMATOCEROS (p. 801)
- Collar setae numerous with a spinulose lobe at the base of the blade or bayonet-shaped
with stout bosses at the base of the blade ........ 6
6 Thoracic membranes reduced or absent. Collar with bayonet-setae OMPHALOPOMOPSIS*
- Thoracic membranes well developed. Collar setae with a spinulose lobe at the base of
the blade ............. 7
8oo POI.VCHAETA OK SOUTHERN Al-RICA
7 Operculum a calcareous plate with branching processes . . SPIROBRANCHUS (p. Roi)
- Operculunt as one or a vertical series of calcareous plates without branchiiii,' prmesscs
POM A TOSTEGUS*
8 Collar setae bayonet-shaped with a separate toothed boss at the base of the blade (hi;.
38.4.dl 9
- Collar setae without a separate boss at the base of the blade . . . . . 12
9 Operculum with a central crown of chitinous spines (fig. 38.4.e) . HYDROIDES (p. 804)
- Operculum without a crown of chitinous spines ....... 10
10 Operculum funnel-shaped with marginal radii. ■Xbdominal capillaries T-shaped
SERPULA (p. 809)
— Operculum pear-shaped or conical without spines or marginal radii, .\bdominal
capillaries geniculate ...........11
11 Thoracic uncini with a single row of large teeth . . . FICOPOMATUS iyi. P:\o)
— Thoracic uncini with three or four rows of small teeth HYALOPOMATUS*
12 Collar setae with serrated blades. Operculum with numerous short spines . . 13
— Collar setae with smooth blades. Operculum without spines ..... 14
13 Thoracic membranes fused across the dorsal surface . . NEOPOMATUS*
— Thoracic membranes not fused ...... MERCIERELLA fp. 812)
14 Setae oi Apomalus (fig. 38. 6. e) among the notosetae. Operculum conical and hornv with
internal septa VERMILIOPSIS Ip. Ki 2)
— Xo setae o( Afiomnliis. Operculum spherical or concave, not horny; no internal septa
NEOVERMILIA (p. 814)
POMATOLEIOS Viy^vW, 1913
Body SNiiinifliic.il witli six thoracic sctigcrs and numerous abdfimiii.il ones.
Operculum flat and calcareous. Opercular stalk winged. Clollar setae absent.
Thoracic notosetae are simple limbatc capillaries. Uncini with a gouge and numer-
ous teeth. Abdominal capillaries obliquely truncate with one side produced as a
long spine. Tube with a projection over the mouth ; worms often gregarious, forming
coralliform masses.
Type species: Placostegus caiiiiifiiiis \ar. haiissii Baird, 1865.
Pomatoleios kraussii (Baird, 1865)
(fig. ;58.3.a-r)
Placostegus cariniferus var. kraussii Baird, 1865 : 14.
Pomatoleios crosslandi Pixell, 1913 : 85, pi. g fig. 10.
Pomatoleios kraussii: Day, 1955 : 449.
Length up to 25 mm. Tubes gregarious forming blue coralliform masses (fig.
38.3.3). Individual tubes (fig. 38. 3. b) with a flattened dorsal ridge projecting
forward over the entrance. Operculum (fig. 38. 3. f) a flattened or slightly concave
calcareous plate. Opercular stalk broad and flattened with smooth wings. Branchial
lobes each with 13-16 radiolcs which are banded blue and white and united by a
web for half their length and end in bare tips, dollar notched deeply laterally and
very broad ventrally. A separate dorso-lateral tube-building fillet inside the collar.
Thoracic membranes extend to the end of the diorax and arc united and free from
the body Neutrally. C!(illar setae absent. No eye-spots. Thoracic nf)tosetae have
simple striated blades. Thoracic tiiicini (fig. 38. 3. c, d) witli a gouge and eight to ten
SERPULIDAE 801
teeth in a single row. Abdominal neurosctae (fig. 38. 3. e) obliquely truncate with
fine teedi and one side produced into a long spine.
Type locality : Cape of Good Hope.
Records: Cape (from 34/18/1 to 32/28/i) ; Natal (from 31/29/i to 27/32/i) ;
Mocambique (26/32/i).
Distribution : Tropical eastern Africa (i) ; Madagascar (i) ; Madras (i) ;
Red Sea (i) ; Japan.
POMATOCEROS Phihppi, 1844
Operculum flattened, sometimes with a few low spines. Opercular stalk winged.
Collar setae small and capillary. Body symmetrical with seven thoracic segments
and numerous abdominal ones. Thoracic notosetae are capillaries. Uncini with
numerous teeth, the first one being enlarged and gouge-hke. Abdominal capillaries
trumpet-shaped with one side produced into a long spine. Tube triangular usually
adnate with the upper side keeled and ending in a projection over the mouth.
Type species : Serpula triquetra Linnaeus, 1 767.
Potnatoceros caeruleus (Schmarda, 1861)*
(fig. 38.3.g)
Placoslegtis cafTuletis : Schmarda, 1861 : 29, pi. 21 fig. 178.
Pomatoceros slrigiceps Morch, iSB-j : 412; Mcintosh, 1885 : 520, pi. 55 figs. 3-4 ; pi. 31 A figs. 26-28 ;
Ehlers, 1905 : 67, pi. 9 figs. 1 1-19.
Tube blueish, ridged dorsally, often with a projection over the entrance. Gills
and body blue. Often gregarious. Operculum (fig. 38. 3. g) flattened, sometimes with
two tiers of plates. Opercular stalk with small smooth wings. Collar setae are a
few fine, limbate capillaries. Subsequent notosetae are normal limbatc capillaries.
Thoracic uncini widi a single row of 10 teeth and a basal gouge ending in a pair of
divergent prongs. Abdominal capillaries expanded at the end and one side produced
as a long spine. Thoracic membranes from collar to end of thorax.
Type locality : New Zealand.
Records: Schmarda's original record from "Cape of Good Hope" is very
doubtful.
Distribution: New Zealand (i).
SPIROBRANCHUS BlviinvMe, 1818
Body symmetrical with seven thoracic setigers and numerous abdominal ones.
Operculum as a calcareous plate bearing a group of branched spines. Opercular
stalk winged. Collar setae often hirsute. Subsequent thoracic notosetae are hmbate
•According to Baird (1865), P. caeruleus Schmarda is a sy-nonym off*, cariniftrus Gray, 1843.
8o2
l'UL\ t:llAE lA OF SOUrill-.R.\ AI'RIOA
Flo. ;i8.3. Pomnloltios kraiinii. (a) Part of colony, (n) Tube of solitary individual, (c, n)
Thoracic uncinus. (r.) .Xbdominal capillary, (f) Anterior end with operculum. Pomaloceros
(oeruleui. {<;) Operculum (modified from Mcintosh, liiS', as P. ilrigiceps). Spirobrtituhin
gignnlrii^. In) Anterior end. (i) Collar seta, (j, k) Thoracic uncinus. Spirobra'ichin Irlra-
ceros. (1., M; Thoracic uncinus. (n) Operculum.
SERPULIDAE 803
capillaries. Uncini with numerous teeth, the first one enlarged and gouge-like.
Abdominal neurosetae are trumpet-shaped and denticulate with one side produced
as a long spine.
Type species : Serpula giganlea Pallas, 1 766.
Key to Species
1 Branchial lobes spiral with four to six whorls of radioles (fig. 38. 3. h) .... 2
- Branchial lobes flat with one circle of radioles. (Uncini with 13-16 teeth) ... 3
2 Thoracic uncini with 15-18 teeth. Operculum with two to four branching processes
5. giganteus
- Thoracic uncini with 26 teeth. Operculum with one branching process . S. gardineri*
3 Operculum without branching processes . . . . . . . S. maldivensis*
- Operculum with four to six branching processes (fig. 38. 3. n) . . . S. tetraceros
Spirobranchus giganteus (Pallas, 1 766)
(fig. 38.3-h-k)
Serpula gigantea Pallas, i 766 : 402.
Spirobranchus giganteus : Pixell, 1913 : 80, pi. 8 fig. 6.
A large species reaching 120 mm. with a stout, ridged, cylindrical tube and four
to six whorls of branchial radioles usually coloured red. Opercular stalk (fig. 38. 3. h)
with small, blunt wings. Operculum oval with two or four short, branching, antler-
like processes. Collar usually blue and divided laterally with a fillet at the incision.
Collar setae (fig. 38.3.1) with a hispid boss at the base of the blade which becomes
more spinulose distally. Thoracic uncini (fig. 38.3.J, k) with 15-18 teeth in a
single row. Abdomen two to three times as long as the thorax and has 200-300
segments.
Type locality : \Vcst Indies.
Records: Natal (30/30/i) ; Mocambique (26/32/i).
Distribution : ^Vest Indies ; Suez (i) ; Tropical Indian Ocean ; Japan (s) ;
New Caledonia.
Spirobranchus tetraceros (Schmarda, 1861)
(fig. 38.3.1-n)
Pomatoceros tetraceros Schmarda, 1861 : 30, pi. 21 fig. 179.
Spirobranchus tetraceros : Johansson, 1918: 7, fig. 2.
Spirobranchus scmperi Morch, 1863 : 405, pi. 2 figs. 24-25.
Length up to 30 mm. Tube stout and ridged. Branchial lobes flattened with a
single whorl of radioles. Opercular stalk (fig. 38. 3. n) broad with the wings dentate
on their inner distal margins. Operculum arched or flat with four long branching
antlcr-like processes. The anterior pair are deeply branched and the branches
may separate forming six processes. Collar not divided ventrally but produced
8o4 1H)LVC:IIAF. TA Ol- SOI'TIIKRN AFRICA
forward as a loiiguc-sliapcd membrane. Thoracic uncini (fig. 38.3.!, m) witii 13-16
lecth in a single row and a large expanded gouge.
Type locality: New Sciuth Wales.
Records: Mcx ainbiqiie (■26321); .Madagascar (s).
DisTRiBV HON : Tropical Indo-west-Pacific from Zanzibar (i) to the Great Harrier
Reef (i).
HYDROIDES (■unmriis, 1768
(including EUPOM ATI'S Philippi, 1844)
Body sxnmielrical with se\en thoracic setigers and numerous abdominal ones.
Tube circular in section, often erect. Opercular stalk slender and without wings.
Operculum with a basal funnel effused radii from which arises a central crown of
several horny spines, dollar split laterally forming a \-entral lobe and a ])air of
lateral lobes which continue back as thoracic membranes past the end of the thorax,
curve round and unite ventrally. Collar setae include bayonet-shaped forms, with
two or more coarse teeth at the base of the blade and a few fine capillaries. Noto-
setae of setigers 2-7 are narrow-bladed lapillaries. Thoracic uncini with about
six coarse teeth in a single row and a basal gouge. Abdominal imcini similar.
.Abdominal capillaries with short, triangular blades with a toothed margin.
Type species: Hydroidcs nonegica Gunnerus, 1768.
Kf-v to Species
1 .Spines of opercular (Town all equal in sizr ........ 2
- One or more spines of ojKTCular crown enlarged ....... 4
2 Spines of opercular crown wiili tuo to four pairs of lateral spinules. (Radii of funnel hluiil)
(tig. 38. 4. c) H. norvegica (p. 805)
- Spines of opercular crown without lateral spiniiles ....... 3
3 Spines of opercular crown with curved, pointed ends (fig. 38.4.h) H, uncinata (p. 805)
- Spines of opeixular crown with anchor-shaped ends (fig. 38. 4. k) . H. lunuli/era (p. 807)
4 Only one enlarged hooked spine in the opercular crown ...... 15
- Two to three enlarged spines in the opercular rrowTi (fig. 38.4.ni) . H. raluniiana (p. 807)
5 Radii of opert ular funnel with simple pointed ends ....... 6
- Radii of open ul.n fmniel uilli o\ oid or am hor-shaped ends (hg. 38. 4. 1)
//. heteroceros (p. 807)
6 Enlarged Ijook with ,1 pair of lateral hooklets. ( )|)ercular funnel oval and slanting (lig.
38.4.01 .......... H. monoceros (p. 808)
- I'.nlarged hook simple. ( )piercular fiumel ratlially symmetrical ..... 7
7 .Smaller spines of operetilar crown end in two outwardly directed hooklets (fig. j8.,(.q'i
H. bifurcata (p. 808)
- Smaller spines of opercular crown end in one tjutwardly directed and one inwardly
directed ho.>klet (fig. 38.4. ti //. rfi'/joma (p. 809)
SERPULIDAE 805
Hydroides norvegica Gunnerus, 1 768
(fig. 38-4-a-g)
Hydroides norvegica Gunnerus, 1768: 53; Fauvel, 1927: 356, fig. 122 i-o.
Body (fig. 38.4.b) slender and up to 30 mm. long. Tube fragile, often erect.
Opercular funnel (fig. 38.4. c) radially symmetrical with about 50 blunt radii.
Opercular crown of 10-20 equal spines each with two to five pairs of lateral spinules,
a pointed tip and sometimes one to two inner spines at the base. Collar incised
laterally with a small fillet in the notch. Dorsal lobe of collar continuous with the
thoracic membrane. Collar setae include plain capillaries (fig. 38. 4. e) and bayonet-
like setae (fig. 38.4. d) with serrated shaft heads culminating in a pair of stout
bosses at the base of the blade which becomes spinulose distally. Subsequent
thoracic notosetae are limbate capillaries. Thoracic uncini (fig. 38. 4. f) have a single
series of six to seven teeth and a slight gouge. Abdominal uncini smaller, with five
teeth. Abdominal ncurosetae have expanded, truncate and toothed blades (fig.
38. 4. g). Tubes cause fouling on slow moving ships.
Type locality : Norway.
Records: South West Africa (26/14/d to 28/16/s) ; Cape (from 31/16/d to
34/18/s, d and 33/27/s) ; Natal (29/31/s, d).
Distribution: Cosmopolitan apart from polar seas (s, d, vd). Also world wide
on ships' hulls.
Hydroides uncinata (Philippi, 1844)
(fig. 38.4.h-i)
Eupomatus uncinatus Philippi, 1844: 195, pi. 6 fig. 9.
Hydroides uncinata: Fauvel, 1927: 357, fig. 122 a-h.
Length up to 60 mm. Opercular funnel (fig. 38.4.11) radially symmetrical with
about 30 radii ending in triangular points. Opercular crown of 8-12 equal horny
spines curving towards the centre. Spines all similar in size with plain hooked tips.
All spines are broad basally and may have a spinule low down on the inner side.
Bayonet setae with a pair of simple bosses at the base of the smooth blade. Tube
stout, often adnate on shells.
Type locality : Mediterranean seas.
Records: Mocambique (23/35/s).
Distribution: Bay of Biscay (i, s) ; North Carolina (s) ; Morocco (s) ; Senegal
(s) ; Congo (i) ; Mediterranean ; Japan.
8o6
I'Ot.VtiHAK lA OF .S(.>L: 1 IIKRN Al RKIA
I'r<;. ;^fJ.4. HvJnn'/i-s noricgica. (a) Tubes, (b) Entire worm (five times life size), (c) Oper-
culum, ft)} Bayonet-seta from collar, (e) Normal thoracic capillary, (f) Thoracic uncinus.
{<'.) Abdominal neuroseta. Hydroides uncinata. (ii) Operculum, (i) Part of bayonet-seta.
Hydroidis lunuli/fia. (]) Part of bayonet-seta, (k) Operculum. Hydroidts heteroceros.
(i.) Opert ulum. Hydroides ralumiana. {m. m') Operculum. (n) Part of bayonet-seta.
Ilydnndts inonocero^. (o) Operculum, {pj Part of bayonet-seta. Hydroides bifiircata. (Q, Q^)
Operculum, (k) Part of bayonet-seta. Hydtoides dif^oma. (s| Operculum. (t) Opercular
spine.
SERPULIDAE 807
Hydroides lunulifera (Claparede, 1868)
(fig. 38.4-J-k)
Eupomatus lunulifera Claparede, 1868: 441, pi. 31 fig. 3.
Hydroides lunulifera : Fauvel, 1927: 358, fig. 122 p-s.
Body about 20 mm. long. Opercular funnel (fig. 38. 4. k) radially symmetrical
with about 40 radii having incurving pointed tips. Opercular crown of 10-14
similar horny spines, each smooth, rather flattened and of uniform width except at
the end which is abruptly expanded like a very broad T or crescent moon. Tube
stout, rugose, often adnate. Bayonet setae (fig. 38.4.J) w-ith a pair of simple blunt
bosses at the base of the smooth blade.
Type locality : Italy.
Records: Natal (29/31/s).
Distribution: Mediterranean; Suez Canal ; Madras (i).
Hydroides ralumiana Augener, 1927
(fig. 38.4.m-n)
Hydroides (Eupomatus) ralumianus Augener, 1927: 150, fig. 5; Fauvel, 1947: 85.
A small species about 10 mm. long with a bi-ridged tube. Opercular funnel
(fig. 38. 4. m) almost symmetrical with 25-30 short pointed radii. Opercular crown
widi 8-10 unequal liooks all curving inward like a clenched fist. The two to
three largest hooks are smooth and curve over the others but the seven to eight
smaller ones each have an external knuckle-like projecdon at the point of inflection.
There may be smaller basal spinules. Bayonet setae (fig. 38.4.n) with a pair of
simple bosses at the base of the smooth blade.
Type locality : New Pomerania.
Records: Mocambique (26/32/i, 24/34/s, and 23/35/s).
Distribution: Pacific (New Pomerania and New Caledonia).
Hydroides heteroceros (Grube, 1868)
(fig. 38.4.I)
Serpula {Eupomatus) heteroceros Grube, 1868 : 639.
Hydroides heteroceros : Pixell, 1913 ; 75, pi. 8 fig. 2 a-c ; Fauvel, 1953 : 459, fig. 24. C.
Serpula (Hydroides) uncinata: (non Philippi) Gravier, 1902 : 114, pi. 8 figs. 286-7, text-figs. 463-466.
Body up to 40 mm. long ; tube thick and rugose with faint longitudinal ridges.
Opercular funnel (fig. 38. 4. 1) composed of 30-50 radii whose tips are T-shaped
with a pair of lateral points. Opercular crown with seven to nine inwardly cur\'ing
spines. One spine is larger than the rest and is shaped like a plain hook without
lateral spines. The others have a pair of lateral spines and a single inwardly directed
spine near the base. Branchiae with 16-18 radioles ending in long naked dps.
Collar large and incised laterally ; ventral lobe entire. Collar setae include stout
8o8 I'OI.VCiHAEI A OF SOU HI KRN Al RK: A
bayonet setae with a pair of plain, blunt bosses at the base of the tapered blade and
slender forms without obvious blades. Notosetae of sctigers 2-6 as normal capill-
aries. Thoracic uncini with five to seven teeth, .\bdominal uncini with six to scN'cn
teeth. Abdominal capillaries with tranvesrely expanded and serrated tips.
Type LOCALrrv : Red Sea.
Records: Madagasdar (25/46/s).
DiSTRiBniON' ; Tropical Indian Ocean (i, s).
Hydroides monoceros Gravier, 1908
(fig. 38.4.0-p)
Serpula (Hnlroitiei) monoieros Graxier, 1906a : 115, pi. 8 tig. 288, text-figs. 4(17-472.
Bcidv about 20 nmi. long. Operculai funnel (fig. 38. 4.0) oval and slating and
composed of 16-20 stout, pointed radii of unecjual size. Opercular crown of six to
eight small spines and one very large dark hook with a pair of booklets at its tip.
C!ollar incised laterally. C^ollar setae include fine capillaries and bayonet setae
(fig. 38. 4. p) with a pair of plain bosses at the base of the smooth blade. Thoracic
uncini xvith five to se\cn teedi and an ob\ious gouge. Tube stout, rugose and
adnatc.
Type LOCAi.rrv : Red Sea.
Rec:ords: Mocambique (26/32/i).
DiSTRiBUTio.x : Red Sea (i) ; India ; Cleylon ; Zanzibar ; Mocambique Is. ;
Gambier (S. Pacific)
Hydroides bifxircata (Pixell, 191 3)
(fig- jST-q-i'i
Eujiomnlui hijmcalm Pixell, 1913 : 78, pi. 8 fig. 5.
Hydroidei btfimala: Day, Kiji : 64, fig. 8 f-g.
Body about 30 mm. long. Opercular funnel (fig. 38. 4. q) of 40 pointed radii.
Opercular crown of one enlarged hook and 10 smaller equal spines (fig. 38. 4. q')
each ending in a double hook whose di\crgent prongs both point outwards and each
with an inwardly directed spinule at the base. Each branciiial lobe \vith about 20
radioles. CloUar deeply incised laterally with a small fillet in the notch. Bayonet
setae (fig. 38.4.rj with four bosses at the base of the blade, two larger and two
smaller.
Type i.ocAi.rrY : liuli.in Ocean.
Records: Xat.il (29/31/1); Mocambique (26/32/i).
DiSTRiEi Tlo\ : Tropical Indian Ocean.
SERPULIDAE 809
Hydroides dipotna (Schmarda, 1861)
(fig. 38.4-s-t)
Eupomaliis dipoma Schmarda, 1861 : 29, pi. 21 fig. 177.
Eupomatus spinosus Pixell, 1913 : 78, pi. 8 fig. 5.
Body about 30 mm. long. Opercular funnel (fig. 38.4.5) sUghtly oblique with
30-40 long, pointed radii. Opercular crown with nine to eleven tall spines including
one enlarged hook and eight or 10 smaller spines each with a terminal double
hooklct of which one prong points inward and one outwards (fig. 38.4.1). An
inwardly directed spinulc or curved lobe may be present at the base of each spine.
Each branchial lobe with 10-14 radicles ending in short tips. Collar entire and edged
with brown, the thorax red with black bars marking the uncigerous rows. Collar
setae include a few fine capillaries and numerous bayonet setae with smooth blades
and a pair of simple bosses at the base. Thoracic uncini with six to seven teeth in a
single row. Tube stout and rugose.
Type locality : Cape of Good Hope.
Records: Cape (from 33/18/i to 34/18/i, s and 33/26/i, s).
Distribution : Mediterranean ; Senegal ; Angola.
SERPULA Linnaeus, 1 758
Body symmetrical with seven thoracic setigers and numerous abdominal ones.
Operculum funnel-shaped and soft with marginal serrations formed by the ends of
numerous fused radii. Opercular stalk smooth. Collar setae include capillaries and
bayonet setae with two to four conical bosses at the base of a hispid blade. Sub-
sequent dioracic notoscate are plain limbate capillaries. Uncini with only a few-
large teeth in a single row. Abdominal capillaries trumpet-shaped or T-shaped.
Type SPECIES : Serpula vermicularis Linnaeus, 1767 ("designated" Hartman, 1959:
592).
Serpula vermicularis vermicularis Linnaeus, 1767
(fig- 38-5-a-l^)
Serpula vermicularis hinnacus, 1767: 1266; Fauvel, 1927: 351, fig. 120 a-q.
