rT

J. H. DAY

A REVIEW OF THE FAMILY
AMPHARETIDAE (POLYCHAETA)

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A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA)

By
J. H. Day
Koology Department, University of Cape Town

CONTENTS

PAGE

Introduction AM Ass NG oan Cy)
Discussion of diagnostic characters LOS
Division into sub-families and genera .. 103
Sub-family Melinninae .. ae LOO
Table of type-species . . a LOA!
Key to genera ae oe BH 1 LOS)
Generic definitions .. ae ee LOO
Sub-family Ampharetinae 5 56!) UO
Table of type-species . . oe LOS)
Key to genera sia ob ceselEh
Generic definitions .. Bs ol), WES)
Summary .. Si Me a So MLC)
References .. 36 56 a6 6. IHG)

INTRODUCTION

The most complete account of the family Ampharetidae is that of Hessle
(1917). He reviews earlier work including that of Malmgren (1865) and Fauvel
(1897) on family and generic characters, Nilsson (1912) on the nervous system,
Wiren (1885) on the alimentary canal, and Meyers (1887) on the nephridia.
After giving further evidence from his own researches on the nephridia and
structure of the stomach he goes on to discuss generic characters. While he
follows the main lines laid down by Malmgren he finds that the genera were so
narrowly defined by the latter worker that they seldom include more than one
or two species, so that broader generic definitions are necessary. He criticizes
Fauvel’s work on the enumeration of the anterior segments and his genera as
being based partly on internal anatomical characters. Surprisingly enough
Hessle’s own generic divisions may be criticized on the same lines. Although
he stresses that the number of nephridia may vary from species to species in
the same genus he has grouped the genera largely on the relative sizes and
arrangement of these internal structures.

Since the publication of Hessle’s work many new genera have been erected,
particularly by Augener (1918), Chamberlin (1919), Benham (1921),
Annenkova (1930), Caullery (1944) and Eliason (1955). In general these later
workers focused attention on external features, particularly the development

97
Ann. S. Afr. Mus. 48 (4), 1964, 97-120. A ty RY 2 1064

98 ANNALS OF THE SOUTH AFRICAN MUSEUM

of the paleae and the number of thoracic setigers, and they do not describe the
nature of the nephridia nor the morphology of the stomach. As a result it is
difficult to correlate the genera they describe with those defined in Hessle’s
monograph. There is also another source of confusion. Nilsson and Hessle
recognize only a single two-ringed buccal segment in front of the paleal or
first branchiferous segment, so that the latter becomes segment II. This inter-
pretation is followed by Annenkova (1930), Eliason (1955), and Uschakov
(1957). On the other hand Malmgren (1865), Fauvel (1897 and 1927) and
Caullery (1944) recognize two segments in front of the paleae, so that the latter
which also bears the first pair of branchiae becomes segment III. At first sight
it would seem of little taxonomic importance whether the first branchiferous
segment is labelled segment II or III, but as many workers do not state which
system of numbering they are using, statements such as ‘setae present on seg-
ment III’ or ‘nephridia absent from segment V’ or ‘notosetae of segment XIII
modified’ lead to confusion.

To date about 150 species have been described, distributed through about
49 genera. Fifty-six species are ascribed to the three genera Amphicteis, Amage,
and Ampharete, and the remaining 94 are distributed through the remaining
46 genera, many of which are monotypic. The need for revision is obvious.

In the account that follows the various diagnostic characters are discussed,
the grouping of genera is considered, and tables and keys are provided for the
whole family.

I wish to thank Dr. Olga Hartman for constructive criticism during the
preparation of this paper, and both Mr. R. Sims of the British Museum and
Mr. J. B. Kirkegaard of the Copenhagen Museum for allowing me to examine
material housed in their respective institutions. The Trustees of the South
Afritan Museum are grateful to the South African Council for Scientific and
Industrial Research and the University of Cape Town for grantis in aid of
publication.

DIscUSSION OF DIAGNOSTIC CHARACTERS

The head and buccal tentacles. The Ampharetidae are deposit feeders and
the food particles are collected by mobile projections that arise from the roof
of the buccal cavity. The most common type are buccal tentacles which are
either smooth with a longitudinal groove on one side as in the Terebellidae or
papillose; often these papillae are pinnately arranged, but not always, and
buccal tentacles of this type are best described under the more general term
‘papillose’. Apart from tentacles, folded or probosciform feeding membranes
have been described in Amythas membranifera Benham and Pabits deroderus
Chamberlin. An examination of Jsolda whydahensis Augener showed a long
ribbon-shaped structure protruding from the mouth with the distal end split
into a number of short tentacles.

As stated, these feeding organs arise from the roof of the buccal cavity and
in some cases the mouth cavity leads straight back into the pharynx; in other
cases, however, an internal shelf or horizontal septum separates the tentacular

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 99

cavity above from the pharynx below. According to Hessle this septum
corresponds to the dorsal lip of the Terebellidae.

Above the mouth there is a hood-shaped lobe which Hessle refers to as the
‘Tentakelmembran’ and Fauvel as the prostomium. Since it bears the eyes and
nuchal grooves and contains the cerebral ganglia the latter term will be used
here. In some genera it also bears a pair of glandular ridges which diverge
anteriorly and project from the antero-lateral margins as a pair of blunt
processes.

When the buccal tentacles are retracted the prostomium may appear
bluntly pointed with obvious lateral grooves, so that the whole prostomium
(or tentacular membrane) has been described as trilobed. When the buccal
tentacles are fully extended, however, the whole head is stretched, the anterior
margin is straightened, and the lateral grooves disappear. This mobility of the
head means that the exact shape of the prostomium is of little systematic value.
The glandular ridges, however, are always recognizable and the eye-spots can
usually be found.

The segmentation of the head region. As described earlier (Day 1961) the region
behind the prostomium is telescoped and the segments are distorted. ‘The
branchiae are often grouped on a transverse branchial ridge which partially
overhangs the segments immediately behind the prostomium. The ventral
surface in Melinna and related genera is distorted by the formation of a pair of
lateral folds which slope back from the mouth region towards the dorsum of
segment VI where they unite to form a transverse dorsal crest. On the basis of
dissection and examination of external features I have accepted the interpreta-
tion of