Body up to 70 mm. long. Operculum (fig. 38. 5. a) a deep symmetrical funnel
formed about 40 fused radii with blunt tips. Thirty to forty branchial radioles in a
semi-spiral united by an obhque web for about one-fifth of their length. Collar
trilobed with a small fillet in the lateral notch. Thoracic membranes extend the
full length of the thorax. Collar setae include fine capillaries (fig. 38.5.C) plus stout
bayonet setae (fig. 38. 5. b) which have two bosses at the base of the blade. Thoracic
uncini (fig. 38. 5. g) with six teeth of increasing siee in a single row. .\bdominal
uncini similar in shape but with four to eight teeth. Most abdominal neurosetae
(fig. 38. 5. f) have shafts with a denticulate blade set transversely to form a rough T.
8io POLVCHALTA OF SCJUTHKRN AFRICA
Xeurosctac at the end of the abdomen (fig. 38. 5. c) are slender wingless capillaries
with cur\cd tips. Tnbe (fig. 38. 5. h) circular in section often ridged externally.
Type loi:ai itv : Western Europe.
Records: Cape (from 32/18/d to 34/20/i, s and 34/23/c, s, d, to 32/28/i) ; Natal
(2931/1); Mocambique (26/32/i).
Distribution : Cosmopolitan (i, s, d).
Serpula I'ermicularts echinata Linnaeus, 1788
(Hg- 3«-5-i)
Sfrfiiila rchinntti lAnn:iC\i^, i/HB: 3744.
Sfrfnila vermiculiuis echinnta ; Famcl, 1927: 352.
Generally similar to the nominate form but the tube (fig. 38.5.1) has fi\e to seven
longitudinal ridges bearing recurved teeth.
Type locality : Mediterranean Sea.
Records: Clape (34/18, s and 34 2i,s).
IJiSTRiBtTiox : Mediterranean.
FICOPOMATUS ^nMhnn, 1921
IJody sN'mmelrical with seven thoracic and numerous abdominal segments. Oper-
culum pear-shaped to conical, soft or chitinous but without spines. Opercular
stalk without wings. Collar setae include some with a toothed boss at the base of
the blade and others with simple serrated blades. Subsec[uent thoracic notosetae
are limbate capillaries, there being no setae of Afmniatiis with denUculate tips.
Uncini with relatively few large teeth in a single row. Abdnminal capillaries have
geniculate toothed blades.
Type spe(;ies : Firojximaliis maoodon .Southern, 1921.
Key to Spiciks
I Thoracic membranes reach past setiger 7. Operculum fig-shapcd . . F. macrodon*
- Thoracic membranes stop at setiger 3. Operculum an elongate cone (fig. 3fi-5.j) . F. capensis
Ficopomatus capensis Day, 1961
(fig- 3^-50"")
Ficojtuwiilii\ fnpfn\i\ Day, iffGi ; 552, fig. 17 h-n.
Body 15 nun. long. Tube adnate and tiiangiihu- in section. Operculum (fig.
38.5.J) an elongate chitinous cone without internal septa. Collai deeply incised
laterally and the dorsal lobes extend back as thoracic membranes which end at
setiger 3 so tint there is no free ventral membrane at the end of the thfuax. Collar
setae of two typrs : 'a) large setae (fig. 38. 5. k) with stout shafts and a boss with three
SERPULIDAE
8ii
Fig. 38.5. Serpula vermkularis. (a) Entire worm (five times life size), (b) Bayonet-seta.
(c) Slender collar seta, (d) Normal thoracic capillary. Capillary from end of abdomen.
(f) .Abdominal T-shaped neuroseta. (c;) Thoracic uncinus. (h) Tube. Serpula vermkularis
echinata. (l) Part of lube attached to Relepora. Fkoporrmlus capensis. (j) Operculum.
(k) Bayonet-seta, (l) Abdominal capillary, (m, n) Thoracic uncinus. Mercierella enigmatka.
(o) Thoracic uncinus. (p) Stout ts-pe of collar seta, (q) .Abdominal capillary, (r) Tubes,
(s) Anterior end.
8i2 POLVCHAEIA OV SOrTHKRX AIRICA
to fi\c icclh at the base of the narrow serrated bhide, (b) slender geniculate setae with
hispid blades. Notosetae of sctigers 2-7 are normal limbate capillaries. Tlioracic
uncini i^lig. 38.5.ni, n) with a single row of eight teeth the last of which extends
beyond the blunt recurved basal prow. Abdominal capillaries (fig. 38. 5. 1) are
geniculate with minutely toothed blades.
Type locality: Agulhas Bank, Soutii Africa.
Records: Cuipe (34 21 s).
Distribution' : A single record only.
MERCIERELLA Fauvel, 1923
Operculum saucer-shaped and crowned with concentric rows of horny spines.
Opercular stalk wingless. Clollar setae include fine capillaries and others with
toothed bases. Subsequent thoracic notosetae are limbate capillaries. Uncini witJr
a few teeth in a single row, the first being enlarged and gouge-like. Tube cylindrical
with a series of rings representing earlier trumpet-shaped mouths.
Type species: Mercicrella cnigmalka Fauvel. 1923b.
Mercierella enigmatica Fau\el, 1923
(fig. 38.5.0-s)
AlcriiercUa cnis^matiai F.invel, 1923b: 4'.i4. tig. i ; Fauvel, 1927: 360, tig. 123 a-o.
Body up to 25 mm. long. Branchial lobes each \vith six to ten stout radioles ending
in naked tips. Operculum (fig. 38. 5.5) oblique, concave and edged with about 25
dark chitinous spines.* Opercular stalk smooth and triangular in section. Gollar
large, reflected back but not incised, dollar setae include fine capillaries and
numerous stout forms (fig. 38. 5. p) with bluiu teeth set in twf) rows along the tapered
blades. Subsequent dioracic notosetae are capillaries with finely hispid blades.
Uncini (fig. 38.5.0) with a .gouge and a single row of five to seven teetli. Abdominal
ncurosctae (fig. 38. 5. q) are geniculate ca]3illaries with denticulate blades. Tube
(fig. 38. 5. r) cylindrical with a trumpet-shaped mouth and Uie position of earlier
mouths is shown by a series ol rings. Twisted masses of tubes are attached to hard
substrata in estuaries.
'Fype locality : Caen, France.
Records: Oape (from 33/18/e to 32/28/e) ; Natal (from 30/30/e to 28/32/e).
DlSTRiBUTiox : World wide in warm estuaries.
VERMILIOPSIS Saint-Joseph, 1894
Operculum a cylindiical or conical In any caj) with internal septa. Opercular
stalk without wings. Collar setae are limbate capillaries. Body of seven thoracic
*Fau\'--l dosciiljcd llic lAiroptan furm as lia\ing tliree rows of sj)iiirs.
SERPULIDAE 813
setigcrs and numerous abdominal ones. Thoracic notosetae include limbate capil-
laries and setae of Apomalus with distally toothed blades. Thoracic uncini with a
single row of teeth, the first of which is enlarged and blunt. Abdominal capillaries
are geniculate with toothed blades.
Type species : Serpida infundibulum Linnaeus, 1 788.
Key to Species
1 Thoracic membranes stop at setiger 2 (fig. 38.6.3) ....... 2
- Thoracic membranes continue past setiger 2 ....... . 3
2 Operculum smoothly conical with numerous (10-20) septa. Tube with four to five low
longitudinal ridges .......... V. pygidialis*
- Operculum not smooth but with about five steps corresponding to the septa. Tube with
three denticulate ridges (fig. 38.6.f) ....... V. babylonia
3 Thoracic membranes continue to setiger 5. Operculum bluntly conical with three to four
internal septa (fig. 38.6.!) ......... V. glandigerus
- Thoracic membranes continue to setiger 7. Operculum terminating in a point (fig. 38. 6. k)
V. acanthophora*
Vermiliopsis babylonia sp. nov.
(fig. 38.6.a-f)
Vermiliopsis pygidialis : (non Willey) Day, ig6i : 552.
Body pale, about 4 mm. long. Tube (fig. 38. 6. f) sinuous, attached throughout
and with thiee well marked longitudinal ridges each bearing a series of rectangular
teeth. Operculum (fig. 38. 6. a) like the tower of Babylon with seven successive
cylindrical towers each with vertical walls and steps with an outer flange marking
the position of the internal septa. All but the youngest or basal annulus are yellow
and heavily chitinised. Branchial lobes each with seven radioles. CoUar apparently
continuous laterally and ventrally but with a dorsal gap. The sides of the collar
continue as rounded flaps to setiger 2. Collar setae (fig. 38.6.b) are four limbate
capillaries and four smaller slender capillaries. Later thoracic notosetae are limbate
capillaries plus two to three setae of Apomatus (fig. 38. 6. c) in each of the last four
bundles with the blade abruptly narrowed and toothed for most of its length.
Thoracic uncini (fig. 38. 6. e) with a single series of 1 1 teeth and a well marked gouge.
Holotype: B.M. (N.H.) Reg. No. 1966. 26.8.
Type locality: \'ema Sea Mount (32^S/IO°E).
Records: Cape (32/i0;s; 34/18/s).
Vermiliopsis glandigerus Gravier 1908,
(fig. 38.6.g^)
Vermiliopsis glandigerus Gravier, 1908: 121, pi. 8 figs. 290-291, text-figs, 476-481.
Length 25 mm. Operculum (fig. 38.6.1) ovoid to conical, horny and with three
to four septa internally but smooth on the surface. Collar deeply incised laterally
to form a square ventral lobe and a pair of large lateral ones which continue back
as thoracic membranes to end at setiger 5. Collar setae are long, limbate capillaries
fii4 POLYCHAETA OK SOUTHKRX AIRICA
and fine capillaries. Subsequent thoracic notosetae are of three types : (a) elongate
limbate capillaries (b) setae of Apomatus with the distal three-quarters of the blade
very finely toothed and (c) very fine wingless capillaries. Thoracic uncini (fig.
38. 6. h) arc quadrangular plates with 13-14 teeth preceding the enlarged rectangular
terminal tooth. Abdominal neurosetae (fig. 38. 6. g) are finely toothed geniculate
capillaries and, in posterior segments, very elongated capillaries. Tube cylindrical,
slightly rugose.
Type locality: Djibouti, Gulf of Aden.
Records: Cape (from 33 ly/s and 34/18,1, s, d to 32/j8,i).
DisTRiBLTiON : Western Africa (Annobon Is.) ; Red Sea (s) ; tropical Indian
Ocean, li, s, d) ; Tasmania (d).
Vermiliopsis acanthophora Augcner, 1914
(,fig. sB-b.j-k)
I'trmilio/ins aianlkaplwra .Vugener, 1914: 153, pi. i figs. 122-124, text-fig. 19; Fauvel,i953: 467,
fig. 243 c-c.
Length jo mm. Tube (fig. 36.6.J) sinuous, attached throughout, surface with four
low ridges and marks of successive apertures. Opercular stalk broad, often
wiinkled ; operculum a short broad cone (fig. 38.6. k) with three to four external
ridges and often a terminal thorn-hkc spike. No internal septa. Cbllar with a large
ventral and two lateral lobes which continue back as thoracic membranes to the
end of the thorax (setiger 7) but are not continuous across the ventrum. Collar
setae are limbate capillaries. Subsequent thoracic notosetae include smootli-bladed
capillaries and setae of Apomatus with three-quarters of the blade minutely toothed.
Thoracic uncini with about nine to ten teeth and a terminal gouge. .Abdominal
neurosetae geniculate with minutely toothed blades. A glandular patch at the end
of the abdomen.
Type locality : Sharks Bay, Western .Australia.
Records : Xot recorded from southern Africa.
Distribution : Tropical Indian Ocean (Mombasa (i) ) ; Ceylon (i) ; \\'cstern
Australia (ij.
NEOVERMILIA Day, 1961
Body svmmelrical with seven thoracic and numerntis abdominal setigers. Oper-
culum spiierical or funnel-shaped ; opercular stalk without wings. Branchial lobes
sometimes spiral. Collar setae are limbate capillaries similar to those of the following
thoracic segments. Xo setae o^ Apomatus. Thoracic uncini with a few large teeth in
a single row. Abdominal neurosetae are geniculate capillaries with finely toothed
blades.
Type species: .Xcoicrmilia capetnis Day, 1961.
SERPULIDAE
815
Fig. 38.6. Vermiliopsis babylonia n.sp. (a) Anterior end. (b) Limbate collar seta, (c) Seta
of Apomalus. (d) .'\bdominal capillary, (e) Thoracic uncinus. (f) Tube. Vermiliopsis
glandigerus. (g) .Abdominal capillary, (h) Thoracic uncinus. (i) Operculum. Vermiliopsis
acanthophora (modified from .-Xugener, 19 14). (j) Tube, (k) Anterior end. Neovermilia
capensis. (l) Collar seta, (m) Thoracic winged capillar)', (n) .\bdominal capillary, (o)
Operculum, (p, q) Thoracic uncinus. (r) .Xnterior end.
8i6 roLVcniAEi A or soliiii.r.\ A1'RK:A
Key to Species
I OpcrcLiIuni sj:)lu'rical .......... N. falcigera*
- (_)i>ercniiim runnel-shapcti (fig. :i8.6.()) ....... N, capeyisis
Neovermilia capensis Day, 1961
(fig. 3S.6.1-r)
.\eovcrmilia tii/mtsis Day, 1961 : 551, fig. 17 a-g.
Length up to 30 mm. Operculum (fig. 38.6.0) a siuiKjth, shallow rumiil witlnuu
radii. Opercular stalk smooth and wingless. Branchial lobes spiral with four whorls
of radioles. Xo palps. CloUar (fig. 38. 6. r) widel)' o]5en dorsally, incised laterally and
forming a ventral lobe and two lateral lobes which continue back as tlmracic
membranes to the end of the thorax and unite vcntrally. C'oUar setae (fig. 38. 6.1)
smaller than subsequent notosctac and have narrower blades. Notcisctae of sctigers
2-7 with rather broader blades (fig. 38. 6. m). No setae oi Apomalus. Thoracic uniini
(fig. 38. 6. p, q) quadrangular with a single series of seven teeth of decreasing size
above the main fang. Abdomen with .ibout 60 segments. Abdominal neurosetac
are geniculate capillaries with finely toothed bl.ides (fig. 38.6.0). J'ube unknown.
Type loc.xlity : False Bay, South Africa.
Records: Cape (34/i8,s).
DisTRiBUTio.N : A single record.
Subfamily FILOGRANINAE Rioja, 1923
Body symmetrical with five to twehe thoracic segments. Prostomium sometimes
distinct and may bear eyes. Palps sometimes present. Operculum either absent or
poorly developed with tin- stalk bearing later. d ])innules similar to those of other
radioles. Thoracic luicini may have more than one row of teeth.
Records from southern Africa
Filiiniatm implfxd Hvrkclcy ..... .).4Ca, 5iClsd
as Salmacina dyiteri (Huxley) .... 26Ai
as Salmacina dviteri var. iiii'iii\ttiiis C^Lqjarede . .].o.\i
Prolula bispiralis (Savigny) ..... ;;8Ca, 35Ca, fjiC^s
as Protula tiibularia var. capfihis (non .\h Intosh) . l3C'i
I'rotula tuhidaria (\hw\.\n\) ..... 45X1, 5iC;sd
as Prntula capen\h Mcintosh .... loCld
Piolida tiihiilnria annniala D:\y . . . . iK'i
I HE MAIN DIAGXOSIIC ( :H ARAL. lERS
The operculum may be entirely absent as in Piutula or formed by a slightly modified
radiole which retains pinnules on its stalk but becomes expanded at its apex to form
a soil globular or ( up-sha]jed structure. The opercuhmi is ni\rr large or (dli ified
SERPULIDAE 817
and in Filograna which reproduces by fragmentation the common growth phase lacks
an opercuktm and has been regarded as a separate genus {Salmacina) even though
it occurs in the same colony with the operculatc form {Filograna).
Thorax. The number of thoracic setigcrs varies from five to twelve in different
genera and m Filograna the number varies from one individual of a colony to another.
Setae. Collar setae are always present and may be represented by simple capillaries
or limbate capillaries with a dentate fin at the base of the blade. Subsequent thoracic
notopodia often have setae oi Apomatus among the smooth-bladed capillaries particu-
larly in posterior segments. Uncini are provided with one to four rows of numerous
small teeth and an elongated terminal gouge. In the genus Protula they may be
lacking not only from the collar segment but the next two to three segments as well.
Abdominal uncini are similar to those of the thorax and abdominal capillaries aie
geniculate with finely serrated blades but become more elongate and often smooth
towards the end of the abdomen.
Key to Genera
1 Five thoracic setigers. Operculum vase-shaped with strengthening ribs . JOSEPHELLA*
- More than five thoracic setigers. Operculum either absent or membraneous -2
2 Collar setae with a dentate lobe at the base of the blade. (Operculum either absent or
cup-shaped. .Animal gregarious forniing interlacing tubes) . FILOGRANA (p. 817)
- Collar setae are narrow-bladed capillaries without a dentate lobe .... 3
3 Operculum a globular vesicle ........ APOMATUS*
- Operculum absent PROTULA (p. 818)
FILOGRANA Berkeley, 1835
Sinall gregarious worms whose twisted tubes form a lacy coralliform mass. Body
symmetrical with six to twelve thoracic setigers and about 20 abdominal ones.
Branchial lobes with few radioles which are not united by a basal web. The dorsal
pair of radioles retain their pinnules but sometimes bear cup-shaped membraneous
opercula at their ends. Two palps. A prominent prostomium with a pair of eye-
spot.s. \ well developed collar. Collar setae with a dentate expansion at the base of
the blade. Uncini rectangular with two to fourr ows of teeth and an enlarged
terminal tooth or gouge. Abdominal capillaries geniculate with minutely dentate
blades.
Type species: ''Serpula implexa Turton"' (Berkeley, 1835) = Serpula complexa
Turton, 1819.
Filograna implexa Berkeley, 1835
(fig. 38.7.a-h)
Serpula filograna h'lnnaeus, 1766: 1265; Berkeley, 1828: 230, pi. 18 fig. 3.
Serpula cnmplexa Turton, 1819: 153.
Filograna implexa Berkeley, 1835: 427; Fauvel, 1927: 376, fig. 129 a-b.
Protula dysleri Huxley, 1855 : 113, figs. I-i i.
Salmacina dysteri \ Fauvel, 1927: 377, fig. 129 c-k.
8i8 l'ol.^■e;llAl: 1 A oF' .sofrnKRiX ai-ru:a
Body (fig. 38.7.c'l about 4 mm. long wilh 25-35 setigcrs and pink when alive.
Branchial lobes carh wilh four radioles of which the dorsal one on each side may end
in a small bixalve operculum 1 fig. 38. 7. f). In the Saliiiacina form opercula are absent.
Collar setae include normal capillaries and geniculate forms with a dentate expansion
at the base of the blade (fig. 38.7.g). Subsequent thoracic notosetae include smooth-
bladed capillaries (fig. 38.7.11) and setae oi Apmnatus (fig. 38.7.1) with finely toothed
blades. Thoracic uncini (fig. 38. 7. d, e) with two to four rows each with six teeth
and a large terminal tooth or gouge. Abdominal capillaries (fig. 38.7.J) geniculate
with dentate blades. The worms are gregarious and reproduce asexually by trans-
verse fission, young forms having few segments and large ones up to 12 thoracic
setigers. They commonly occur in large coralliform masses (fig. 38.7.3) formed of
anastomosing biuidles of tubes (fig. 38. 7. b).
Type lo<:ality : \Ve\nioiuh, England.
Records : Cape (from 29 16 i to 34 23 i, s, d and 32'28'i).
Distribution : Atlantic from Greenland (d) and .Sweden (d) south to New
England (i), the Gulf of Mexico (s) and tiopical western Africa (i, s) ; Mediterra-
nean (s) ; Red Sea ; India ; S.\\'. Australia (s) ; Tasmania (d) ; New Zealand (s) ;
Japan.
PROTULA Montagu, 1804
Body symmetrical with seven thoracic and numerous abdominal segments. No
operculum. Branchial lobes flat or spiral with several whorls of radioles. Thoracic
membranes continue to the end of the thorax and form a free flange ventrally. Collar
setae as simple capillaries. Subsequent thoracic notosetae include limbate capillaries
and sometimes setae of Apaniatui. Uncini with numerous teeth and long slender
gouge. Abdominal neurosctae arc dentate capillaries with either sickle-shaped or
bayonet-shaped blades. Tube cylindrical, often erect.
Type species: Protula tubularia Martini, 1776.
Rev to Species
1 Branchiae spiral with a central axis and three lo ten when Is of radioles . . P, bispiralis
- Brinchiae not spiral, no central axis ......... j
2 Dcntate-bladed abdominal capillaries with a notch at the base of tlic bhidc
P. tubularia anoinala
~- Dentate-bladed abdominal capillaries without a notch at the base of the blade P. t. tubularia
Protula bispiralis (.Savigny, 1820)
(fig. 38.7.k-n) '^
Strpula bispiralis Savigny, 1820: 7*1.
pT'.ilula bi'ipiralis ', Fauvel, I92'2 : 498, fig. a-f; Day, 1933: 44'|.
A lar.^c s])rci(.'s reaching a lenejth of 180 mm. and with nxl <j:ills. Branchial lubes
(fig. ^B.j.mj spiral wilii a central axis and three lo ten whorls (ti" radioles united by a
SERPULIDAE
8i9
Fig. 38.7. Filograna impUxa. (a) Colony (natural size), (b) Part of same showing individual
tubes, (c) Entire worm (30 times life size), (d, e) Thoracic uncinus. (f) Operculum of
Filograna form, (g) Collar seta, (h) Thoracic winged capillary, (i) .Seta of Apomatus.
(j) Abdominal capillary. Prolula bispiralis. (k) Thoracic capillary, (l) .Abdominal
capillary, (m) Anterior end. (n) Thoracic uncinus. Protula tubularia. (o) .\nterior end.
(p) Seta of Apomatus. (q) Thoracic uncinus. 1 r) .Abdominal capillary. Prolula tubularia
anomala. (s) Abdominal capillary.
8-H) i'()i.\c;ii.\i',r.\ oi' SOL' 1 iii'-.RN \iric:a
web lor (Uic-lliinl lluii length. CliilLir im isnl NriUni-latcrally lurniiiig twi) lateral
k)bi\s and a median ventral one. Thoraeie membranes strengthened at segmental
intervals. Thoraeie notosetac are all smootli-bladed capililarcs (fig. 38.7.!;) and
setae oi Apnmalus are absent. Uncini (fig. 38. 7. n) with a single row of 15-20 teeth;
they are lacking from the first three thoracic sctigers. Abdominal ca])illarii-s (fig.
38. 7. 1) with finely dentate blades. Tube rugose mainly adnate.
Typk locai.hy: Indian Seas.
Records: C:ape (from 3418 i, s to 3.2/28/1).
Distribution: Indian Ocean.
Protula tubularia tubularia (.Montagu, 1803)
(fig. 38.7.0-r)
Serpulii lubiiltirii] Montagu, 1803; 513.
Protuld liibularia: Fauvcl, 1927: 382, fig. 130 a-1.
Length up to 50 mm. Branchial Icibes (fig. 38.7.0) senii-circular with a single
whorl of long, wideK" Hanged radioles. Collar incised \entro-later,dly to form a
ventral and t\\<i hiter.d lobes. Thoracic notosetae arc smooth-bladcd capillaries in
the first three to four segments and smooth-bladed capillaries plus setae o\' Apnmnlus
in the remainder. Individual setae o( Apnmaliis (fig. 38. 7. p) with a frilly edge bearing
very minute teeth. Lhicini (fig. 38.7.0]) with a single row of 20-25 tt-cth ; the first
row is on setiger 3 or .). Blades of abdominal capillaries (fig. 38. 7. r) dentate and
frilly but smoothly continuous with the shaft.
Type LOf;.M,ri v : Engl.md.
Records: C:ape (31/16/d and 35/i8,d) ; Natal (29/31/s).
Distribution : .-Xtlantic from Grecnaind (d) and Scotland (s) south to the English
Channel (s), the Gulf of .Mexico (d), Morocco (s, d) and Senegal (s) ; Mediterranean
(s) ; Intli.ui Oce.ui ; .\e\v Caledonia ; J.ip,in.
Protula tubularia anomala Day, 1955
(fig. 3B.7.S)
PtoIuIh tubularia var. aiioiiiala Day, 1955 : 449, tig. 8 I-m.
This is a small form about 15 mm. long which is generally similar to the stem
firm but has a notch at the base of the dentate blade of the abdominal capillaries
(fig. 38.7.S).
Type loc.m.iiy: Langcbaan Lagoon, South Africa.
Rec:ords : Cape (33/18,1 and 34/23/i).
DisTRiBt'TKjN : No Other records.
GLOSSARY OF TECHNICAL TERMS
821
GLOSSARY OF TECHNICAL TERMS
abdomen
achaetous
acicular seta
aciculum(a)
anal cirrus(i)
antenna
apodous segment
arborescent
aristate seta
auricular
avicular (seta)
the posterior region of the body beliind ilie thorax and sometimes
followed by a caudal region or "tail"
without setae
a very stout projecting seta homologous with other setae but similar
in thickness to an internal aciculum (fig. 0.3. 6. v)
a stout internal chitinous rod which supports each of the two lobes
of a parapodium (fig. 0.3. i.c)
one or more elongated projections from the pygidium or terminal
segment on which the anus opens
a sensory projection arising from the anterior or dorsal surface of the
prostomium
a segment without a foot or parapodium
branching like a tree
a stout seta with a smooth shaft and a tuft of fine hairs at the end
(fig. i.iy.e)
ear-shaped
beaked (fig. 0.6. 7. v)
biarticulate
bidentale (seta)
bifurcate (seta)
bilabiate
bilimbate capillary
bipinnate
biramous
biramous parapodium
branchial crown
branchial vesicles
buccal
buccal tentacles or cirri
two-jointed
with two teeth (fig. o.a.S.v)
ending in two prongs (fig. o.5.4.d)
with two lips
a pointed seta with two wings or flattened margins like a feather
(fig. o.6.6.d)
a structure such as a feather with a main axis and two rows of side
branches (fig. o.4.4.d)
having two rami or forks
a foot or parapodium with two bundles of setae, one in the noto-
podium and one in the neuropodium (see also uniramous) (fig.
0.3. I.c)
a circle of filaments (radioles) for filter feeding and respiration
which arises from the head of a sabellid or serpulid worm
soft baggy papillae on the parapodia with a respiratory function
(fig. i.iy.i)
pertaining to the mouth
elongate or finger-shaped food gathering appendages either in or
around the mouth (fig. 0.6.6.B)
calicinate
canaliculate
capillary
capillary seta
caruncle
caudad
cephalic cage
cephalic rim
cephalic veil
like the calyx of a flower
with fine canals
hair-like
strictly a hair-like bristle but often used to cover all long slender
tapering setae (see types of notosetae shown in fig. 0.4)
a sensory lobe extending behind the prostomium (figs, o.a.a.b and
3-b)
towards the tail
long, forwardly directed setae which enclose and protect the head
(fig. 0.6.2. b)
a flange encircling the head (fig. o.j./.b)
a delicate hood-like membrane in the family Pectinariidae which
separates the opercular paleae from the buccal tentacles (fig.
0.6.4. b)
822
POLYCHAETA OF SOL'Tlll.RN Al RICA
cephalisation
ceratostyle
chromatophll gland
chromatophore
ceratophore
cirriform
cirrigerous (segment)
cirrophore
cirrostyle
cirrus
clavate
compound (seta)
compressed
copragogiie
cordate or cordiform
crenulate (seta)
crotchet
ctenidium(a)
cultriform
the modification and fusion of anterior segments to form a head
the distal joint of an antenna (fig. 1.4.C)
a densely staining gland in the ventral pinnule of Tomo/ilfris (fig.
0-3-3-c)
a special cell or group of cells carrying pigment
the basal joint of an antenna (fig. 1.4.C)
shaped like a cirrus
a segment bearing a cirrus
a basal projection on which a cirrus is moimted (fig. 0.2. i.c)
the distal part of a cirrus (fig. 0.2.1.C)
a sensory projection (usually tapered) derived from tlic superior part
of the notopodium (dorsal cirrus) or the inferior part of tlie neuro-
podium (ventral cirrus) (fig. 0.3.2.C)
club-shaped
a jointed seta
flattened in the lateral plane
a groove along the posterior region of a tubicolous worm along which
the faecal pellets pass before being voided from the tube
heart-shaped (fig. 0.3.8.C)
with a series of small cusps (fig. 0.4. 7. d)
a long-shafted seta with a hooked or curved end
a branchia or respiratory organ
shaped like a knife (see blade of seta, fig. 0.4. 3. d)
deciduous
dentate
dentate-crested hooks
denticle
denticulate
depressed
detritus
digitiform
distal
dorsum
liable to fall off like a leaf
toothed
setae with the apex of the shaft toothed (e.g. fig. 0.5. 7. v)
a structure like a minute tooth
with minute teeth or denticles
flattened in the dorso-ventral plane
particles of organic origin on the sea bed
finger-shaped
the part of a structure towards llie end
the dorsal surface of the body
echinulate
elytrigerous segment
elytron(a)
elytrophore
entire (margin)
eversible (proboscis)
e.xcision
facial tubercle
falciform
falciger
felt
filamentous
filiform
flanged fseta)
prickly like a sea-urchin
a segment bearing an elytron or scale
a dorsale scale-like structure found in the .'\phroditidae (fig. 1. 3. a.
and c)
a projection above a parapodium bearing an elytron
smooth edged, i.e. without papillae or other projections
capable of being extended by turning the inner part outwards (e.g.
fig. 0.3.8. b)
a part cut out
a projecting lobe on the upper lip below the prostomium (see pro-
jection below median antenna in fig. I.I.r)
hook-shaped
a compound seta having a stout, hooked blade or apex
matted hairs (setae) produced by the notopodia in some species of
Hermoninae (fig. i.i.n, o)
shaped like a filament or fine thread
thread-like, very slender
an elongate seta with a Matlcncd edge or margin (fig. o.5.i.d)
GLOSSARY OF TECHNICAL TERMS
823
foliaceous
frontal peak
fusiform
leaf-like
antero-lateral, often chitinised projections of the prostomium found
insomespeciesofPolynoinae (see also prostomial peaks) (fig. 1.2. a)
spindle-shaped or cigar-shaped
geniculate
genital papilla
glabrous
bent like a knee
projection below the neuropodium on which a reproductive duct
opens
smooth and glistening
harpoon seta
hastate
heterogomph (compound
seta)
hispid
homogomph (compound
seta)
hooded hook
hook
hyaline gland
a stout pointed seta with recurved barbs near the apex (fig. 1.1. g)
shaped like the blade of a spear
a compound seta with a slanting or asymmetrical joint between
shaft and blade (fig. 0.3. 2. d)
minutely furry
a compound seta with transverse or symmetrical joint between shaft
and blade (fig. I4.!.d, e, 1)
a stout, blunt or apically toothed seta with the apex protected by a
delicate chitinous envelope or guard (fig. 0.4. i.v)
a broad term used to cover a wide range of simple setae which have
stout shafts and blimt or toothed apices (e.g. figs. 0.4. i. v. 6.v, 7.V)
a gland which occurs in the pinnules of certain species of Tomopteris
and appears relatively transparent, sometimes with a yellow spot
in the centre (see notopodium of fig. 8.2.k)
imbricating
inferior
intermediate cirrus
intersegmental
introvert
overlapping like tiles
the more ventral of two structures
a cirriform projection between the notopodium and the neuro-
podium (fig. o.4.8.ca)
between segments
the anterior part of the body which can be withdrawn inwards
lamella
lanceolate
ligule
limbate (seta)
a flattened or plate-like structure
pointed and shaped like a lance
a compressed conical lobe of a parapodium (fig. 0.3.2.C)
a seta with a flattened margin to the blade (fig. o.5.i.d)
mammiliform
medial
median
membraneous
metastomium
raoniliform
multiarticulate
multidigitate
neotenic
nephridial papilla
neuropodium
neuroseta
notopodium
notoseta
shaped like a breast
on the side nearer the median or mid-line of the body
in the mid-line
thin and flattened like a membrane
the segmented body of an annelid worm, i.e. the whole length
between the prostomium and the pygidium
like a string of beads
with many joints (see antennae of fig. o.2.5.b)
with many finger-like lobes or projections
pertaining to a young or immature stage which reproduces sexually
a projection on which the excretory organ opens, usually posterior
or ventral to the parapodium
the lower or \'entral part of a parapodium
a seta arising from the neuropodium
the upper or dorsal part of a biramous parapodium
a seta arising from the notopodium
niiclial e]iauli'tli-
nuchal organ
occipital
occipital collar, fold or
lappet
occipital papilla
ocular
ocular peduncle
omnialophorc
ctperculuui
o\'oid
palea
palmate
palps or palpi
palpode
]3alpophorc and palpostyle
papillose
paragnaths
parapodial trimk
parapodiunifaj
]3ectinate
pcnicillate
peristomium (peristome)
pharynx
pilose
pinnate
pinnules
piriform
pluriarticulate
polygonal
postsetal
post-trochal
presctal
pretrochal
proboscis
POLVCHAF.TA OF SOI' rill'.RX AFRUIA
a raised and elongated sensory organ projecting jiostero-latcral to
the prostomium
a sensory organ on the prostomium or extending hack from it usually
in the form of a groove or ciliated ridge
pertaining to the posterior part of the prostomium
a proinincnt fold immediately overlapping the posterior part of the
prostomium (fig. i.3.g)
a papilla found on the posterior margin of the prostomium of some
species of PhvUodocc which represents a reduced median dorsal
antenna (fig. o.3.8.b)
pertaining to the eye
the projection supporting an eye (fig. i.iy.g)
see ocular peduncle
a lid or stopper which closes an opening ; part of the head of a
tubicolous polychaet which plugs the tube when the worm retracts
egg-shaped
a broad flattened type of seta (fig. 0.2. 3.b, c, d)
like the fronds of a palm
paired projections growing out from the sides of the head. In errant
polychaetes they arise from the ventral surface of the prostomiimi
and have a gustatory function but in sedentary polychaetes they
arise from the peristome and are usually grooved and adhesive
and pass food into the mouth
a tapering anterior projection of the prostomium (fig. o.3.j.b)
tlie basal and distal jjarts of a jiiinlcd palp (fig. o.3.2.b)
with papillae
horny or chitinous granules in the pharyngeal ( a\ity of the Nercidae
which help to grip the prey
the proximal, undi\ ided part i.f an elongate parapodium (fig. 0.3.3.1:1,
c)
segmental, foot-like projections bearing setae
with a series of projections arranged like the teeth oi a comb (fig.
o.2.8c)
like a little jDaint brush
the segment behind the prostomium, which is modified to form part
of the head and surrounds the mouth. Only the first segment
forms the true peristomium but in the families Xereidae, Hesioni-
dae and others the possession of more than two pairs of tentacular
cirri shows that two or more segments have fused to form the head
the posterior part of the mouth cavity leading on to the oesophagus
velvety ; covered with very short hairs
feather-like; with a main axis and lateral branches (fig. i.iS.s')
a series of side branches of some feather-shaped organ (fig. i.i8..s')
with a broad bulbous base and tapered tij) ; onion-shaped
many jointed (see multiarliculate)
many sided
posterior to the setae
posterior to the prototroch of the trochophorc larva
anterior to the setae
anterior to the prototro* h
the anterior part of the alimentary canal derived trom the slonia-
daeum which can be everted to projcc t forwards
GLOSSARY OF TECHNICAL TERMS
825
prostomial peaks
prostomium
protolroch
proximal
pseudo-penicillate setae
pygidial
pygidium
chitinised antero-lateral projections of the prostomium of certain
Polynoinae (fig. 1.2. a)
the anterior lobe in front of the mouth bearing eyes and antennae
the main girdle of cilia anterior to the mouth of trochophore larva
which is responsible for locomotion
part of a structure nearer to the base or main body; cf. distal
setae of the family Polyodontinae intermediate between true pen-
nicillate forms with a terminal tuft of fine spinules and forms with
hairy blades and tapering tip projecting beyond the tuft of spinules
(fig. 1.17.I)
pertaining to the anal segment or pygidium
the anal segment or terminal part of the body
radiole
ramose
ramus
receptaculum seminis
reniform
retort organ
rhomboidal
rosette gland
ringent seta
rostrum
rugose
one of the main radii or tentacles on the head of a fan worm (Sabel-
lidae or Serpulidae). The radioles normally bear two rows of side
branches or pinnules (fig. 0.6. 7. a)
branched
a branch or prong ; the notopodium and neuropodium which form
the two parts of a parapodium are often termed the two rami
a female organ to contain received spermatozoa
kidney-shaped
a clear club-shaped gland in the head of the Typhloscolecidae which
opens on the roof of the buccal cavity
irregularly four-sided
yellowish star-shaped glands in the pinnules of some species of
Tomopleris. They are situated next to the apices of the parapodial
rami (fig. S.i.c)
a seta with a series of annular serrations on both prongs (fig. 3.1.x)
the enlarged first tooth or main fang of a seta
rough or lumpy
sabre-like seta
scaphe
secondary tooth
segmental organ
sesquiramous
sessile
setal
setiger
setigeroiis lobe
simple seta
spathulate
spindle-shaped
spiniger or spinigerous (seta)
spinning gland
spinous pocket
spinule
AA
a seta with a broad curved blade
a flattened caudal appendage bearing the anus in Peclinaria
the second of two teeth, the first being the apical or terminal one
organs occurring in segmental arrangement. In the .\lciopidae they
are swellings (often pigmented) at the bases of the parapodia
a parapodium with a notopodium reduced to a dorsal cirrus, an
aciculum and sometimes one or two setae (see also sub-biramous)
without a stalk
pertaining to setae of bristles
a segment with setae
that projection or part of the notopodium or neuropodium which
bears the setae
an unjointed seta
like a spathula ; flattened, and blade-like with a blunt tip
elipsoid ; cigar-shaped with pointed ends
a compound seta whose blade tapers to a fine point (fig. 0.2.4.V)
a modified setigerous gland opening between the notopodium and
neuropodium of Polyodontinae which produces the chitinous threads
of which the tube is made
an enlarged serration on a seta, e.g. in Scalisetosus (Polynoinae) whose
edge is divided into spinules which surround a pocket-like cavity
(fig- i-7-j)
a small spine
826
rOLVt:llAETA OF SOU'lHERN AFRIOA
spinulose
spirally-serrulate (seta)
spur-gland
slriae
striated
stylet
stylode
sub-biramous
subeqiial
subspiral
subterminal
subtriangular
subuate
SLibuluncini
superior
provided with spinules
a seta whose blade is eiKirclcd uilh spirally arranged serrations ((ig.
i.i/.n)
a gland found in some species of Tomoptfris whose pointed end pro-
jects from the edge of the pinnule usually next to the chromophil
gland (fig. 8.I.C)
very fine parallel marks or lines
with striae
a small pointed, tooth-like structure
a botanical term here used to mean a fmger-likc projection usually
on a parapodium (fig. I.i8.r)
a parapodium which is neither completely uniramous nor biramous
approximately ecjual
approximately spiral
almost at the end
approximately triangular
an elongate, blunt and tapering projection
setae with a stout shaft suddenly tapering to a slender li|) irUer-
mediale between capillaries and uncini (fig. 23.2.e)
the more dorsal of two structures
lelotroch
tentacle
tentacular cirrus
tentacular formula
tcntaculophore
tessellated
thorax
tori of parajDodia
trochophore larva
the ciliated girdle around the anus of a trochophore larva
a slender outgrowth from the head
a cirrus arising from the peristome which is elongated to act as a
tactile organ (fig. o.3.8.b)
a series of letters and numbers used to indicate the arrangement of
the tentacular cirri and setae in the Phyllodocidae and .Mcio-
pidae
the basal projection on which a tentacle is mounted (fig. o.^.i-b)
a surface with a network of grooves like a tiled wall (fig. o.j.b.b)
the anterior region of the body
ridges from which the setae arise (see neuropodium of fig. 0.5.6.C)
the larval stage of an annelid or mollusc which develops from the
gastrula
with the end cut olT; not tapering
uncigerous
uiK inus
uniramous
urite
ventrum
winged capillary seta
bearing uncini
a general term used to cover sharp claw-like setae ; they may be
square or oval plates with several curved teeth or S-shaped with a
single tooth and a broad base (e.g. fig. o.6.3.d, 4.V, 5.V, 6.v, B.av)
with a single lobe or prong where two might be expected ; the oppo-
site of biramous. (Uniramous parapodia lack one fif the two
setigerous lobes)
a projection from the anal segment or pygidiiun
the ventral surface of the body
a simple, unjointed seta whose blade has an axial ril) but ilic margins
are flattened and tapering (fig. o.6.6.d)
REFERENCES 827
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■90.')- Report on the Polychacta collected by Professor Herdman at Cleylon in ic|o2. Siippl.
Rep. Ceylon Pearl Oyster Fish, 4 : 243-324.
WiLLi.vMs. T. 1851. Report on the British Annelida. Rep. Br. Ass. advrnl. Sci. 21 : 159-272.
Wilson, U. P. 1928. The post-larval development o( Loimia medusa Sav. J. mar. biol. Ass. U.K.
15: 129-144.
REFERENCES 841
1958. The polychaete Magelona alteni n.sp. and a reassessment o{ Magelona cincia Ehlers. J.
mar. biol. Ass. U.K.
WiREN, A. 1883. Chaelopoder fran Sibiriska Ishafvet och Berings Haf insanlade under Vega-
Expeditionen 1878 — 79. Vega-exped. vetensk. laktlag. 2: 283-428.
WoLLEBAEK, A. igi2. Nordcuropaeiske Annulater Polychaeta. I. Ammocharidae, Amphic-
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Kl.) 19U (18): 1-144.
Fmj
INDEX OF SCIENTIFIC NAMES
PARTS I & 2
[Part I. pp. 1-458; Part 2. pp. 459-841)
Family and sub-family names are primed in SM-VLL CAPITAL letters, new names are printed in
bold t\-pe. other valid names are printed in Roman type and names listed in synonymy are printed
in italic type. Only the principal reference is usually given to each name.
Abarenicola, 610
— aftinis africana, 61 1
— gilchristi, 61 1
aberans. Xotomastus, 599
aberrans, Loandaha. 220
aberrans. Lumbrineris. 439
abuwia, Paranereis, 301
abranchiata, Leaena. 721
abranchiata, Streblosoma, 723
abyssale, OKphanostomum, 703
acantholepis, Hermenia, 77
acantholepis. Polynoe, 77
acanthophora, X'ermiliopsis, 814
accraensis. Amaeana, 720
Acholoe, 52
— aslericola, 52
— orbiculata, 36
— squamosa, 52
acicula, Caulleriella, 507
acicularum, Marphysa, 378
Acrocirrus, 501
aculeata, Aphrodita, 33
acuminata, Ophelina, 579
acuta, Lumbrineris, 439
acutifrons. Ampharete, 699
acutifrom, Amphicteis^ 699
acutus, Than.'x. 504
adenensis. Marphysa, 399
adeonella. Spirorbis. 794
Aedicira, 563
- belgicae, 563
acnea, Alarphysa^ 378
aequiseta. Ariltrwe. 66
(iffjinsfla. HiT7fi/jthot\ bb
aequiseta aequiseta, Harmothoe. 66
— africana, Harmothoe, 68
afer, Autolytus, 283
afer, Cirralulus, 315
afer, Cirriformia, 515
afRnis africana. .\barenicola, 61 1
ajjinis^ Abarenicola as'^imilis^ 6r i
affinis, Clymeue^ 644
affinis, Flabelligcra, 655
qffinh luctator, Flabelligera. 652
— marenzdleri. Flabelligera, 652
ajjinis^ Pariospilus, 170
affmis, Praxillclla, 644
affinis, Samythella. 704
ojra. Dodecaceria, 498
afra, Eunice, 392
afra afra. Eunice, 392
— paupera, Eunice, 393
— punctata, Eunice. 393
africana. Aberenicola affinis, 61 1
africana, Eunereis, 293
africana, Glycera, 352
africana, Glvcera convolula, 352
africana, Harmothoe, 68
africana, Harmothoe aequiseta. 68
africana, Nicolea venustula, 73b
africana. Ophelia. 575
africana, Phyllodoce, 137
africana, Pisione, 133
africana, Sternaspis sciitata. 648
africanus. Cirratulus. 51 i
africanus, Leiochrides, 600
africanus. Thelepus sclosus. 722
agassizi. Rhamphobrathuini. 418
INDEX OF SCIENTIFIC NAMES
843
agilis, Ophiodromus, 226
agUis, Podarke, 226
Aglaophamus, 34 1
— dibranchis, 34 1
— macroura, 343
— malmgreni, 343
Aglaurafulgida, 426
Aglaurides, 426
— erythraeensis, 426
agnae, Polynoe, 38
agnesiae, Goniadopsis, 368
agnesiae, Nereis, 321
agulhana, Harmothoe, 74
agulhana, Hipponoa gaudichaudi, 122
agulhana, Nereis, 318
agulhana, Ophelia, 573
agulhasensis, Ampharete, 700
agulhasemis, Lysippe, 700
alata, Plotohelmis, 193
alba, Glycera, 360
alba, j\ereis, 360
albidenlata, Liimbrkonereis, 434
albidentata, Lumbrineris, 434
albifrons, Lumbriconereis, 435
albifrons, Lumbrineris, 427
albini, Synelmis, 215
albomaculatus, Corynocephalus, 1 77
albopicta, Eulalia, i^j
Alciopa, 179
— canlrainii, 1 76
— lepidota, 1 79
— petersii, 192
— reynaudii, 180
ALCiopiDAE, 172
Alciopina, 176
— parasitica, 177
alcocki, Sabellaria, 672
alcocki, Sabellaria spinulosa, 672
alcyonaria, Eurythoe, 1 28
Alcyope Candida, 1 88
Alentia, 44
— australis, 44
Alitta, 32 1
alleni, Halosydna, 88
Allmaniella, 42
— inluKa, 86
— marquesensis, 50
Alluandella, 279
— madagascariensis, 280
aha, Aphrodita, 35
alveolala, Sabella, 671
Amaea, 718
— Irilobata, 718
Amaeana, 718
— accraensis, 720
— trilobata, 718
Amage, 695
Amblyosyllis, 257
— formosa, 259
— lineolata, 233
— rhombeata, 257
ambrizettana, Nephtys, 349
ambylodonta, Nereis, 333
arnica, Syllis, 243
.^mmotrypane, 579
— aulogaster, 579
— longicaudala, 577
Ampharete, 699
— acutifrons, 699
— agulhasensis, 700
— capensis, 702
— griibei, 699
— kerguelensis, 700
AMPHARETIDAE, 686
AMPHARETINAE, 693
Amphicora medilerranea, 757
— sabella, 782
Amphicteis, 695
— acutifrons, 699
— gunneri, 695
Amphictene, 683
Aniphiduros, 223
Amphiglena, 757
— mediterranea, 757
Amphinome, 122
— incarunculala, 1 20
— pallasi, 123
— rostrata, 123
AMPHINOMIDAE, I20
AMPHITRICACEA, 731
Aniphitrite, 746
— bombyx, 767
— cincinnata, 727
— cirrata, 746
— cristata, 738
— gunneri, 695
— nesidensis, 744
— pauciseta, 746
— plumosa, 658
— reniformis, 764
— vesiculosa, 758
ampulUfera, Harmothoe, 47
ampuUi/era, Polynoe, 47
ampulliferus, Lepidonotus, 47
ampulliferus, Paralepidonotus, 47
Anaitides, 143
analisjjasmineira, 779
atuUis, Sabella, 774
844
POI.VCHAETA OF SOUTHERN Al RICA
Ancistrosyllis, 215
— constricta, 215
— groenlandica, 2 1 5
— parva, 21G
— robusta, 216
angflini, Callizorui, 172
angflini, Krohiia, i f|(i
angelini, Rhyiichonorclla, 190
angolacnsis, Mystides, 141
angolaiia, Dorvillea, 458
angolana, Stauronereis, 458
angrapequemis, Alicia, 543
angrapequensis, Orbinia, 543
Anguillosyllis. 27!
— capcnsis, 271
tinguiti/rons, Irma, 224
angustifrons, Ophiodromus, 224
angusti/rons, Podinke, 224
ankylochaeta, Opisthosyllis, 252
ankyloseta, Laeonereis, 303, 323
ankyloseta, TrypanosylUs, 255
annaiidalei, Euclymene, 641
annulosus, Tharyx, 506
aiiomala, OpheHa, 572
anomala, Protula tubularia, 820
anomala, Pseudonereis, 333
anops, SylUs, 243
Anotomastus, 594
antimtica, Leptoecia, 412
antarctica, Maldanella, 626
antarctica, Paronuphis, 412
antarctica, Vanadis, 181
antennata, Eunice, 384
antennata, Exogonoidcs, 279
antennata, Leodice, 384
antennata, Polydora, 473
Anthoitoma fragile, 546
— hexajihylhan. 533
antilopis, Harmothoe, 66
Antinoe, 57
— aeqiiiseta, 66
— epitoca, 57
— lactea, 58
— microps, 57
— ivaahli, 72
antipoda, Pectinaria, 685
Aonides, 478
— aiiricularis, 478
— gracilis, 56b
— oxycepliala, 478
Aphlebina hacmalodes, 717
Aphrodita, 33
— aculcata, 33
— alta, 35
— data, 79
— complanata, 1 28
— Jiava, 124
— minuta, 100
— rostrata, 123
Aphrodite cirrosa, 61
APHROniTIDAE, 30
APHRODITINAE, 30
aphroditois, Eunice, 389
aphroditois. Nereis, 389
APISTOURANCHIDAE, 52 1
Apistobranciuis, 521
Apomatus, 817
appendiculatus, Lopadorhynchus, 161
apsteini, Tomoptcris, 199
Arabella, 444
— iricolor cacrulca, 447
iricolor, 446
— niutans, 446
— novecrinita, 446
ARABELLIDAE, 442
ARAIiELLINAE, 442
Aracoda caerulea, 447
— capensis, 442
— multidenlala, 443
arborifcra, Dendronereis, 302
arenaceodonla. Nereis, 32 i
Arenicola, 608
— assimilis ajfmis, 61 1
— bombayensis, 610
— claparedii, 6 1 1
— loveni loveni, 610
sudaustraliensis, 610
— marina, 610
arenincola, Rhynchospio, 478
ARENICOLIDAE, 6o6
Arenicolides, 607
arenosus, Piromis, 664
arenosus, Stylarioides, 653
Arete capensis, 335
argm capensis, Dasyclwne, 751
— chefmae, Dasychone, 751
Aricia angrapequensis, 543
— hioreti, 545
— cwieri, 543
—foetida, 542
amtratis, 542
ligustica, 542
— laevigata, 539
Aricidea, 557
— belgicae, 563
— capensis, 562
— cur\'iseta, 557
INDEX OF SCIENTIFIC NAMES
845
— fauveli, 560
— fragilis, 557
— fragilis, 560
— jeffreysi, 558
— longobranchiata, 560
— pacifica, 563
suecica simplex, 558
suecica, 557
Armandia, 576
— cirrhosa, 576
— intermedia, 577
— leptocirrus, 577
— longicaudata, 577
— melanura, 576
armandi, Fabricia, 785
armata, Polydora, 466
armata, PuUiella, 595
armata, Syllidia, 227
armatus, Idanthyrsus, 675
armiger, Ltimbricus, 554
armiger, Scoloplos, 554
armillaris, Nereis, 249
armiUaris, Syllis, 249
Artacama, 733
— proboscidea, 733
ARTACAMACEA, 731
articulata, Psammolyce, 105
Asclerocheilus, 586
— capensis, 586
assimilis affinis, Abarenicola,(>ii
assimilis, Eunoe, 64
assimilis, Eurylhoe, 120
assimilis, Eusyllis, 266
astericola, Acholoe, 52
Asterope Candida, 188
Asychis, 645
— atlanticus, 646
— capensis, 647
— dorsofilis, 646
allanlica. Lumbriconereis, 440
adattlica, Lumbrineris heteropoda, 440
atlanticus, .\sychis, 646
atra, Lysidice, 378
atrocollaris, Cirratulus, 499
altemiala, Polynoe, 38
Auchenoplax, 695
Audouinia auslralis, 517
— filigera, 518
capensis, 498
meridionalis, 499
— punctata, 5 1 7
— saxalilis, 515
— semicincta, 517
— lentaculata, 515
Augeneria, 43°
— lentaculata, 430
aulogaster, Ammotrypane, 579
auranliaca, Proceraea, 283
aurantiacus, Autolytus, 283
aurantiacus, Polycirrus, 715
auricularis, Aonides, 478
australis, Alentia, 44
auslralis, Ariciafoetida, 542
auslralis, Audouinia, 5l^
auslralis, Cirratulus, 5iy
australis, Eunice, 385
australis, Heteronereis, 305
australis, Hololepida, 44
australis, Lciochrides, 600
australis, Notocirrus, 448
australis, Phylofoetida, 542
australis, Platynereis, 305
Australonereis, 297
Austrolaenilla, 42
AUTOLYTINAE, 237
Autolytus, 28 1
— afer, 283
— aurantiacus, 283
— bondei, 285
— charcoti, 283
— inermis, 233
— maclearnus, 284
— pictus, 284
— proliter, 284
tuberculatus, 281
.\xiothea catenata, 629
.\xiothella, 629
— jarli, 629
quadrimaculata, 629
babylonia, N'ermiliopsis, 813
bairdi, Dasychone, 751
bairdi, Grymaea, 723
bansei, Fabricia, 782
bansei, Oriopsis, 788
BarantoUa, 594
Barrukia, 42
belgtca, Nereis cylindraria, 680
belgicae, Aedicira, 563
belgicae, Aricidea, 563
belgicae, Paraonis, 563
bella, Sabellaria, 672
benguellana, Glycera, 358
benguellana, Glycera capilata, 35»
benguellana, Syllis. 249
benguellarum, Sphaerodor.dium, 28
benguellarum, Sphaerodorum, 289
benthaliana, Laetmonice, 33
846
POLVCHAETA OF SOUTHERN AFRICA
benthaliana. Laitmonue firoducta, 33
berrisfordl, Ciphiociromus. 224
Uhauaiiia, 1 17
— cn/)to{e/iluila. 1 18
— goodci. 1 18
— myrialcpis, 1 18
Ijiarliculala, Protodorvillea, 452
bicornis, Ophelia, 572
bidentata, Mystides, 143
bifilaris, Lumbrineris, 431
bifurcala, Hydroides, 808
bifurcata, Marphysa, 397
bijurctiltis, Eii/mmnliis, 808
bilincata, Eulalia, 154
hilineala, Hypnculalia, 154
hilineata, Phyllodocc, 154
biodtlatum, Bramhiomma, 760
bioculatum. Mcgalomma, 760
bioculatus, ClauUericlla, 509
bioadatusy Cirnitutiis, jOg
bioculatus, Hclerochrus, 509
bioreti. Ariciti, 54",
bioreti, Orbinia, 545
bipartitus, Dasybranchus, (103
biparlilus, Oncoscoltx, 603
bipes, Rhamphobrachuim, 418
Bispira, 756
— volutacomii, 7-,3
bispiralis, Prolula, 81H
bispirali^, Scrpiila, 818
blomstrandi, Eusyllis, 266
boa, ,Sii^alion, ukj
boa, Sthenelais. log
Boccardia, 462
— ligerica. 463
— polybranchia, 463
— pseudoiiatrix, 46-j
bocki, Prionospio. 490
bombayensis. Arenicola, Gio
bombyx, Aiiiphitritc, 767
bombyx, Spio, 474
bomb\'x, Spiopliancs, 474
bondei, Autolytus, 285
borealis, Mystides, 141
borealis, Spirorbis, 793
borealis, Spirorbis, 792
bou\'ieri, Syllis, 244
brachychaela, Syllis, 249
Brada, 656
— %-illosa, 652
rapensis, 656
brancliiatus, Oirrophorus, 563
branchiatus. Parasderocheilas, 585
Branchiocapitella, 594
Branr hiomaldane, 608
— \inccnti, 6u8
Branrhiomma, 767
Branchionuna, 757
— bioculatum, 7(10
— capcnsis, 770
— mushaensis, 758
— natalensis, 767
— nigromaculata, 770
loandensis, 73!
— quadrioculatmn, 7;,8
— serratibranchis, 768
— vesiculosum, 758
— violacea, 768
Brancliioasychis, 618
BramhoiColex craspidochaetus, 391
— oligobranchui, 591
— sphaerochat'tus, 591
Brania, 267
— furcelligera, 269
— pusilla, 267
— rhopalophora, 269
brevibranchia, Pista, 737
brevicirra, Lumbrineris, 433
brcvicirrus, Lumbricoucreis, 433
brevicirrus, A'otocirrus, 433
brcvis, Lopadorliynchiis, 162
brunnea, Lcpidasthenia, 92
brunnea. Nereis indica, 320
brunnea, Opisthosyllis, 233
caducus, Dasybranchus, 603
caducus, Dnsymallus, 603
caeca, Maiipasia, 164
caeca, Polydora, 469
caecum, Leucodorwu, 469
caerulea. Arabella irirolor, 447
caerulea, Aracoda, 447
caeruleus, Placoslegus, 801
caeruleus, Pomatoceros, 801
caeruleu\, Pomatoceros, 798
californicnsis, Mediomastus, 600
callaoana, .Kfrei\. 292
Callizona angriini, 172
— mbbii, 1 89
— moebii, 1 89
— jiasula, 189
— setosa, 192
CalUzonella lepidola, 179
calodonta, Platynereis, 306
candela, Jasmineira, 779
Candida, Alcyope, 1B8
Candida, Asterope, 188
Candida, Chlotui, 120
IXDEX OF SCIENTIFIC NAMES
847
Candida, Torrea, 188
CASEPHOKIDEA, 710
cantabra, Nerindcs, 485
cantrainii, Alciopa, 1 76
cantrainii, Naiades, 176
capense, Rhamphobrachium, 418
capense, Sigalion, 103
capense, Sphaerodidium, 289
capense, SphaerodoTum, 289
capensis, Ampharete, 702
capensis, Anguillosyllis, 271
capensis, Aracoda, 442
capensis. Arete, 335
capensis, Aricidea, 562
capensis, Asclerocheilus, 586
capensis, Asychis, 647
capensis, Audouinia jUigera, 498
capensis, Brada villosa, 656
capensis, Branchiomma, 770
capensis, CauUeriella, 507
capensis, Chaeloplerits, 522
capensis, Cirraliihis, 5 1 7
capensis, Cirriformia, 51 7
capensis, Clymene praetermissa, 613
capensis, Dasychone, 770
capensis, Dasychone argus, 751
capensis, Dasychone violacea, 751
capensis, Diopalra neapolitana, 413
capensis, Diplocirrus, 666
capensis, Dodecaceria, 504
capensis, Drilognathus, 444
capensis, Eriphyle, 377
capensis, Euchone, 776
capensis, Eulalia, 154
capensis, Eulalia viridis, 154
capensis, Eunice, 400
capensis, Eunoe, 38
capensis, Euphrosine, 127
capensis, Eurythoe, 120
capensis, Fabricia, 784
capensis, Ficopomatus, 810
capensis, Glycera convoluta, 359
capensis, Glycinde, 371
capensis, Gunnarca, 673
capensis, G)ptis, 231
capensis, Hermella, 673
capensis, Heierocirrus, 498
capensis, Heierocirrus capuiesocis, 507
capensis, Hypsicomus, 761
capensis, Lanassa, 722
capensis, Lipobranchius, 588
capensis, Loandalia, 220
capensis, Lumbrineris, 432
capensis, Lumbrineris, 427
capensis, Lysidice, 402
capensis, Lysippe, 702
capensis, Maclovia iricolor, 443
capensis, Alagalia, 227
capensis, Magelona, 497
capensis, Maldanella, 627
capensis, Marphysa, 400
capensis, Mediomastus, 600
capensis, Melinnopsides, 693
capensis, Alelinnopsis, 693
capensis, Mesochaetopterus, 532
capensis, Micronereides, 298
capensis, Xeovermilia, 816
capensis, Nephtys, 344
capensis, Nereis, 320
capensis, J\fereis cylindraria, 683
capensis, Nerine cirratulus, 460
capensis, .Nicomache, 6 1 4
capensis, JVicomache tumbricalis, 6 1 4
capensis, Notocirrus, 443
capensis, Ophelia, 573
capensis, Oridia, 784
capensis, Oxydromus, 231
capensis, Pallasia, 667
capensis, Panthalis oerstedi, 97
capensis, Parahalosydna, 86
capensis, Paraonides lyra, 568
capensis, Paraonis lyra, 568
capensis, Parasclerocheilus, 585
capensis, Parmensis, 66
capensis, Pectinaria, 683
capensis, Perinereis, 335
capensis, Phyllodoce, 148
capensis, Pliylo, 540
capensis, Pisla cristala, 732
capensis, Polydora, 466
capensis, Polymnia, 732
capensis, Polynoe, 86
capensis, Praxilla, 644
capensis, Praxillella, 644
capensis, Praxillella praetermissa, 6 1 3
capensis, Prionospio, 459
capensis, Protomystides, 143
capensis, Prolula, 816
capensis, Protula tubularia. 8 1 6
capensis, Ranzania, 532
capensis, Ranzanides, 532
capensis, Sabella, 678
capensis, Sabellaria, 673
capensis, Sabellides, 698
capensis, Scolaricia, 549
capensis, Scoloplella, 536
capensis, Spermosyllis, 270
capensis, Sphaerosyllis, 276
K.tfi
POLVCHAEIA OI' SOrilllRX Al RICA
capensis, Sphaerosyllis hvilrix, 276
capcnsis, Spirorbis, 796
ccipensis^ Stylntioiiks, 664
capensis, Syllidia, 227
caperuis, Syllis, 235
capensis, Telotholepus, 7;; i
capensis, Tra\'isiopsis, 207
capemis, Trophonia, 664
capitala, Capitclla, 595
capitata, Glycera, 355
capitala henguellana, Glycera. 358
capitata, Plotohelmis, 195
capitala. Rhynchonerella, 195
capitatiis. Lwnbricus, 595
Capitclla, 594
— capitata, 593
— Jili/ormis, 601
CAPITELLIDAE, 59 I
Capitobranchus, 594
Clapitoniastus, jr)4
caputesocii capensis, Heterrocirrus, -,07
capiilleonis, Polynoc. 38
carini/crin firiinii, Placoitegus, 800
rarinifcriis, Pomatoceros, 801
carinulala. Pulynoe. 81
I arinulatus, Lcpidonotiis, 81
(Uirobia castanea. 149
carpcnteri, Toniopteris, 204
Castalia claparedii. 230
castanea, Carobia, 149
castanea. Genetvllis. 149
castanea, Phyllodoce, 149
catenata, /Vxiothea, 629
caudata, Jasmineira, 780
caudata. Nereis, 321
Caulleriella, 506
— aciciila, 507
— bioculatus, 509
— capensis, 507
— zetlandica, 509
cavalli, Tomopteris, 206
caialli. Tomopteris. 204
cavifrons. Lwnbriconereis. 438
cavifrons, Lunibrineris, 438
cavifrons, Lunibrineris. 427
celox, GrccJJia, 180
Ccnothrix nnitan s , 446
ceramensis, Scalisetosus, 58
Ceratoccphale, 297
Ceratonereis, 324
— costae, 325
— ehlersiana, 291
— erythraeensis, 327
— falcaria, 309
— hircinicola. 327
— keiskania, 327
— mirabilis, 324
— pachycliaeta, 325
Chaetacanthus, 43
chaf;topteridae, 522
Chaetopterus, 529
— capensis, 522
— hamaliis, 522
— varieopedalus, 529
Chaetozone, 509
— setosa, 510
— zetlandica, 509
charcoti, Aulolytus, 283
chejinae, Daychone nrgns, 751
chilensis, Eiirythoe, 130
chtlen\is. Euthelepus, 725
chilensis, Ninoe, 430
ciiiiensis, Notocirrus, 448
chilensis. Paraeurythoe, 130
chincnsis, Lcf>crates, 230
chilensis, Slreblosoma, 725
Chloeia, 123
— Candida, 120
— flava, 124
— fusca, 124
— gilchrisli, 120
— inermis, 123
— natalensis, 120
Chhraenta telragona, 652
Chone, 776
— collaris, 777
— filicaiidata, 776
— infimdibuliforniis, 776
— letterstedti, 777
chrysncoina. Hermione, 35
chrysocoma, Pontogenia, 33
chrysodernia, Cirratulus, 51 1
chrysodon. .Sabella, 678
chrysolepis, Palcanoius, ii(3
CIIRYSOPETALIDAE, 115
Chryso/ietalnni. 1 1(3
— debile, i 1 7
— ehtersi. 1 1 7
chiini, Rhaniphobrachium, 420
cidariophiUini, Phalacrostcinma, 669
ciliala, A^creis, 340
ciliata, Polydora, 469
cilialus, Leucodore, 469
ciricinnala, Amphitrile, 727
cinrinnalus, Thelepiis, 727
cincta, Eunice, 382
cincta, Magelona, 495
cincta, .Vicidion, 382
INDEX OF SCIENTIFIC NAMES
849
cingulala, Dasychone, 751
cirrata, Amphitrite, 746
cirrala, Clymene, 623
cirrata, Glycera, 359
cirrata, Kefersteinia, 228
cirrata, Laonice, 480
cirrala, Nerine, 480
cirrata, Pectinaria koreni, 682
cirrala, Psamathe, 228
CIRRATULIDAE, 498
CIRR,\TULINAE, 5OI
Cirratulus, 510
— "fir, 5^5
— africanus, 51 1
— alrocoliaris, 499
— auslralis, 5 1 7
— bioculatus, 509
— capensis, 5 1 7
— chrysoderma, 5 1 1
— cirratus, 513
— concinnus, 514
— dorsobranchialis, 506
— filiformis, 5 1 1
— gilchristi, 513
— punclalus, 5 1 7
— tentaculus, 499
mtridionalis, 499
— sp, 498
cirraliilus, Lumbricus, 483
cirralulus, Nerine, 483
cirraliilus capemis, Nerine, 460
cirratus, Cirratulus, 513
cirratus, Lumbricus, 5 1 3
cirrhosa, Armandia, 576
cirrifera, Prionospio, 486
Cirriformia, 514
— capensis, 517
— filigera, 518
— maryae, 514
— punctata, 517
• — saxatilis, 515
— tentaculata, 515
ctrrobranchiata, Eunice, '^■j-j
cirrobranchiata, Prionospio, 488
Cirrophorus, 562
— branchiatus, 563
cirropunctata, Syllis, 250
cirrosa. Aphrodite, 6 1
cirtosa, Larymna, 425
Cirrosytlis picla, 233
— luberculata, 28 1
claparedii, Arenicola, 61 1
claparedii, Caslalia, 230
claparedii, Leocrates, 230
claparedii, Nicolea, 732
claparedii, Sphaerodorum, 289
clava, Aphrodita, 79
clava, Lepidonotus, 80
clava, Lepidonotus, 37
clava, Lepidonotus semitecta, 85
clavator, Exogone, 272
clavigera, Gastrolepidia, 51
closterobranchia, Syllis, 234
Clymaldane, 617
Clymene ajfinis, 644
— cirrata, 623
— glandularis, 639
— lombricoides, 636
— luderitziana, 64 1
— lyrocephala, 638
— microcephala, 6 1 4
— monilis, 633
— mossambica, 639
— natalensis, 638
— oerstedii, 635
— praetermissa, 642
capensis, 6 1 3
— producta, 633
— saldanha, 6 1 4, 634
— ^/i, 6 1 3
Clymenella, 618
clymenoides, Johnstonia, 631
Ciymenura, 623
— tenuis, 623
coasta, Cossura, 581
coccinea, Eunice, 389
coccinea, Lumbriconereis, 436
coccinea, Lumbrineris, 436
coccinea, Nereis, 436
collaris, Chone, 777
collaris, Lysidice, 402
colonia, Polydora, 468
Colymmatops granulatus, 731
comans, LamellisyUis, 257
comata, Terebella, 729
comatus, Thelepus, 729
communis, Scalisetosus, 38
compacta, Pionosyllis, 261
complanala, Aphrodita, 128
complanata, Eurythoe, 128
complexa, Serpula, 8 1 7
concharum, Dodecaceria, 502
conchilega, Lanice, 743
conchilega, Nereis, 743
conchylega, Nothria, 425
conchylega, Onuphis, 425
concinna, Dindymene, 576
850
i'Oi,vc:H.\i:r.\ OF sol ihkr.x ai rica
concinna, Travisia, 576
concinnus, C.irratulus, 514
congoensc, Pycnodcrma, (156
congoensis, Goniada, 365
coniceps, Plotobia, 210
coniceps, Travisiopsis, 210
constricta, Ancistrosyllis, 2 if)
convoluta, Glycera, 360
— afrkana, Glycera, 352
— capeiuis. Glycera, 359
corallina, Marphysa, 400
coraliuia, Xauphanta, 400
cornuta, Polydora, 465
comiita, Sagitella, 212
cornuta, Syllis, 244
corollifera, Daiychone, 751
conalophila, Harmothoe. 72
Corynocephalus albomaciilaliis, 177
— lemiii, 193
Cossura, 581
— coasta, 581
— longocirrata, 581
cossuRiDAE, 581
costae, Ceratonereis, 325
coilae, jWm'j, 323
costarum, Spiochaclopterus, 328
costarum, Telepsavm, 528
coutierei. Nereis, 31 1
cra^pidochaetus, Branchoscolex, 591
crassa, Enmenia, 588
crassa, Pulyphysia. 588
crasucornis, Syllis, 234
crinita, Notopygos, 130
criilata, Amphitrite, 738
cristata, Melinna, 691
cristata, Pista, 738
cristata capemis, Pista, 732
cristata, Polynoe, 82
cristata, Sabellides, 691
cristatus, Lepidonotus, 82
crocea, Phyllocomus, 702
crosctemis, Harmothoe, 36
croislandi, Pomaloleios, 800
cryplocephala, Bhatrania, 1 18
Cryptopomatiis, 675
— geayi, 667
crystallina 1 rystalliiia, Vanadis. iBa
— inornata, \'anadis, iBa
CTtNODRILINAE, 50I
ctenostoma, Odonlosyllis, 261
clenostoma, Odonlosylli^, 260
ciiltrtjira. .Nereis, 337
cultrifcra, Perinereis, 337
cultrifcra hcUeri, Perinereis, 292
culveri. Nereis, 303
cuprea. Nereis, 417
cuproa cuprea, Diopatra, 417
— punctifcra, Diopatra. 418
curtum, Pcdinosoma, 158
curviseta, Aricidea, 557
cuvieri, Aricia, 543
cuvieri, Orbinia, 543
cygnochaetus, Proscoloplos, 538
cvlindraria bclgica, Nereis, 680
— capensis. Nereis, 683
cylindricauda, Lumbriclymene, 625
cvlindrijer, Scolnplos, 546
Dasybranclius, 603
-- bipartitiis, 603
— caducus, 603
Dasychone, 767
— argus capensis, 751
chefmae, 751
— bairdi, 751
— capensis, 770
— cingulata, 751
— corollifera, 751
— loandensis, 751
— natalensis, 767
— nigromaculata, 770
— odhneri, 771
— orientalis, 751
— serratibranchis, 768
— violacea, 768
capends, 751
Dasvmallus caducus, 603
debile, Chrysnpelalnm, 1 1 7
dcbilis, Lumbrincris, 432
debili^, Lumbrinens, 427
debilis, Palcanotus, 1 17
debilis. Palmyra, i [ 7
decipiens, Leonnates, 330
dendrolepis, Slhenelais, 98
dendrolepis, 'I'halenessa, 107
Dendroncreides, 302
— heteropoda, 302
— zululandica, 302
Dendronereis, 301
— arborifera, 302
denlala, Diopatra, 405
depressa, Eunice. 378, 395
depressa, Marphysa, 395
Desdemona, 788
— ornata, 790
Dexiospira, 797
Dindymene concinna, ',76
dibranchis, Aglaophamus, 34 1
IIVDEX OF SCIENTIFIC NAMES
851
dibranchis, Nephtys, 341
dictyophora, Harmothoe, 65
dictyophorus, Polynoe, 65
difficilis, Lumbrineris hetcropoda, 440
Diopatra, 412
— cuprea cuprea, 417
punctifera, 418
— dentata, 405
— dubia, 415
— holobranchiala, 405
— monroi, 415
— musseraensis, 4 1 7
— neapolitana capensis, 413
neapolitana, 413
— punti/era, 418
— viridis, 405
Diplocirrus, 664
— capensis, 666
diponia, Eupotnalus, 809
dipoma, Hydroides, 809
Disoma, 519
DISOMIDAE, 519
Dispio, 481
— magna, 482
— uncinata, 482
Ditrupa, 799
diversicolor, Nereis, 317
diversidenlata, Peririereis, 293
diversosetosuni, Rhamphobrachium, 418
djiboutiensis, Scyphoproctus, 604
djibotitiensis, Sjllh, 235
Dodecaceria, 501
— afra, 498
— capensis, 504
— concharum, 502
— fistulicola, 499
— laddi, 502
— pulchra, 502
dorsipapillata, Pholoe, 100
dorsobranchialis, Cirratulus, 506
dorsobranchialis, Tharyx, 506
dorsofilis, Asychis, 646
Dorvillea, 454
— angolana, 458
— gardineri, 455
— neglecta, 455
— rubrovittata, 457
— rudolphi, 457
DORVILLEI.NAE, 45O
Dricschia, 75
— pelagica, 75
Drilognalhus, \\\
— capensis, 444
Drilonereis, 447
— falcata, 447
— monroi, 448
dubejiiy Lumbrineris^ 427
dubia, Diopatra, 415
dubia, Orbinia, 547
dubia, Prionospio rnalmgreni, 489
dubia, Scolaricia, 547
dubia, Travisiopsis, 212
duccii, Tomopteris, 201
dumerilii, Mereis, 306
dumerilii, Platynereis, 306
dunckeri, Tomopteris, 201
durbanensis, Lcpidonotus, 85
durbanemis, Alarphysa, 378
Dysponetus, 1 16
dysleri, Protula, 8 1 7
dysteri, Salrnacina, 8 1 7
dysleri incrustans, Salrnacina, 816
Dystilia violacea, 75 1
echinata, Serpula, 810
echinata, Serpula vermicularis, 810
Echinorhynchus scutalus, 648
edwardsi, Glycera, 359
edwardsi, Sigalion, 107
egena, Protodorvillea, 454
egena, Stauronereis, 454
ehlersi, Chrysopetalwn, 1 1 7
ehlersi, Euleanira, loi
ehlersi, Oriopsis, 787
ehlersi, Potamilla, 752
ehlersi, Prionospio, 490
Ehlersia, 243, 244
— femigina, 244
ehlersiaeformis, Pionosyllis, 263
eklersiaeformis, Pionosyllis, 264
ehlersiana, Ceratonereis, 291
ehrenbergi, Terebella, 748
eimeri, Oria, 787
eimeri, Oriopsis, 787
elegans, Jasmineira, 780
elegans, Lcpidasthenia, 90
elegans, Phalacrostemma, 669
elegans, Polynoe, 90
elegans, Pygospio, 475
elegans, Tomopteris, 205
elioti, Phyllochaetopterus, 525
elisabethae, Euphione, 77
elongata, Samythella, 704
elongalus, jXeosabellides, 703
emerita, Goniada, 365
Enapteris, 197
• — euchaeta, 202
enigmatica, Mercicrella, 812
l'()l.VC:H.\K r.\ Ol- SOUTHERN' ATRKIA
Ktiipio rhomb igfrti, 54
Eone gracilis, $60
Ephesia gracilis, 288
Epidiopatra, 408
— gilchristi, 409
— hujiferiana, 409
hupfcriana, 40()
monroi, 41 1
— papillosa, 408
epitO( a, .\ntinoe. 57
eremita, Onuphis, 422
Ereutho plumosa, 718
erinaceus, Sphaerosylli*;, 277
Eri/ihyle cajiensis, 377
ervtliraccrtsi^, A^lnurides, 426
erytliraccnsis, Ceratoiiereis, 327
erxthrotacnia. Hcmilepidia, 55
eryllirotacnia, Polynoe, 55
Eteone, 139
— foliosa, 141
— ornata, 140
— siphonodonta, 140
— spclsbcrgcnsis, 136
— s\phoiiodonta, 140
— sp, 141
Etconidcs, 139
Eucanmculata grubei, 120
euchaeta, Eiiapleriu 202
euchaeta, Tomopteris, 202
Euchone, 774
— rapensis, 776
— rosea, 774
Euclymcnc. 634
— annaiidalei, 641
— glandularis, 639
— lombricoides, 636
— kideritziana, 641
— lyrocephala, 638
— mossambica, 639
— natalensis, 638
— oerstedi, 635. 636
— quadrilobata, 638
— sp, 6 1 3
EL'CLVMF-NINAE, 6l 7
eugeniae. Nereis, 312
eugcniae, .\icon, 312
Eulalia, i")i
— albopictn, 137
— bilineata, 154
— capensis, 154
— falsa, 155
— macroceros, 1 52
— microcephala, 1 ",2
— sanguinea, i ',"",
— trilineata, 152
— viridis, 1 36
capensis, i 54
Eulcanira, loi
— ehlersi, loi
Eiilepis geayi, 43
— hamifera, 45
Eumenia, 586
— crassa, 588
Eumida, 155
— sanguinea, 155
Eunereis. 323
— ofricana, 293
— hardyi, 323
Eunice, 380
— fl/ra, 392
afra, 392
paupera, 393
punctata, 393
— antennata, 384
— aphroditois, 389
— australis, 385
— capensis, 400
— cincta, 382
— cirrobranchiala, 377
— coccinea, 389
— depressa, 378, 395
— filamentosa, 392
— filamenlosa, 377
— floridana, 388
— grubei, 391
— indica, 386
— kinbergi, 377
— macrobraiuhui, 377
— ntossambica, 395
— jnjirrayi, 385
— norvegica, 388
— paupera. 393
— pennata, 388
— pmiclala, 393
— rousseaui, 389
— savignyi, 378
— schemacephala, 384
— siciliensis, 382
— tentaculata, 391
— torquata, 389
— tubifex, 386
— vittata, 385
EfNiciDAE, 374
EfNICINAE, 377
Eunoe, 61
— assimilis, 64
— capensis, 38
— luibrcchti, 62
INDEX OF SCIENTIFIC NAMES
853
— macrophthalma, 64
— nodulosa, 62
Eupanthalis, 94
— kinbergi, 94
— tubijexy 93
Euphione, 76
— elisabcthae, 77
— lenuiselosa, 82
Euphrosine, 126
— capensis, 127
— myrtosa, 127
— pohbranchiay 120
Eupolymnia, 744
— nebulosa, 744
etz/wmaloides, Sabellaria, 667
eupomatoides, Sabellaria spinulosa, 667
Eufiomatus, 804, 807
— bifuTcaliis, 808
— dipoma, 809
— lunulifera, 807
— spinosiis, 809
— uncinatus, 805
Eurato sarictijosephi, 771
Eurysyllis, 238
Eurythoe, 127
— alcyonaria, 1 28
— assimilis, 120
— capensis, 120
— chilensis, 130
— complanata, 128
— matthaii, 128
— parvecarunculata, 1 28
El'SYLLINAE, 237
Eusyllis, 264
— assimilis, 266
— blomstrandi, 266
— monicornis, 233
— sctabulemis, 235
Euthalemssa insignis, 107
Euthelepus, 726
— chilerisis, 725
— kinseniboensis, 726
— setabulensis, 726
Eiarne hubrechti, 62
exilis, Syllis, 250, 253
eximia, Goniada, 372
eximia, Ophioglycera, 372
Exogone piisilla, 267
Exogone, 271
— clavator, 272
— gemmifera, 274
— hfterochaeta, 233
— heterosetosa, 274
— naidina, 271
— normalis, 275
— verugera, 272
EXOGONINAE, 237
Exogonoides, 279
— antennata, 279
exlemmla, Lagisca, 36
Fabricia, 782
— armandii, 785
— bansei, 782
— capensis, 784
— filamentosa, 784
— mossambica, 78 1
FABRICIIN.^E, 755
Fabriciola, 781
— mossambica, 781
JalcaTia, Ceralonereis, 309
falcaria. Nereis, 309
falcata, Drilonereis, 447
falcata, Pilargis, 218
falcigera, Neovermilia, 816
falsa, Eulalia, 155
falsa, Nereis, 317
falsovariegata, Perinereis, 335
fasciata, Pista, 742
fasciala, Terebella, 742
fauveli, Aricidea, 560
Jatweli, Lfinice, 732
fauveli, Notomastus, 597
FaitvelieUa pukhra, 132
felix, Phylo, 540
ferrugina, Ehlersia, 244
ferrugina, Syllis, 244
fenuginea, Syllis, 244
fibrillata, Maldanella, 627
Ficopomatus, 810
— capensis, 810
— macrodon. 810
filamentosa, Eunice, 392
jUamentosa, Eunice, 377
filamentosa, Fabricia, 784
filibranchia. Tharyx, 505
filicaudata, Chone, 776
filicornis, Laetmonice, 32
filicornis, Laetmonice, 30
filicornis, J\'ereis, 48 1
filicornis, Spio, 481
fili/ormis, Capitella, 601
filiformis, Cirralulus, 51 1
filiformis, Heteromastus, 601
filigera, Audoninia, 518
filigera, Cirriformia, 518
filigera capensis, Audoninia, 498
— meridionalis, Audoninia, 499
854
POI.VCHAL lA OF SOUTHKRX AFRICA
filigertis, Lumbricus. 518
Kilograna, 817
— iniplexa. 81 7
Jitograna, Scrpula. 817
FILOGRANINAE, 8 1 1)
Jdiim. Lumhriconereis. 447
/islulicolii. Doilfcaceria, 499
Flabclligera. 655
— affinis. 655
lucla/or. 652
marenzelleri, 652
— luctator, 652
— xanlhotrichn, G52
FLARELLIGERIDAE, 652
/flirt, .i/i/irorlila. 124
flava. Chloeia, 124
flaia, .Vereis, 139
Hava, Polydora, 468
flava, Psammolyce, 105
flaroiiridu, Helerocirrus, 498
flexuosa, Js^ereis, 223
floridana, Eunice, 388
floridana, Lumbrineris, 427
fot'tida, Aricia, 542
fnelidti auslralis, Aricia, 542
foetida auslralis, Phylo, 542
— foetida, Phylo. 542
foetida lignsticn, Orbinia, -,33
foetida ligustica, Phylo, 542
— ligNslica, Phvlo, 542
Joliifera, Sabella, 75 1
foliigera, Pisia, 740
foliosa, Eteone, 141
folioia, Pliylloduce, i 5 i
foHosum, Notopliyllum, j-,!
foraminosus, Spirorbis, 797
forbesii, Travisia, 575
formosa, .'VmblyosylHs, 259
fortnosa, Pteroiyllis, 259
formosa, X'anadis, i8r
fragile. Anthostoma, 546
fragile, Hcrmadion, 59
I'ragilis, .\ricidea, 557
fragilis. Aricidea, 560
fragihs, Haploscoloplos, 546
fragilis, Lumbricus, 431
fragihs, Scahsetosus, 59
fraserthomsoni, Harmothoe, 74
fristedti, Phyllodoce, 147
Juciclua, Sabellaria, 672
fucicola, Sabellaria spinulosa, 672
fuigens, Paraonis. =165
fi/lgem, Rhvnchmerclta, 19-,
fulgida. Aiiliiiii,!. 426
fulgida, Ocnone, 42b
fulgoris, Poecilochaetus, 520
fulgurans, Syllis, 260
fuliginosa, Malacoceros, 477
furcelligera, Brania, 269
furcelligera (Jrubea, 269
fusca, Chloeia, 124
fusca, Sabella, 764
fuscapunctata, Vanadis, 184
fiisicornis, .Syllis, 235
fiisifera, \erois, 323
fusiformis, Ovvenia, 649
gallafiagens^, Puudonereis, 331
gardineri, Dorvilea, 4-15
gardineri, Spirobranchus, 803
gardineri, Stanrrophaliii. 453
Gastrolepidia, 51
— clavigera, 51
Gattyana, 61
— mossambica, 6r
gaudichaudi, Hipponoa, 122
gaudichaudi agulhana, Hipponoa, i:
— gaudichaudi, Hipponoa, 122
geqvi, Cryptopomatus, (567
geayi, Eulepis, 45
geayi, Pareulepis, 45
gelatino^a, Polvnoe, 44
gemmifera, Exogone, 274
gemmipara, Trypanosyllis, 256
gemmulifera, Trypanosyllis, 2"-,6
genelta, Hesione, 221
Genctyllis, 149
— ca^tanea, 149
geophiliformi^, .Nothria, 424
geophiliformis, Onuphis, 424
giardi, Polydora, 471
gibba, Odontosyllis, 261
gigantea, Glycera. 362
gigantea, Ophioglycera, 372
gigantea, Serpula, 803
gigaiitem, jVotomaitm. 597
giganteus, Spirobranchus, 803
gilchristi, Aharenicola, 61 1
gilchriiti, Chloeia, 120
gilchristi, Cirratulus, 513
gilchristi, Epidiopatra, 409
gilchristi, Harmothoe, (38
gilchriui, Lygdamii indicia, 667
gilchristi, Lygdamis murata, (177
gilchristi. Nereis, 315
gil< hristi, Xerindes, 485
gllfhri\li, Sabellaria ipinulom, 667
gilchusti. Teliere^ murata, 677
INDEX OF SCIENTIFIC NAMES
855
girardi, Malacoceras, 477
glacialis, Trichobranchos, 7"
glandigerus, N'ermiliopsis, 813
glanduiaris, Clymene, 639
glandularis, Euclymene, 639
glauca, Trophonia, 666
glandulosa, Nereis, 321
glauca, Polynoe, 84
glaucus, Lepidonolus, 84
glebifex, Maldane, 645
glutaca, Rhynchospio, 478
glutaea, Scolecolepis, 478
Glycera, 355
— ajricana, 352
— alba, 360
— benguellana, 358
— capitata, 355
benguellana, 358
— cirrata, 359
— convoluta, 360
africana, 352
capensis, 359
— edwardsi, 359
— gigantea, 362
— goesi, 352
— krauisi, 352
— lancadivae, 359
— longipinnis, 356
— natalensis, 356
— papulosa, 358
— parashadi, 359
— prashadi, 359
— rouxi, 362
— subaenea, 363
_ tesselata, 359
_ tridaclyla, 352
— unicornis, 362
GLYCERIDAE, 332
Glycinde, 371
— capensis, 371
— kameruniana, 371
_ maskallemis, 369
— multidens, 371
Glyphanostomum, 703
— abyssale, 703
goesi, Glycera, 352
Goniada, 363
— congoensis, 365
— emerita, 365
— eximia, 372
— maculata, 367
Goniadella, 367
— gracilis, 368
COSIADWAE, 352
Goniadopsis, 368
— agnesiae, 368
— incerta, 369
— maskallensis, 369
goodei, Bhawania, 118
goreensis, Harmothoe, 69
Gorekia, 42
gracile, Sphaerodorum, 288
gracilior, Rhodine, 619
gracilis, Aonides, 566
gracilis, Eone, 368
gracilis, Ephesia, 288
gracilis, Goniadella, 368
gracilis, Haliplanes, 164
gracilis, Maupasia, 164
gracilis, Onuphis, 412
gracilis gracilis, Paraonis, jbb
oculata, Paraonis, 565
gracilis, Phyllochaetopterus, 524
gracilis, Rhynchonerella, 189
gracilis, Syllls, 241
granulata, Nereis, 309
granulatus, Colymmatops, 731
graveleyi, Odontosyllis, 260
Gravierella, 630
— multiannulata, 630
— s/)-, 613
Greeffiacelox, 180
— oahuensis, 1 72
groenlandica, Ancistrosyllis, 215
Grubea, 267
— JurcelUgera, 269
— pusilla, idl
— rhopalophora, 269
grubei, Ampharele, 699
grubei, Eucarunculata, 120
grubei, Eunice, 391
grubei, Pista, 732
Grymaea bairdi, 723
— persica, 725
guinensis, Sabellaria, 672
Gunnarea, 673
— capensis, 673
gunneri. Amphicteis. 695
gumeri, Amphitrik, 693
Gyptis, 230
— capensis, 231
— propinqua, 231
— sp., 232
haematodes, Aphlebina, 7 ' 7
haemasoma. Marphysa, 378
haemasoma. Marphysa sangmnca, 378
haematodes. Polycirrus, 7 1 7
8s6
POLVCHAKIA OF SOUIHKRN AIRKIA
Haliplanes gracilis, 1 64
— magtia, 164
Halitkea hyslrix, 3.!
Halla, 426
Halosydna, 88
— alleni, 88
— patagonica, 88
Halosyiimlla, 88
hamatiis. Chai'loplerus, 522
hamifera, Eulepis. 45
Haploscoloplos, 546
— fragilis, 546
— kcrguelensis, 547
Haplosyllis, 240
hardyi, Eunereis, 323
Harmopsidcs, 42
Harmothoe, 64
— aeqitist'ta, 66
aequiseta, 66
africana, 68
— africana, 68
— agulhana, 74
— ampullifera, 47
— antilopis, 66
— corralophila, 72
— crosetetuis, 36
— dictyophora, 65
— frascrthomsoni, 74
— gilchristi, 68
— goreensis, 69
— lagiscoides serrata, 69
— lunulata, 71
— profunda, 75
— saldanha, 71
— spinosa, 64
— waahli, 72
— sp., 37
hartmani, Liimbriconereis, 440
hartmani, Lumbrincris, 440
Hauchiclla. 714
— iribullala, 714
helenac, Sthenelais, in8
hclgolandica, Tomopteris, 199
hclleri, Perinereis cuUrifcra. 292
Hemilepidia eryllirutaenia, 55
— tuberculata, 38
Hemipodus, 355
hcmipodus, Xotomastus, 599
henseni. Lopadorhynchus, 161
henseni, Reibiichia, 161
herdmani. Phyllochaetopterus, 524
Htrmaditm fragile, ',9
Hermella capemis, 673
Hermenia, 77
— acantholepis, 77
— verruculosa, 77
Hermione chrysocoma, 33
— hystrix, 30
HERMIOS'liVAE, 30
Hcrmodice, 122
— striata, 1 3 1
Hermonia, 31
— hystrix, 32
Hesione, 227
— genetta, 221
— pantherina, 228
— splendida, 228
HESIONIDAE, 22 I
hesslei, Streblosoma, 725
heteroceros, Hydroides, 807
hiieroceros, Serpula, 807
heterochaeta, Exogone, 233
Helerocirrus bioculatus, 509
— capensis, 498
— ca/}ulesocis capensis, 507
— flauoviridii, 498
— marioni, 505
— zetlandica, 509
Heteromastus, 601
— rilil'ormis, 601
Heleronereis auslralis, 305
Heieropale, 1 16
heteropoda, Dendroncreides, 302
heteropoda, Liimbriconereis, 440
heteropoda allanlica, Lumbrineris, 440
— difficilis, Lumbrineris, 440
— heteropoda, Lumbrineris, 440
heterosetosa, Exogone, 274
HETF.ROSPIONIDAE, 518
Hetromastides, 594
hewilli, Platynereif, 306
hcxapliyllum, Anihosloma, 533
hexaphyllum, .Kainereis, 533
hexaptiylliim, Scolopkn, 533
hexaphyllum, Theodisca, 533
hiltoni, Phyllocomus, 703
hiltoni, Schiiloconuis, 703
Hipponoa, 122
— gaudiehaudi, 122
agulhana, 122
gaudu haudi, 122
hircinicola, Ceratonereis, 327
hircinicola, .Nereis, 327
kolobranchiala, Dio/ialra. 405
holobranchiata, Onuphis, 424
Hololepida, 44
— aiistralis, 44
HuhiUpidelhi nigiopmuldta, 34
INDEX OF SCIENTIFIC NAMES
857
I
hombergi, Nephtys, 344
hoplura hoplura, Polydora, 468
_ inhaca, Polydora, 468
horsli, Perinereis, 334
hupi/eri, Lepidonolus, 37
huhferiaiia, Epidiopalra, 409
hupfcriana hupferlana, Ep.d.opatra, 409
_ monroi, Epidiopatra, 41 1
hubrechti, Eunoc, 62
hubrechti, Evarne, 62
hubrechli, Lagisca, 62
hyalina, Syllis, 246
Hyalinoecia, 411
— tubicola, 411
— sp, 405
Hyalopomaws, 800
Hyboscolex, 588
— longiseta, 588
Hydroides, 804
— bifurcata, 808
— dipoma, 809
— heteroceros, 807
_ lunuUfera, 807
— monoceros, 808
— norvegica, 805
— ralumiana, 807
— uncinata, 805
multispinoia, 789
uminalus macronyx, 798
Hydrophanes krohni, 159
Hyperhalosydna, 88
Hypoeulalia, 154
— bilineat'J, 154
Hypsicomus, 760
— capensis, 761
phaeotaenia, 761
— pigmentatus, 761
hyslncis, Leanira, 112
hystricis, Nephtys, 345
hystrix, Halithea, 32
hyslrix, Hermione, 30
hystrLx, Hermonia, 32
hystrix, SphaerosylUs, 275
hysirix capensis, Sphaerosyllis, 27b
Idanthyrsus, 675
— armatus, 675
— pennatus, 675
impalit-ns, Lwnbricomreis, 439
impatiens, Lumbrineris, 427
implcxa, Filograna, 817
implexa, Serpula, 817
incarunculala, Amphinome, 120
incerta, Goniadopsis, 369
incl^a, Leanira, 113
incisa, Sthenelais, 113
incisa, Sthenolcpis, 113
incruslans, Salmacina dystert, bib
indica, Eunice, 386
indica, Lagisca, 48
imlica, Lycashs, 301
indica, Namalycastis, 301
indica brunnea. Nereis, 320
indica, Pisionella, 133
indica, Pisionidens, 133
indica, SabeUa,i>l?>,n^
indica, Sabellastarte, 771
indica, Scolelepis, 477
indicus, Leonnates, 328
indicus. Lygdamis, 677
indicus gilchnsli,Lygda>ms,bb-]
indicus, Malacoceros, 477
indicus, Paralepidonotus, 48
inermis, Chloeia, 123
inermis, Autolylus, 233
inflala, Lumbriconereis, 435
inHata, Lumbrineris. 435
inBatum, ScaUbregma, 59°
infundibuliformls, Chone, 77b
infundibulum, Myxicola, 773
infimdibidum, St-ypida, &vi
infundibulum, Tcrebella, 773
ingens, TrypanosylUs, 235
inhaca, AUmaniella, 86
inhaca, Polydora hoplura, 4t>8
inhaca, Polynoe, 86
inhaca, Pseudopolynoe, Bb
inontala, Pholoe minula, 91
inomata, Vanadis crystalhna, .82
insignis, Euthaknessa, 107
intermedia, Armandia, 577
intermedins, Lipobranch.us, 58b
intoshi, Sabellaria, 672
inloshi. Sabellaria spinulosa, 672
lOSPILIDAE, 168
lospilus, 168
_ phalacroides, 1 70
Iphione, 43
— muricata, 43
— ovata, 43
— spinosa, 37
Iphitime, 426
Irana, 688
iricolor caerulea, Arabella, 447
_ iricolor, Arabella, 446
iritolor capensis, Maclovia, 443
iricolor, Nereis, 446
Inna angustifrons, 224
cc
858
POLYCHAEIA Ol' SOUTHERN AFRICA
Irniula, 2G6
— spissipcs, 267
isochaeta, Maiq^asia, 164
IsoUla, 6gi
— pulchclla, 691
— wliydahacMisis, 692
isolita, I'lalyiiorcis, 307
jacksoni, I.epidoiiotus, 80
jacksoni, Nereis, 31 1
jn/ionica, Leanira^ 1 12
japonica, Sthenolepis. 1 12
jarli, ;\xiothclIa, 629
Jasmincira, 779
— aualis, 779
— candela, 779
— caiidata, 780
— elegans, 780
jefTrcysi, Aricidea, 558
jelTieysi, Tachytrypane, 580
jejfreyiii, Scolccolepis, 558
Johnstoiiella, 198
johnsoni, Poecilochaetus, 520
johnstonci, Scoloplos, 550
Jolinstonia, 630
— clymcnoides, 63 1
— knysna, 631
Josephella, 817
jousseaumei, Lconnates, 330
jukcsii, Lepidonotus, 80
jukesii, Thorinora. 80
kameruniana, Glycinde, 371
kauikrni, .Nereis, 309
h'tbuUa. 588
kefcrsteini, Stauiciteplialus, 452
Kefersteinia, 228
— cirrata, 228
hfenteinii, TuinopliTi^, 205
kciskama, Ceratoncreis, 327
keinpi, Polydora, 473
kenipi, Tomopteris, 202
kcrguclensis, Ampharete, 700
kergueleiisis, Haploscoloplos, 547
kcrguelensis, Nereis, 309
kergm-lemis, .Vereis, 318
kerguelensis, Scoloplos, 547
kifihergi, Eunue, 377
kinbergi, EupaiUlialis, 94
kinsemboensis, Eullielepus, 726
knysna, Jolinstonia, 631
koreni cirrata, Pectinaria, 682
— koreni, Pectinaria, G81
koreni, Lagis, tj8i
krampi, Tomopteris, 204
kraussi, Glycera. 3")2
kraiissii, Placoslegus cariniferus, 800
kraussii, Poniatoieios, 800
krulirii, Hydroplianes, 159
krohni, Lopadoriiynchiis, 159
kwhni simplex, Lopadorhytuhus, 156
krohni, Syllis, 234
krohni, Trypanosyllis, 253
Krohnia, 177
— angetini, 1 90
— lepldota, 179
kovvalewskii, SagilcUa, 209
kroyeri, Spiophanes, 474
Labidognatlius, 443
lactea, Antinoe, 58
Lacydonia, 350
LACVDONHDAE, 35O
laddi, Uodecaceria, 502
Lacoiiereis, 303
— ankylosela, 303, 323
Laeospira, 796
Lacospira, 794
Laetmonice, 32
— benthaliana, 33
— filicornis, 32
— Jilicornis, 30
— prodiicta, 32
— prodiicta, 33
benthaliana, 33
wyvillei, 33
— wyvillei, 3 1
laeve, Siphonostonntm, 6G1
laevigata, Arieia, 539
laevigata, Naineris, 539
laevis, Lcptoncrcis, 303
laevis, Opisthosyllis, 253
laevis, Pherusa, 661
laevis, Polyeunoa, 54
laevis, Spirorbis, 796
laevis, Slylarioides, 661
Lagis, 681
— koreni, 68 1
Lagisca, 72
Lagisca, 64
— extenuata, 3G
— hubreehti, 1)2
— indica, 48
lagiscoides sorrata, Ilarmotlioe, 69
lagosiana. Ninoe, 431
Lamellisyllis, 237
— comans, 257
lamellosa, Nereis, 314
INDEX OF SCIENTIFIC NAMES
859
laminosa, Phyllodoce, 143
Lanassa, 721
— capensis, 722
— nordenskioldi, 721
lancadivae, Glycera, 359
lanceolata, Travisiopsis, 212
Langerhansia, 243
Inngi, Leodice, 37B
Lanice, 743
— conchilega, 743
— fatwcli, 732
— wollebaeki, 743
Lanicides, 733
Laonice, 480
— cirrata, 480
Laonome, 756
Laphania. 714
lapidaria. Tercbella, 747
lapulatm, Tmbella, 732
Larymna cirrosa, 425
latericeus, Notomastus, 599
lnlipalt>a, Masligonereis, 291
talifalpa lypica, Neanlhes, 292
lalipalpa, Nmis, 292
latreilli, Lumbricomreis, 43B
latrellli, Lumbrineris, 438
Leaena, 721
_ abranchiata, 721
— sp., 721
Leanira, 1 1 1
— hystricis, 112
— incha, 1 1 3
— japonica, 1 1 2
— quatrefagesi, 1 1 1
— telragona, 1 1 3
lefebvreu Nerine. 483
lefebvrei, Scolelepis, 483
Leiochone tenuis, 623
Leiochridcs, 600
— africanus, 600
— australis, 600
Leocrates, 230
— chinensis, 230
— claparedii, 230
Leodamas, 550
Leodice antcnmla, 384
— langi, 37**
Leonnales, 328
— decipiens, 330
— indicus, 32B
— jousscaumei, 330
— persica, 328
Lepidametria, 88
Lepidasthenia, 88
— brunnea, 92
— elegans, 90
— maculata, 92
— microlepis, 90
— mossambica, 89
— sp., 37
Lepidaslheniella, 88
Lepidonola semilecla, 85
Lepidonotus, 79
— ampulliferui, 47
— carinulatus, 81
— clava, 80
— claia, 37
semilecla, 85
— cristatus, 82
— durbancnsis, 85
— glaucus, 84
— hupferi, 37
— jacksoni, 80
— jukesii, 80
— magnatuberculatus, 81
— nalalensis, 82
— platycirrus, 84
— purpureus, 84
— semicinclus, 37
— semitcctus, 85
tenuisetosus, 82
— trissochaetus, 37
— ivahlbergi, 85
lepidola, Alciopa, I79
lepidola, Callizonella, 1 79
lepidota, Krohnia, 179
Leprca plerochaeta, 732
Icptocirrus, Armandia, 577
Leptoecia, 412
— anlarctica, 412
Leptonereis, 305
— laevis, 305
letterstedti, Chone, 777
lelterstedii, Parachonia. 777
Leucodore cHiatus, 469
Leucodorum caecum, 469
leucoplerum, MacrophjUum. 137
Icvenseni, Travisiopsis, 213
ligerica, Boccardia, 463
ligulata, Tomopteris, 204
Uguslica, Orbinia foetida, 533
liguslica, Phylo foetida, 542
liguslica, Phylo foetida. 542
limicola. Sigalion, 1 1 1
limicola, Sthenelais, 1 1 1
lineala, SylHs, 234
lincolala, Amblyosyllis, 233
linguicoUaris, Potamilla, 765
86o
rOLVCHAETA OF SOUTHERX AFRICA
lApobramhius, 586
— ctipaisis, 5M8
— intermedins, 586
— loiigistiu^^ 58J
Loaiidalia. 219
— aberrans, 220
— capcnsis. 220
loandonsis. Brant Iiiomma nigrnmaculata,
loandtmis, Danrlwne, 751
lobala, Pherccardia, 131
lobifcra, SngilcUn, 210
lobifera, Travisiopsis, 2 1 3
Loimia, 742
— medusa, 743
lombricoides, Chnwie, 636
lombricoides, Euclymcne, 636
longa, Sabella, 771
longa, Sabellastarte, 771
longicaudata^ Ammotrypatu\ 577
longicaudata, Armandia, 577
longicinn, Polynof, 58
longicirrata, Pelagobia, 163
longicirrns, Scalisetosus, 58
longipes, Phyllodoce, 144
longpinnis, Glyccra, 356
longirostrosis, Malacoccros, 484
longiseta, Hyljciscolex, 588
lorigiseli/s, LijmbranMus, 583
longissima. Nereis, 323
longissima, Rhynchonerella, 186
longissima, Syllis, 243
longissima, \'anadis, 186
longobranchiata, Aricidea, 560
longocirrata, Cossura, =j8i
longocirrata, Pionosyliis, 264
longocirrata. Syllis, 259
longocirrata, Syllides, 259
LOSCOSOMIDAE. 518
LOPADORHVNCHINAE, I 36
Lopadorhynchus, 158
— appendiculatiis. iGi
— brevis, 162
— hcnscni, i6t
— krohni, 159
simplex, 1 56
— nationalis, 162
— uncinatus, 159
loveni loveni, .Vrenicola, 610
— sudaustraliensis, Arenicola, 610
loveni, Lysilla, 720
loveni, Rhodinc, 619
liicida, Oenone, 426
lucipeta, j\'ereis. 292
luctator, Flabelligera, 652
luclator, Flabelligera affinis. 652
luclalor, Tecturella, 652
hidcriizi, Plerampharele, 697
liideritzi. Sabellides, 697
liiderilziana, Clymene, 641
luderitziana, Eurlymene, 641
himbricalis, Nicomache, 621
lumbricalii eapensis, Michomache, 614
Iwnbricalis, Sabella, 62 1
Lnmbriclymene, 625
— cylindricauda, 625
— minor, 626
Lumbriclymenella, 616
LUMBRICLVMENINAE, 6l6
Lumbriconercis albidenlala, 434
— albi/rons, 435
— atlantica, 440
— brevicirrus, 435
— carifrons, 438
— coecinea. 436
—filum, 447
— harlntani, 440
— heteropoda, 440
— impaliem, 439
— injlata. 435
— lalreilli, 438
— meleorana, 434
— oculala, 436
— paptllijira. 442
— leranrus, 439
— unicornis, 403
Lunibricus armiger, 554
— capitatu-i, 595
— cirralus, 5 1 3
— cirratulus, 483
— filigeriis, 518
—fragilis, 431
— marimis, 608
— squamalus, 483
— syphodonla, 140
LIMBRINERINAE, 426
Lumbrineris, 431
— aberrans, 439
— acuta, 439
— albidentata, 434
— albi/rons, 427
— bifilaris, 431
— brevicirra, 435
— eapensis, 432
— eapensis, 427
— cavifrons, 438
— cavifrons, 427
— coecinea, 436
— debilis, 432
INDEX OF SCIENTIFIC NAMES
86i
— debilis, 427
— dubeni, 427
— floridana, 427
— hartmani, 440
— heleropoda atlanlica, 440
difficilis, 440
heteropoda, 440
— impaliens, 427
— inflata, 435
— latreilli, 438
— magalhaensis, 432
— meteorana, 434
— nardonis, 427
— oculata, 436
— papillifera, 442
— pettigrewi, 427
— tetraura, 439
— letraura. 427
lunulata, Harmothoe, 71
lunulala, Polynoe, 7 1
lunulifera, Eupomatus, 807
lunulifera, Hydroides, 807
Lycastis indica, 30 1
— quadraliceps, 299
Lycoris, 3 1 2
Lygdamis, 675
— indicus, 677
gilchrUli, 667
— murata gilchristi, 677
lyra capensis, Paraonides, 568
— lyra, Paraonides, 568
lyra, Paraonh, 568
lyra capensis, Paraonis, 568
lycrocephala, Clymene, 638
lyrocephala, Euclymene, 638
lyrochaeta, Nephtys, 341
LYSARETINAE, 425
Lysidice, 400
— air a, 378
— capensis, 402
— coUaris, 402
— natalensis, 402
— ninetta, 403
Lysilla, 720
— loveni, 720
— ubianensis, 72 1
Lysippe, 695
— agulhasensis, 700
— capensis, 702
Lysippides, 695
Macellicephala, 44
— mirabilis, 45
macgregori, Unanereis, 299
macintosh!, Marphysa, 396
macintoshi, Nicomache, 6 1 4
macinloshi, Petaloproctus, 614
maclearnus, Autolytus, 284
Maclovia tricolor capensis, 443
tnacrobranchia, Eunice, 377
macrobranchia, Nicolea, 736
macrobranchia, Terebella, 736
macroceros, Eulalia, 152
macroceros, Phyllodoce, 152
macroceros, Sige, 132
Macrochaeta, 501
Macroclymene, 633
— monilis, 633
— saldanha, 634
Macrodymenella, 617
macrodon, Ficopotamus, 810
macrolobata, Pista, 738
macronyx, Hydroides uncinatus, 798
macrophthalma, Eunoe, 64
macrophthatma, Phyllodoce, 149
Macrophyllum leucopterum, 137
— splendens, 151
macroura, Aglaophamus, 343
macroura, Nephtys, 343
niaculata, Goniada, 367
maculata, Lepidasthenia, 92
maculata, Polydora, 472
madagascariensis, .\Iluandella, 280
madagascariensis, Scoloplos, 552
madeirensis, Phyllodoce, 145
magalhaensi, Steggoa, 1 36
magalhaensis, Lumbrineris, 432
magalhaensis, Platynereis, 305
Magalia capensis, 227
— perarmaia, 227
Magelona, 494
— capensis, 497
— cincta, 495
— papillicornis, 495
MAGELONIDAE, 494
magna, Dispio, 482
magna, Haliplanes, 164
magnatuberculatus, Lepidonotus, 81
magnidens, Pionosyllis, 263
magnus, Spio, 482
majorii, Perinereis, 337
majungaensis, Perinereis nimtia, 292
Malacoceros, 477
— fuliginosa, 477
— girardi, 477
— indicus, 477
— longirostris, 484
malardi, Spirorbis, 794
8G-
polvchai:ta of southern Africa
Maldane. 645
— glebifcx, 643
— sarsi, 645
Maldanella, 6'j()
— antarctica. 6'j6
— capensis, 6-^7
— fibrillata, 6:27
MALDAMDAE, 6 I ;}
MALDANINAE. 617
maldivensis. Spirobram hus, 803
malmgrcni, Prionospio, 492
mabngreni diibia, Prionospio, 489
Malmgrcnia, 48
— marqiicsensis, 50
— purpurea. 50
— whiteavesi, 48
mabngreni, Aglaophanius, 343
malmgreni, Nephtys, 343
malmgrcni, Phyllodocr, 147
malmgreni, Pionosyllis, 2O4
Manayunkia, 756
— pacifica, 781
marenzfUcri, FUihtUigaa ajfinis, 632
marina, Arenicola, bio
jnaiini/s, hnnbriius, bo8
rnarioni, Hetcrocirrw^, 505
mariimi, 'Fharyx. 505
Marpbysa, 393
— acicularum, 378
— adenensis, 399
— acnea. 378
— bifurcata, 397
— capcnsis, 400
— corallina. 400
— deprcssa, 395
— durbam-n\is, 378
— haemasoma, 378
— macintoshi, 396
— mossambica, 395
— posterobranchia, 399
— purcellana, 397
— sanguinca, 396
kaemasomn, ;',78
— simplex, 378
marquesen^i^, AUmanuUa, 50
marqucsensis, Malmgrrnia. 30
marsujiiabs, Scoloplos, -,-,0
maryae, Clirrifnrmia, 514
maskallensis, Glvdnde, 3b9
maskallcnsis, (i(niiadf)psis, 369
Alaslignnrreiu 331
— lalipnipa, 291
— nperta, 291
— quadridentata, 293
— striala, 293
malhildae, Sigabnn, 103
matthaii, Eurythoe, 128
Maupasia, 164
— caeca, 164
— gracilis, 164
— isochaeta, 164
maxiUosa, Phyllodoce, 93
Mediomastus. 600
- — californicnsis, 600
— capcnsis, 600
mediterranea, Aphicora, 757
mediterranea, Amphiglcna, 757
medusa, Loimia, 743
medusa, Polycirrus, 715
medusa, TcrcbeUa, 743
Megalomma, 757
- — bioculatum, 760
— quadrioculatum, 758
— vesiculosum, 758
melanonotus, Panthalis, 96
mclanonotus, Polyodontes, 96
mclanura, Armandia, 576
Melinna, 689
— - cristata, 691
— monoccroides, G89
— palmata, G89
MF.LINNINAE, 688
Molinnopsidcs, 692
— capcnsis. 693
Molinnopsis, 689
— capcnsis, 693
nictidax, Acreis, 293
niendaw Pcrinereis, 293
Mercierella, 812
— enigmatica, 812
mcridionalis^ Audouinia /iligrra, 499
rticridionalis, drralubts /rritaculiis, ^99
Mesocbaetopterus, 531
■ — capcnsis, 532
— minutus, 531
- — sagittaria, 531
— taylori, 531
mcleorana, Lumbriconrrei'i, 434
mcleorana, Lumbrineris, 434
michaelscni, Myxicola, 732
microcephala, Clymcne, 614
microcephala, Eulalia, 132
microcephala, Pscudcur^-thoc, 126
Microclymcnc, 617
microlepis, Lepidastbenia, 90
Micromaldane, 6ig
— ornithochaeta, 619
INDEX OF SCIENTIFIC NAMES
863
— sp, 62 1
Micronephthys, 347, 349
Micronereides, 298
— capensis, 298
Microiiereis, 297
Microphthalnius, 223
rnicrops, Anlinoe, 57
Microsamytha, 695
minor, Lumbriclymene, 626
tninula, Aphrodita, 100
minuta, Pholoe, 100
tninula inornala, Pholoe, 97
minuta, Vanadis, 184
minuta, Vanadis, 184
minutus, Mesochaetoptcrus, 531
mirabilis, Ceratonereis, 324
mirabilis, Macellicephala, 45
mirabilis, Polynoe, 45
misakiensis, Tryjianosyllis, 256
mobii.Callizona, 189
mobii, Rhynchonerella, 189
moebii, Callizona, 189
moebii, Rhynchonerella, 189
monilaris, Syllis, 239
monilicornis, Eusyllis, 233
moriilis, Clymene, 633
monilis, Macroclymene, 633
monoceroides, Mclinna, 689
monoceros, Hydroides, 808
monoceros, Serpula, 808
monroi, Diopatra, 415
monroi, Drilonereis, 448
monroi, Epidiopatra liupferiana, 41 1
monroi, Orbinia, 545
monroi, Phenisa, C60
monroi, Stylarioides, 660
rnossambica, Clymene, 639
mossambica, Euclymene, 639
rnossambica, Eunice, 395
mossambica, Fabricia, 781
mossambica, Fabriciola, 781
mossambica, Gattyana, 61
mossambica, Lepidasliicnia, 89
mossambica, Marphysa, 395
mossambica, Nereis, 318
mossambica, Nicomache, 622
mossambica, Sabella, 752
muelleri, Typhloscolex, 208
multiannulata, Gravierella, 630
multidens, Glycinde, 371
mullidentala, Aracoda, 443
muliiJUis, Tharyx, 499
rnulli.-ipinosa, Hydroides uncinata, 798
murata gilchristi, Lygdamis, 677
murata gilchristi, Tetreres, 677
muricata, Iphione, 43
muricata, Polynoe, 43
murrayi, Eunice, 385
mushaemis, Branchiomma, 758
musseraensis, Diopatra, 417
mutans, .Arabella, 446
mitlans, Cenothrix, 446
myrialcpis, Bhavvania, 1 18
Myrianida, 285
— phyllocera, 287
— pulchella, 287
Myriochele, 649
Myriowenia, 649
myrtosa, Euphrosine, 127
Mysta, 140
— siphonodonia, 1 40
Mystides, 141
— angolaensis, 141
— bidentata, 143
— borealis, 141
Myxicola, 773
— infundibulum, 773
— michaeiseni, 752
MYXICOLINAE, 755
Naiades, 176
— cantrainii, 176
naidina, Exogone, 271
J^'aidonereis, 533
Nainereis hexaphyllum, 533
Naineris, 539
— laevigata, 539
— quadricuspida, 539
Nais, picta, 579
— quadricuipida, 539
Namalycastis, 301
— indica, 301
Nainanereis, 299
— quadraticeps, 299
nardonis, Lumbrineris, 427
nasuta, Callizona, 1 89
natalensis, Branchiomma, 767
natalerisis, Chloeia, 1 20
natalensis, Clymene, 638
natalensis, Dasychone, 767
natalensis, Euclymene, 638
natalensis, Glycera, 356
natalensis, Lepidonotus, 82
natalensis, Lysidicc, 402
natalensis, PharyngeovaKata, 239
natalensis, Sabella, 767
natalensis, Terebellobranchia, 750
nationalis, Lopadorhynchus, 162
864
roi
.VCHAF.TA OF SOUTHERN AFRICA
nationalis, Tiiinopteiis. 198
Xauphanli! coralUna, 400
Ncanlhcs, 317-323
— latipdpa typica, 292
ncapolilana capensis, Diopatra, 413
— neapolitana, Diopatra, 413
neapolitana. Paraonides, 568
ntapoUlana, Ptiraoiiis, 566
neapolitana, Pcctinaria, 681
ncbulosa, Eupolymnia, 744
nebtdom, Polymnia, 744
ncbulosa, Terebclla, 744
ncglecta, DorviUea, 455
ncglecta, Oriopsis, 785
neglecta, Sabella, 764
neglfcliis, Staiirocephalm, 455
nematodes, I'roccrastca, 280
jS'ematonereis, 403
— unicornis. 403
— sp. 378
Neoleprea, 733
Neopomatus, 800
Neosabellides, 695
— etongiitm, 703
Xeovermilia, 814
— capensis, 816
— falcigera, 816
jXephlhys, 34 '"349
NEPHT\'IDA£, 338
Nephtys, 340
— ambrizettana, 349
— capensis, 344
— dibranchis, 341
— hombergi, 344
— hystricis, 345
— lyrochacta, 341
— macroura. 343
— malmgreni. 343
— paradoxa, 347
— sphaerocirrata. 347
— tulearcnsis, 345
NEREIDAE, 29 I
.Xtrcilepas vmicgala, 331
Nereis, 307
— agnesiae, 32 1
— agulhana, 318
— alba. 360
_ ambylodonta, 333
— aphrodiloii. 389
— arena.codonia, 321
— armilhirii, 249
— callanana, 292
— capt'Jisis, 320
— caudata, 321
— ciliala, 340
— coccinea, 436
— conchilega, 743
— coslae, 325
— coulicrci, 31 1
— cuUrifera, 337
— culvcri, 303
— cuprea, 417
— cylindraria bclgica, 680
capensis, 683
— diversicolor, 317
— dumeuUi, 306
— eugeniae, 312
— falcaria, 309
— falsa, 317
— jUicornii, 481
— JIava, 139
— fle\iiosa. 223
— fusifera, 323
— gilchristi, 3 1 5
— glandulosa, 32 1
— granulata, 309
— hircinicola, 327
— indica brunnea, 320
— iricolor, 44O
— jacksoni, 31 1
— kauderni, 309
— kcrguclensis, 309
— kergiielemis, 318
— lamcllosa, 314
— lalipalpa, 292
— longissima, 323
— lucipela, 292
— nicndax, 293
— mossambica, 318
— nigropunctata, 337
— iionegka, 388
— operla, 323
— papillosa, 317
— pelagica, 315
— pennala, 388
— persica, 314
— pinnigcra, 285
— podocirra, 293
— polyodonla. 323
— prolifcra, 284
— rudolphi, 457 ^
— sangidnea, 396
— spUndida, 292
— squamosa, 52
— stimpsonis, 293
— succinea. 321
— tongalabnensis, 293
— trifasciata, 312
INDEX OF SCIENTIFIC NAMES
865
— tubicola, 4 1 1
— unifasciata, 318
— unifasciata, 3 1 2
— lallata, 334
— vancaurica, 334
— variegala, 331
— liridis, 1 5 1
— viltaia, 385
— willeyi, 320
— zonata persica, 314
— ■s/'-3i5
Ncrimyra, 223
Nerindes, 484
— cantabra, 485
— gilchristi, 485
— tridentata, 484
— williami, 485
J^erine, 482
— cirrata, 480
— cirratulus, 483
capensis, 460
— lefebvrei, 483
— oxvcephala, 478
nesideiisis, Amphitriu, 744
Nicidion, 382
— cincia, 382
Nicolea, 735
— ctaparedii, 732
— macrobranchia, 736
— quadrilobata, 740
— venustula africana, 736
venustula, 735
Nicomache, 62 1
— capensis, 6 1 4
— lumbricalis, 621
capensis, 6 1 4
— macintoski, 614
— mossambica, 622
NICHOMACHINAE, 617
JVicon eugeniae, 312
nigromaculata loandensis, Branchiomma, 751
nigromaculata, Dasychone, 770
nigTomaculata, Sabella, 770
nigropharyngea, Syllis, 249
nigropunctata, Hololepidella, 54
nigropunclata, .Xereis, 337
nigropunctata, Perinereis, 337
nigropunctata, Polyeunoa, 54
nigropunciala, Polynoe, 54
ninetta, Lysidice, 403
Ninoe, 430
— chilensis, 430
— lagosiana, 431
nisseni, Tomopteris, 202
DD
nodosa, Polynoe, 6 1
nodulosa, Eunoe, 62
nordenskioldi, Lanassa, 721
normalis, Exogone, 275
normalis, Polydora, 471
norvegica, Eunice, 388
norvegica, Hydroides, 805
non^egica, J\'ereis, 388
Nothria, 424
— conchy lega, 425
— geophili/ormis, 424
Notocirrus, 448
— australis, 448
— brevicirrus, 435
— capensis, 443
— chilensis, 448
— scoticus, 448
— telraurus, 439
Notomastus, 597
— aberans, 599
— fauveli, 597
— giganleus, 597
— hemipodus, 599
— latericeus, 599
Notophyllum, 150
— foliosum, 151
— splendens, 151
Notoproctus, 647
— oculatus, 647
Notopygos, 130
— crinita, 130
— variabilis, 130
novaehollandiae, Perinereis, 333
novecrinita, Arabella, 446
nuntia majungaensis, Perinereis, 292
— vallata, Perinereis, 334
nyanganus, Pseudothelcpus, 722
oahuensis, Greejfia, 1 72
Octobranchus, 71 1
octocirrata. Sabella, 697
octocirrata, Sabellides, 697
oculala, Lumbriconereis, 436
oculata, Lumbrineris, 436
oculata, Paraonis gracilis, 565
oculata, Phyllodoce, 137
oculata, Sthenelais, 97
oculata, Thalencssa, 107
oculatum. Si gallon, 107
oculatus, Notoproctus, 647
odhneri, Dasychone, 771
Odontosyllis, 260
— ctenostoma, 261
— ctenostoma, 260
866
POLVCHAKTA OF SOUTHERN AFRICA
— gibba, 261
— graveleyi, 260
— polyccra, 260
Oenone, 426
— fulgida. 426
— hicida, 426
oerstedi, Panthalis, 96
oerstedi capcnsis, Panthalis, 97
oerstedi, F.uclymene, 635, 636
oerstedi, Pisione, 133
oersledi, Theostoma, 533
Ofrstedii, Clvmerte, 635
oligobrcmcktis. Bratichoscolex, 591
Omphalopomopsis, 799
OncoscoUx, 588
— hif>artiliis, 603
onisciformis, Tomopteris, 197
ONIPHIDINAE, 405
Onupliis, 421
— conchylega, 423
— eremita, 422
— geophiliformis, 424
— gracilis, 412
— holobranchiata, 424
— quadriciispis, 42 1
— quinquedcns, 422
— tubkola, 405
opcrla, Masiigimereis^ 291
operta, Nereis, 323
Ophelia, 571
— africana, 575
— agulhana, 573
— anoniala, 572
— bicornis, 572
— capensis, 573
— peresi, 573
— roscoffcnsis, 572
OPHELIIDAE, 570
Ophelina, 579
— acuminata, 579
Ophiodromus, 223
— agilis, 226
— angustifrons, 224
— berrisfordi, 224
— spinosus, 223
Ophioglycera, 372
— eximia, 372
— gigantea, 372
Ophryotrocha. 451
— pnerilis, 452
Opisthosyllis, 252
— antylociiaeta, 2')2
— brvinnea, 253
— laevis, 253
orbkulata, Acholoe, 36
Orbinia, 543
— angrapequensis, 543
— bioreti, 545
— cuvieri, 543
— dubia, 547
— foetida ligiistka, 533
— monroi, 545
Orbiniella, 535
ORBIMIDAE, 533
ORBINIINAE, 539
Oria eimeri, 787
— parvula, 787
Oridia ca/>ensis, 784
— parvula, 752
orieiilalis. Dasychone, 7 5 1
Oriopsis, 785
— bansei, 788
— ehlersi, 787
• — eimeri, 787
■ — ncglecta, 785
■ — parvula, 787
— parvula, 788
ornata, Desdemona, 790
ornata, Eteone, 140
ornithochaeta, Micromaldane, 619
Orseis, 223
Orthrodromus spirwsiis, 223
ovata, Iphione, 43
Owenia, 649
— fusiformis, 649
OWENIIDAE, 649
oxycephala, Aonides, 478
oxycephala, .Xerine, 478
Oxydromm, 230
— capcnsis, 23 1
— 5/)., 232
pacifica, Aricidea, 563
pacifica, Manayunkia, 781
pachychaeta, Ceratonereis, 325
pacifica. Tomopteris, 199
Faedophylax veruger, 272
Paleanotus, 1 16
— chrysolepis, 1 16
— debilis, 1 17
pallasi, Amphinome, 123
Pallasia, 675
— capensis, 667
— pennata, 667
pnllciccTis, ,Samylha, 703
palmata, Melinna, 689
Palmyra, 1 16
— debilts, 1 1 7
INDEX OF SCIENTIFIC NAMES
867
PALMYRIDAE, I 1 5
Palolo, 382
Panthalis, 96
— rnelanonotiis, 96
— oerstedi, 96
capensis, 97
pantherina, Hesione, 228
papillatus, Polyophthalmus, 570
papiilicornis, Magciona, 495
papilli/era, Lumbricomreis, 442
papillifera, Lumbrineris, 442
Papillodonillea, 454
papillosa, Epidiopatra, 408
papillosa, Glycera, 358
papillosa, Nereis, 317
papillosa, Pectinaria, 685
papillosa, Sthenelais, 108
Parachonia letterstedli, 777
Paradoneis, 566
paradoxa, Nephtys, 347
paradoxa, Paralacydonia, 350
Paraeurylhoe ckitensis, 130
Parahalosydna, 86
— capensis, 86
— sibogae, 86
Parahesione, 223
Paralacydonia, 350
— paradoxa, 350
Paralaeospira, 794
Paralepidonotus, 47
— ampulliferus, 47
— indicus, 48
Paramarphysa, 380
Paramphinome, 121
Paranaites, 148
Paranereis abiuma, 301
PAR-AONIDAE, 555
Paraonides, 566
— lyra capensis, 568
— lyra, 568
— neapolitana, 568
— sp., 555
Paraonis, 565
— belgicae, 563
— fulgens, 565
— gracilis gracilis, 566
oculata, 565
— lyra, 568
capensis, 568
— neapolitana, 566
— tenera, 565
Parasclerocheilus, 585
— branchiatus, 585
— capensis, 585
parashadi, Glycera, 359
parasitica, Alciopina, 1 77
Pareulepis, 45
— geayi, 45
Parheteromastus, 605
— tenuis, 605
Pariospilus, 168
— ajjinis, 170
parmata, Pherusa, 658
pammtus, Stylarioides, 658
Parmensis capensis, 66
— reticulata, 36
Paronuphis, 412
— antarctica, 412
parva, .'Vncistrosyllis, 216
parvecarunculata, Eurythoe, 1 28
parvula, Oria, 787
parvula, Oridia, 752
par\'ula, Oriopsis, 787
parvula, Oriopsis, 788
patagonica, Halosydna, 88
patagonica, Phyllodoce, 137
patagonicus, Spirorbis, 794
paucibranchiata, Pseudeurythoe, 126
pauciseta, Amphitrite, 746
pauliani, Schroederella, 538
paupera, Eunice, 393
paupera, Eunice afra, 393
pavonina, Sabella, 763
Pectinaria, 680
— antipoda, 685
— capensis, 683
— koreni cirrata, 682
koreni, 681
— neapolitana, 681
— papillosa, 685
— pseudokoreni, 68 1
PECTINARIID.\E, 678
pectinata, Sabellaria, 671
Pedinosoma, 158
— curtum, 158
pelagica, Drieschia, 75
pelagica. Nereis, 315
pelagica, Pontodora, 167
Pelagobia. 163
— longicirrata, 163
pellucidus, Scalisetosus, 59
penicillus, Sabella, 763
pennata, Eunice, 388
pennata. Nereis. 388
pennata, Pallasia, 667
pennata, Sabellaria, 675
pennatus. Idanthyrsus, 675
pequcnianus, Thelepus, 730
868
POLVCHAETA OF SOUTHERN AFRICA
perarmala, Mdgalia, 227
pcresi, Ophelia, 573
Perinercis, 333
— capensis, 335
— cultrifera, 337
helleri, 29-2
— diversidentata, 293
— lalsovaricgata, 335
— honti, 334
— majorii, 33"
— mendax. 293
— nigropunctata, 337
_ mvaehollandiae, 333
_ nuntia majungaensis, 292
valuta, 334
— lallala, 292
— vancaurica, 334
— lariegata, 293
perricri, Procerastea, 280
persica, Grymaea, 725
persica, Leonnates, 328
persica, Nereis, 314
persica, Nereis zonata, 314
persica, Streblosoma, 725
perspicax, SphaerosylUs, 277
Petaloproctus, 622
— miicintoshi, 614
— terricola, 622
petersi. Psammolyce, 105
petersi, Rhynchonerella, 192
pelersii, AUiopa, 192
Petta, 680
petligreivi, Lumbrincris, 427
phaeotaenia, H>-psicomus, 761
phaeotaenia. Sabella, 761
phalacroides, lospilus, 170
Phalacrophorus, 170
— pictus, 171
— uniformis, 1 7 1
Phalacrosiemma, G69
cidariophilum, 669
— elegans, 669
Pharyngeovalvata, 239
— natalensis, 239
Phenacia ielosa, 729
Pherecardia, 131
— lohata, 131
— striata, 131
Pherusa, 658
— laevis, 661
— monroi, 660
— parmata, 65B
— saldanha, 663
— swakopiana, 661
— tetragona. 652
— tropica, 663
Phisidia, 714
Pholoe, 99
dorsipapillata, 100
— minuta, 100
inornata, 97
Phragmatopoma, 668
Phyllampliictcis, 695
phylloccra, MyTianida, 287
Phyllochaetopterus, 524
— elioti, 525
— gracilis, 524
— herdmani, 524
— pictus, 522
— socialis, 525
Phyllocomus, 702
— crocea, 702
— hiltoni, 703
phyllodes, Tv^hloscolex, 209
Phyllodoce, 143
— qfricana, 137
— hilineala, 1 54
— capensis, 148
— castanea, 149
— fotiosa, 151
— fristedti, 147
— laminosa, 143
— longipes, 144
— macroceroi, 152
— macrophthabna, 1 49
— madeirensis, 143
— malmgrcni, 147
— maxillosa, 95
— oculata, 137
— paUigonica. 137
— quadraticeps, 143
— rubiginoui, 137
— schmardaei. 149
— tubicola, 148
PHYLLODOCIDAE, 1 36
PHYLLODOCINAE, I 36
Phylo, 540
— capensis, 540
— fclix, 540
foetida australis, 342
foetida, 542
ligustica, 542
Phyzelia, 742
picla, Cirroiyllis, 233
picta, Afl", 579
picla, Proceraea. 284
pictus, Autolytus, 284
pictus, Phalacrophorus, 1 7 1
INDEX OF SCIENTIFIC NAMES
869
piclus, Phyllochaetoptems, 522
pictus, Polyophthalmus, 579
pigmentatus, Hypsicomm, 761
PILARGIDAE, 2I4
Pilargis, 218
— falcata, 218
— verrucosa, 218
pinnata, Prionospio, 488
pinnigera, Nereis, 285
Pionosyllis, 261
— compacta, 261
— ehlersiaeformis, 263
— ehlersiaeformis, 264
— longocirrata, 264
— magnidens, 263
— malmgreni, 264
— sp., 234
Piromis, 663
— arenosus, 664
— sp., 664
Pisione, 133
— africana, 133
— oerstedi, 133
Pisionella, 132
— indica, 133
PisiONiDAE, 132
Pisionidens, 132
— indica, 133
Pista, 736
— brevibranchia, 737
— cristata, 738
capensis, 732
— fasciata, 742
— foliigera, 740
— grubei, 732
— macrolobata, 738
— qolora, 740
— quadrilobata, 740
— typha, 737
— unibranchia, 737
Placoslegiis caerideus, 80 1
— cariniferus kraussii, 800
plagiostoma, Terebella, 729
plagiostoma, Thelepus, 729
planktonis, Tomopteris, 206
plalycirrus, Lepidonotus, 84
Platynereis, 305
— australis, 305
— calodonia, 306
— dumerilii, 306
— hewitli, 306
— isolita, 307
— magalliaemis, 305
— tongatabuensis, 293
Plotohia coniceps, 210
Plotohelmis, 192
— alata, 193
— capitata, 195
— tenuis, 193
plumosa, Amphitrite, 658
plumosa, Ereutho, 718
plumosus, Polycirrus, 718
Podarke, 223
— agilis, 226
— angmtifrons, 224
podocirra, Mereis, 293
Poecilochaetus, 519
— fulgoris, 520
— johnsoni, 520
— serpens, 520
polybranchia, Boccardia, 463
polybranchia, Boccardia, 463
po/ybranchia, Eiiphrosine, 120
polycera, Odontosyllis, 260
polycera, Syllis, 260
POLYCIRRINAE, 713
Polycirrus, 715
— aurantiacus, 715
— haematodes, 7 1 7
— medusa, 715
— plumosus, 718
— swakopianus, 7 1 7
— tenuisetis, 715
— Iribullata, 714
— trilobata, 718
Polydora, 465
— antennata, 473
— armata, 466
— caeca, 469
— capensis, 466
— ciliata, 469
— colonia, 468
— cornuta, 465
— flava, 468
— giardi, 471
— hoplura hoplura, 468
inhaca, 468
— kempi, 473
— maculata, 472
— normalis, 471
— polybranchia, 463
Polydorella, 462
Polyeunoa, 52
— laevis, 54
— nigropunctata, 54
Polymnia capensis, 732
— nebulosa, 744
Polynoe, 55
870
POLVCHAEIA or SOUTHKRN AIRICA
— acantholej>is, 77
— agnae, 38
— amjiulliftra, 47
— attemidta, 38
— cafiemis, 86
— caputleonis, 38
— caririuldta, 81
— cristaia, 82
— dictyophonis, 65
— etegans, 90
— erthrotaenia, 55
— giicithiosa, 44
— glauca, 84
— inhaca, 86
— longicirra, 58
— liimilata, 7 1
— mirabilis, 45
— muricata, 43
— nigropunctata, 54
— nodosa, 61
— scolopendrina, 55
— trvchiscophora, 83
POLYNOIN.\E, 36
pohodunia, J\'ereis, 323
Polyodontes, 94
— mclanonotus, 96
POLYODONTINAE. 93
Polyophthalmus, 579
— papiilalus, 570
— pictus, 579
Polyphysia, 586
— crassa, 588
Pomatoceros, 801
— caeruleus, 801
— caeruleus, 798
— cariniferus, 801
— slrigkeps, 801
— tetraceros, 803
Pomatoleios, 800
— crosslandi, 800
— kraussii, 800
Pomatostcgus, 800
Pontodora, 167
— pelagica, 167
rONTODORIDAE, 1 67
Pontogenia, 35
— chrysocoma, 35
port/era, Sabella, 764
postcrobranchia, Marpliysa, 399
Potaniilla, 764
— ehtersi, 752
— lingiiicollaris, 763
— reniformis, 764
— torelli, 766
praclermissa, Clymene, 642
praetermissa capensis, Clymene, 6 1 3
praelermissa, Praxilla, 642
praclermissa, Praxillclla, 642
praelermissa capensis, Praxillclla, 613
prampramensis, Trypanosyllis, 255
prashadi, Glycera, 359
Praxilla capensis, 644
— praelermissa, 642
Praxillclla, 642
Praxillclla, 634
— alBnis, 644
— capensis, 644
— praelermissa, 642
capensis, 6 1 3
Praxillura, 616
Prionospio, 485
— bocki, 490
— capemis, 459
— cirrifera, 486
— cirrobranchiata, 48H
— chlersi, 490
— malmgrcni, 492
diibia, 489
— pinnala, 488
— saldanlia, 492
— scxoculala, 489
— sicenstrupi, 489
— sp., 492
— ,1/)., 460
proboscidea, Arlacama, 733
Proceraea auranliaca, 283
— picta, 284
Proceraslea, 280
— nematodes, 280
— perricri, 280
Proclca, 714
Proclymene, 617
producia, Clymene, 633
producla, Laelmonice, 32
producia, Laelmonice, 33
producla benlhaliana, Laelmonice, 33
— u-yvillei, Laelmonice, 33
profunda, Harmolhoe, 75
Progciniada, 333
prolifer, Aulolytus, 284
prolifera, Mereis, 284
prolifcra, Syllis, 248
Prolopadorhynchus, 161
propinqua, Gyplis, 231
Proscoloplos, 336
— cygnochaelus, 538
Protoaricia, 333
PROTOARICIINAIL, 536
INDEX OF SCIENTIFIC NAMES
87.
Protocapitetta simplex, 608
Protodorvillea, 452
— biarticulata, 452
— egena, 454
Protomystides, 143
— capensis, 143
Protula, 818
— bispiralis, 8i8
— capensis, 8 1 6
— dysteri, 8 1 7
— tubularia anomala, 820
capemis, 8 1 6
tubularia, 820
Psamathe cirrata, 228
Psammolyce, 105
— articulata, 105
— flava, 105
— petersi, 105
Pseudeurythoe, 124
— microcephala, 126
— paucibranchiata, 126
Pseudocapitella, 594
pseudokoreni , Pectinaria, 68 1
pscudonatrix, Boccardia, 465
Pseudonereis, 331
— anomala, 333
— gallapagensis, 33 1
— variegata, 331
Pseudopolynoe, 86
— inhaca, 86
Pseudoscalibregma, 585
Pseudothelepm nyangamis, 772
Pterampharete, 697
Pltrampharete, 696
— luderitzi, 697
pierochaeta, Leprea, 732
pierochaela. Schmardanella, J^j
pterochaeta, Terebella, 747
pierochaela, Terebella, 732
Pterocirrus, 152
Plerosyllis formosa, 259
puerilis, Ophryotrocha, 452
pulchella, Isolda, 691
pulrhella, Myrianida, 287
pulchra, Dodecaceria, 502
pulchra, Fauveliella, 132
Pulliella, 595
— arniata, 595
punclala, Audouinia, 517
punctata, Cirriformia, 517
punclala, Eunice, 393
punctata, Eunice afra, 393
punclalus, Cirratulus, 5 1 7
punclifcra, Diopaira, 418
punctifera, Diopatra cuprea, 418
purcellana, Marphysa, 397
purpurea, Malmgrenia, 50
purpureus, Lepidonotus, 84
pusilla, Brania, 267
pusilla, Exogone, 267
pusilla, Grubea, 267
Pycnoderma, 655
— congoense, 656
pygidialis, Vermiliopsis, 813
pygidialis, Vermiliopsis, 813
Pygospio, 475
— elegans, 475
qolora, Pisia, 740
quadraliceps, Lycaslis, 299
quadraticeps, Namanereis, 299
quadraticeps, Phyllodoce, 145
quadraliceps, Sphaerodoce, 1 45
quadricuspida, Naineris, 539
quadricuspida, Mais, 539
quadricuspis, Onuphis, 421
quadridenlala, Masligonereis, 293
quadrilobata, Euclymene, 638
quadrilobala, Nicolea, 740
quadrilobata, Pista, 740
quadrimaculata, .Vxiothella, 629
quadrioculalum, Branchiomma, 758
quadrioculatum, Megalomma, 758
quatrefagesi, Leanira, 1 1 1
quinquedens, Onuphis, 422
ralumiana, Hydroides, 807
Ranzania, 531
— capensis, 532
Ranzanides, 531
— capensis, 532
Reibischia henseni, 161
renifonnis, Amphilrile, 764
reniformis, Potamilla, 764
reniformis, Sabella, 764
reliculata, Parmensis, 36
reynaudii, Alciopa, 180
Rhamphobrachium, 418
— agassizi, 418
— bipes, 418
— capense, 418
— chuni, 420
— diversoselosum. 418
Rhodine, 618
— gracilior, 619
— loveni, 6 1 9
RHODINlN.\E, 616
Bya
POLYCHAETA OF SOUTHERN AFRICA
rhombeata, Amblyosyllis, 257
rhombigera, Enipio, 54
rhopalophora, Brania, 269
rhopahphora, Grubea, 269
RhynchonercUa, 188
— an^clini. 190
— capitala, 1 95
—fulgais, 195
— gracilis, 189
— longisnma, 186
— miibii, 189
— moebii, 189
— petersi, 192
Rhynchospio, 478
— arenincola, 478
— glutaca, 478
rigida, Ancistrosyllis , 2 1 -,
robusta, Ancistrosyllis, 216
roscoffensis. Ophelia, 572
rosea, Euchone. 774
rostrata, Aniphinome, 123
roslrata, Aphrodita, 123
roiisseaui, Eunice, 389
rouxi, Glycera, 362
rubiginosa. Phyllodoce. 137
rubrovittata, Dorvillea, 457
rubroviltatm, Staurocephalus, 437
rubroviltalus, Stauronereh, 450
rudolphi, Dorvillea, 457
rudotphi, .Vereis, 457
rudolphi, Staurocephalus, 457
Sabella. 763
— aheolata, 67 1
— analis, 774
— capensis, 678
— chrysodon, 67H
—foliifera, 75 1
— fusca, 764
— indica, 678, 771
— longa, 771
— lumbricalis, 62 1
— mosiambica, 752
— natalemis, 767
— neglecta, 764
— nigrontaculala, 770
— ociocirrata, 697
— paionina, 763
— pcnicillus, 763
— phaeotaenia, 761
— porijera, 764
— renijormis, 7^4
— serralibranchii, 768
— lorclli, 7b*j
— violacca, 768
sabella, Amphicora, 782
Sabellaria, 671
— alcocki, 672
— bella, 672
— capertsis, 673
— eupomatoides, 667
— fucicola, 672
— guinensis, 672
— intoshi, 672
— pectinata, 671
— pennala, 675
— spinulosa alcocki, 672
eupomatoides, 667
fucicola, 672
gilchrisli, 667
intoshi, 672
SABELLARIIDAE, 667
SabcUastarte, 770
— indica, 771
— longa, 77 1
— sanctijoseplii, 771
SABELLIDAE, 751
Sabellides, 696
— capensis, 698
— cristata, 691
— luderitzi, 697
— octocirrata, 697
— 5/-., 693
SABELLINAE, 755
sagittaria, Mesochaetopterus, 531
Sagitella, 209
— cornuta, 2 1 2
— kowalewskii, 209
— lobijera, 2 1 o
saldhana. Clymene, 614, 634
saldanha, Harmothoe, 71
saldanlia, Macroclymene, 634
saldanha, Pherusa, 663
saldanha, Prionospio, 492
Salmacina dysteri, 8 1 7
Salmacina dysteri incruitan^. 816
Samytha, 695
— pallescens, 703
SamythcUa, 704
— affinis, 704
— clongata, 704
sanctijou'phi, Euralo, 771
sanclijosephi, Sabellastarte, 771
sanguinea, Eulalia, 155
sanguinea, Eumida, 155
sanguinea, Marphysa, 396
sanguinea haemasoma, Marphysa, 378
sanguinea, jVereis, 396
INDEX OF SCIENTIFIC NAMES
873
sarsi, Maldane, 645
savignyi, Eunice, 378
saxaiilis, Audouinia, 515
saxatilis, Cirriformia, 515
Scalibregma, 589
_ inilatum, 590
SCALIBREGMIDAE, 583
Scalisetosus, 58
— ceramensis, 58
— communis, 38
_ fragilis, 59
— longicirrus, 58
— pellucidus, 59
schemacephala, Eunice, 384
Schislocomus, 702
— hiltoni, 703
schmardaei, Phyllodoce, 149
schmardaei, Terebella, 748
SchmardaneUa pterochcuta, 747
Schroederella, 538
— pauliani, 538
Sclerocheilus, 585
Scolaricia, 547
— capensis, 549
— dubia, 547
— typica, 547
Scolecolepis glutaea, 478
— jefjreym, 558
Scoielepis, 482
Scolelepis, 477
— indica, 477
— lefebvrei, 483
— squamata, 483
scolopendrina, Polynoe, 55
Scoloplella, 536
— capensis, 536
Scoloplos, 549
— armiger, 554
— cylindrifer, 546
— hexaphyllum, 533
— johnstonei, 550
— kerguelemis, 547 ^
madagascariensis, 552
— marsupialis, 550
— uniramus, 552
— sp., 534
scoticus, Notocirrus, 44a
scutata, Sternaspis, 648
sculala africana, Stemaspu, 648
sculatus, Echinorhynchus, 648
Scyphoproctus, 604
— djibouticnsis, 604
— stcinitzi, 604
semicincla, Audouinia, 517
semicinctus, Lepidonolus, 37
semitecta, Lepidonota, 85
semilecta, Lepidonolus clava, 85
semitectus, Lepidonolus, 85
semiverrucosa, SphaerosylUs, 276
semperi, Spirobranchus, 803
septentrionalis, Tomopleris, 205
serpens, Poecilochaetus, 520
Serpula, 809
— bispiralis, 818
— complexa, 817
— echinata, 810
— filograna, 817
— gigantea, 803
— heleroceros, 807
— implexa.Ril
— infundibulum, 813
— monoceros, 808
— triquelra, 801
— lubularia, 820
— uncinata, 807
_ vermicularis echinata, 810
vermicularis, 809
SERPULIDAE, 791
SERPULINAE, 798
serrata, Harmothoe lagiscoides, bg
serratibranchis, Branchiomma, 7^8
serratibranchis, Dasychone, 768
serratibranchis, Sabella, 768
setabulensis, Eusyllis, 235
setabulensis, Euthelepus, 72b
selosa,Callizona, 192
setosa, Chaetozone, 510
setosa. Phenacia, 729
setosus, Thelepus, 729
selosm africanus, Thelepus, 722
sexoculata, Prionospio, 489
sexoculata, SylUs, 235
sibogae, Parahalosydna, 86
siciliensis, Eunice, 382
Sigalion, loi
— boa, 109
— capense, 103
— edwardsi, 107
— limicola, 1 1 1
— mathildae, 103
— oculalum, 107
— squamatum, 104
— tetragonum, 1 13
SIGALIONINAE, 97
Sigambra, 216
Sige, 155
— macroceros, 1 52
simplex, Aricidea suecica, 558
I'tn.VCHAE lA OF SOUTHER.N Al RICA
simplex, Lo/mJorhynchos krohni, 15G
simplex, Marphysa, 378
simplex, Protocapiteila, 608
simplex, Siheiielais, i 1 3
siphonodonla, Eteone, 140
siphunodonla, Mvsta, 140
Siphofwstomum laeve, 66 1
— villosuTu, 656
socialis, Phyllochactopterus. -j^j
soederstromi, Spiophanes, 473
Sosane, 695
Spermosyllis, 270
— capensis, 270
— torulosa, 270
spehbergensis. Eleone, 136
sphaerochaetus, Branchoscolex. 59 1
sphaerocirrata, \eplitys, 347
Sphiurodoce quadraticeps, 145
SPHAERODORIDAE, 288
Sphaerodoridium, 289
— beiiguellarum, 289
— capense, 289
Sphaerodorum, 288
— benguellarum, 289
— capense, 289
— claparedii, 289
— gracile, 288
Sphaerosyllis, 275
— capensis, 276
— erinaceus, 277
— hystrix, 275
capensis, 276
— perspicax, 277
— semiverrucosa, 276
— sublaevis, 277
Sphinosphaera. 714
spinosa, Harmothoe, 64
spinosa, Iphione, 37
spinosus, Eupomalus, 809
spinosus, Ophiodromus, 223
spinosus, Orlhwdromus, 223
Sl'INTHERIDAE, I 1 9
spinulosa alcocki, Sabellaria, G72
— eupomatoidcs, Sabellaria, 667
— fucicola, Sabellaria, 672
— gilchrisli. .Sabellaria, 667
— intoshi, Sabellaria, 672
Spio, 480
— bnmbyx, 474
— filicornis, 481
— magnus, 482
— lulgaris, 477
— sp., 481
Spiochaetopterus, 527
— costarum, 528
— typicus, 527
— vitrarius, 528
SI'IONIDAE, 459
Spiophanes, 474
-— bombyx. 474
— kroyeri, 474
— soederstromi, 475
Spirobranchus, 801
— gardineri, 803
— giganteus, 803
— maldivensis, 803
— semperi, 803
— tclraceros, 803
Spirographis, 756
SPIRORBINAE, 792
Spirorbis, 793
— adeonella, 794
— borealis, 793
— borealis, 792
— capensis, 796
— foraminosus, 797
— laevis, 796
— malardi, 794
— patagoniciis, 794
— sp., 792
spissipes, Irmula, 267
splendens, Macrophyllum, 151
splendens, iXotophyllum, i-,i
splendida, Hesione, 228
splendida, JVereis, 292
spongicola, Syllis, 240
squamata. Scolelepis, 483
sc|uaniatLim, Sigalion, 104
sqiiamalus, Lumbricus, 483
scjuamosa, Acholoe, 52
squamosa. Nereis, 52
Slaurocephalus gardineri, 455
— kcfersteini, 452
— negleclus, 455
— rubrmillalus, 457
— rudolphi, 437
Stauronereis, 434
— angolana, 458
— cgena, 454
— rubrovittatus, 450
steenstrupi, Prionospio, 489
Steggoa, 154
— inagalhaensi, 136
steinitzi, Sc>-plioproctus, 604
STERNASPIDAE, 648
Sternaspis, 648
— scutata, 648
ajricana, 648
INDEX OF SCIENTIFIC NAMES
875
Sthenelais, 108
— boa, 109
— dendrolepis, 98
— helenae, 108
— incisa, 1 1 3
— limicola, 1 1 1
— oculata, 97
— papiUosa, 108
— simplex, 1 1 3
Sthenolepis, 1 12
— incisa, 1 13
— japonica, 1 12
— tetragona, 1 13
stimpsonis, J^'ereis, 293
Streblosoma, 723
— abranchiata, 723
— chilensis, 725
— hesslei, 725
— persica, 725
Streblospio, 462
Streptosyllis, 238
striata, Hermodice, 1 3 1
striata, Alastigonereis, 293
striata, Pherecardia, 131
strigiceps, Pomatoceros, 80 1
stroemi, Terebellides, 713
Stylarioides arenosiis, 653
— capensis, 664
— laei'is, 66 1
— moiiroi, 660
— parmatus, 658
— swakopianus, 66 1
— tropicus, 663
— xanthotricha, 66 1
subaenea, Giycera, 363
sublaevis, Sphaerosyllis, 277
succinea. Nereis, 32 1
sudaustraliensis, Arenicola loveni, 610
suecica simplex. Aricidea, 558
— suecica, Aricidea, 557
swakopiana, Pherusa, 661
swakopianus, Polycirrus, 717
swakopianus, Stylarioides, 661
SYLLID.\E, 233
Syllides, 259
— longocirrata, 259
Syllidia, 226
— armata, 227
— capensis, 227
sylliformis, Trichosyllis, 234
SYLLINAE, 236
Syllis, 239
— arnica, 243
— anops, 243
— armillaris, 249
— benguellana, 249
— bouvieri, 246
— brachychaeta, 249
— capensis, 235
— cirropunctata, 250
— closterohranchia, 234
— cornuta, 244
— crassicornis, 234
— djiboutiensis, 235
— exilis, 250, 253
— ferrugina, 244
— Jerruginea, 244
— fulgurans, 260
— fusicornis, 235
— gracilis, 241
— hyalina, 246
— krohni, 234
— lineata, 234
— longissima, 243
— longocirrata, 259
— monilaris, 239
— nigropharyngea, 249
— polycera, 260
— prolifera, 248
— sexoculata, 235
— spongicola, 240
— taprobanensis, 246
— trifalcata, 241
— variegata, 248
— vittata, 252
— zebra, 256
— zonata, 248
— •sA>235
Synelmis albini, 215
syphodonta, Lumbricus, 140
syphonodonta, Eteone, 140
Tachytrypane, 580
— jeffreysi, 580
tagensis, Vanadis, 180
Talehsapia, 214
taprobanensis, Syllis, 246
taprobanensis, Tvposyllis, 246
taylori, Mesochaetopterus, 531
Tecturella luctator, 652
Telepsaius, 527
— costarum, 528
— vitrarius, 528
Telothelepus, 731
— capensis, 731
tenera, Paraonis, 565
tentaculata, Audouinia, 515
tentaculata, Augeneria, 430
876
POL\ CHAEIA Ol- SOU IHERN AFRICA
tentarulata, Cirrifornib, 515
tcntaciilata, Eunice, 3<)i
Untaculntci. Terebella, 513
tenlaculus, Cirratulus, 499
— mfridionalis, Cirratulus^ 499
tenuis, Clymenura, 623
teriuis, Corynocfphalus, 1 93
tenuis, Leiochom\ 623
lenuis, Parheteromasliis, 605
tenuis, Plotohclmis, 193
tenuisetis, Polycirrus, 715
tenuisetosa, Euphiorw, 82
tenuisetosus, Lepidonotus, 82
Terebella, 747
— comata, 729
— elirenbergi, 748
— fasciata, 742
— irifuutlibuluiil. 773
— lapidaria, 747
— lapidaria, 732
— macrobrancliia, 736
— nifdum, 743
— rubulosa, 744
— plagiostoma, -jiq
— pterochacta, 747
— pterochaela, 732
— schmardaei, 748
— teutaculala, 515
— Iriserialis, 730
— venmtula, 735
TEREEF.LI.IDAE, 706
Terebellides, 71 i
— strocnii, 713
TEREBELLINAE, 73 I
TerebelIobran( hia, 7jc>
— natalcnsis, /"/i
terricola, Petaloproctus, 622
tesselata, Glycera, 359
tftraceros, Pomatortros, 803
tetraceros, Spirobranchus, 803
tetragona, Chloraema, 652
telragona, Leanira, 1 13
letragorta, Pherusa, 652
telragona, Sthenolepis, 1 13
tetragonuw, Sigalioti, 1 13
tetraura, Lumbrineris, 439
tctraura, Lumbrineris, 427
tetraurus, Lumbriconereis, 439
tetraurus, Notocirrus, 439
Telreres mtirala gikhrisli, 677
Tharyx, 504
— acutiis, 504
— annulosus, 504
— dorsobranchialis, 50G
— filibranchia, 505
— marioni, 505
— multifilis, 499
Thalenessa, 107
— dendroU'i^is, 107
— oculata, 107
THELEPINAE, 722
Thelepus, 727
— cincinnatus, 727
— comatiis, 729
— pequenianus, 730
— plagiostoma, 729
— setosus, 729
africnnus, 722
— triserialis, 730
— sp. 722
Theodisca hexaphyllum, 533
Theostoma oenctedi, 533
Thorrnora, 79, 80
— jukesii, 80
Timarete, 501
— ■f/'-S'S
TOMOPTERIDAE, I 96
Tomopteris, 197
— apsteini, 199
— carpenteri, 204
— cavalli, 206
— cavallii, 204
— diiccii, 2ni
— dunckeri, 201
— elegans, 205
— euchaeta, 202
— helgolandica, 199
— kempi, 202
— kerfersteinii, 205
— krampi. 2f>4
— ligulata, 204
— nationalis, 198
— nisseni, 202
— onisciforniis, 197
— pacifica, 199
— planktonis. 206
— septentrionalis, 205
tongatabucnsis, J^ereii, 293
tongatabuensis, Platynereis, 293
torclli, Potamilla, 7<.)6
torelli, Sabcllii, 766
torquata, Eunice. 389
Torrea, 186
— Candida, 188
torulosa, Spermosyliis, 270
Travisia, 575
— concinna, 576
— forbesii, 575
INDEX OF SCIENTIFIC NAMES
877
Travisiopsis, 210
— capensis, 207
— coniceps, 210
— dubia, 212
— lanceolata, 212
— levenseni, 213
■ — lobifera, 213
— sp., 207
tribuUata, Hauchiella, 714
Iribullala, Polycinus, 7 1 4
trichobkanchidae. 710
trichobranchidea, 710
trichobrjvnchinae, 710
Trichobranchus, 71 1
— glacialis, 71 1
Trichosyllis sylliformis, 234
Tricoelia varieopedalus, 529
tridactyla, Glycera, 352
tridentata, Nerindes, 484
trifalcata, Syllis, 241
trifasciata, Nereis, 312
trilineata, Eulalia, 152
trilobala, Amaea, 718
trilobata, .^maeana, 718
Irilobala, Polycinus, 7 1 8
triguetra, Serpiila, 801
Iriserialis, Terebella, 730
triserialis, Thelepus, 730
trissochaetus, Lepidonotiis, 37
trockiscophora, Polynoe, 85
Trochochaeta, 519
TROCHOCH.\ETIDAE, 5I9
tropica, Pherusa, 663
tropicus, Stylarioides, 663
Trophonia capenxis, 664
— glaiica, 666
— xanthotricha, 66 1
Trypanosyllis, 253
— ankyloseta, 255
— gemmipara, 256
— gemmulilera, 256
— ingens, 235
— krohni, 253
— misakiensis, 256
— prampramensis, 255
— zebra, 256
tuberculata, Cirrosyllis, 28 1
tubercuiata, Hemilepidia, 38
tuberculatus, Autolytus, 281
tubicola, Hyalinoecia, 41 1
tuhicola, jXereis, 41 1
tubicola, Onuphis, 405
tubicola, Phyllodoce, 148
tubifex, Eunice, 386
tubifex, Eupanthalis, 93
tubularia anomala, Protula, 820
— capensis, Protula, 816
— tubularia, Protula, 820
tubularia, Serpula, 820
tulearensis, Nephtys, 345
Tylonereis, 297
typha, Pista, 737
TYPHLOSCOLECIDAE, 2O7
Typhloscolex, 208
— muelleri, 208
— phyllodes, 209
typica, Neanthes latipalpa, 292
typica, Scolaricia, 547
typicus, Spiochaetopterus, 527
Typosyllis, 246-252
— taprobaneusis, 246
ubianensis, Lysilla, 72 1
Unanereis, 298
— macgregori, 299
uncinata, Dispio, 482
uncinata, Hydroides, 805
uncinata multispinosa, Hydroides, 798
uncinata, Serpula, 807
uncinatus, Eupomatus, 805
uncinatus, Lopadorhynchus, 159
uncinatus macronyx, Hydroides, 798
unibranchia, Pista, 737
unicornis, Glycera, 362
unicornis, Lumbriconereis, 403
unicornis, Nematonereis, 403
unifasciata. Nereis, 318
unifasciata. Nereis, 312
uniformis, Phalacrophorus, 171
uniramus, Scoloplos, 552
vallata, Nereis, 334
vallata, Perinereis, 292
vallata, Perinereis nuntia, 334
Vanadis, 180
— antarctica, 181
— crystallina crystallina, 182
inomata, 1 82
— formosa, 1 8 1
— fuscapunctata, 184
— longissima, 186
— minuta, 184
— minuta, 1 84
— tagensis, 180
— violacea, 185
vancaurica. Nereis, 334
vancaurica, Perinereis, 334
variabilis, Notopygos, 130
87«
POLVCHAKI A OF SOUTHERN AFRICA
fdrifgala, ^'ereitepas^ 33 1
lariegnla. Nereis, 331
tariegala, Perinrreis, 2cj3
variepata, Pseudonereis, 331
varlesata, Syllis, 248
varieopedatus, Chaeloplenis, -,21)
larieopedatus, Tricoelia, 529
venustiila africana, Nicolea, 736
— vcnustula, Nicolea, 735
lenuilula, Terebella, 735
vermicularis cchinata, Serpula, 810
— vermicularis, Serpula, 8og
\'ermiliopsis, 812
— acanthnphora, 814
— babylonia, 8 1 3
— glaiuligerus, 813
— pygidialis. 813
— pygidialis, 813
verrucosa, Pilargis, 218
verruculosa, Hcrmenia, 77
vcruger, Paeiiophvlax, 272
verugera, Exogone, 272
vesiculosa, Amphilrile, 758
vesiculosum, Branchiomma, 758
vesiculosum, Megalomma, 758
villosa, Brada, 652
villosa capensis, Brada, 656
lillosum, SiphoiiDsloma, 656
vincenti, Branchiomaldane, 608
violacea, Brancliiomma, 768
violacea, Dasychone. 768
violacea capemis, Dasychone, 751
violacea, Dyslilia, 75 1
violacea, Sabella, 768
violacea, Vanadis, 185
viridis, Diopalra, 405
viridis, Eutalia, 136
viridis capensis, Eutalia, 1 54
viridis. Nereis, 151
vitrarius, Spiochaetopterus, 528
vilrarius, Telepsavus, 528
vitlata, Eunice, 385
villala. Nereis, 385
viltata, Syllis, 252
volulacornis, Bispira, 753
vutgarii, S/iio, 477
waahli, Antitioe, 72
waalili, Harmothoc, 72
wahlbergi, Lcpidonutu^, 85
Watelio, 175
whiteavesi, Malnigrenia, 48
whydahaensis, Isolda, 692
willcyi, Nereis, 320
williami, Nerindes, 485
wollebaeki, Lanice, 743
wyvillei, Laeiinornce, 3 1
u-yvillei, Laetmonice producta, 33
xaiiltiolricha. Flabelligera, 652
xanlholticha. Slylarioida, 661
xaiilhnlncha, Trnphonia, 661
zebra, Syllis, 256
zebra, Trypanosyllis, 256
zetlandica, C^aullierclla, 509
zellandica, Chaetozone, 509
zetlandica, Helerocirrus, 509
zonata pjersica. Nereis, 314
zonata, Sylln, 248
zululandica, Dendronercidcs, 302
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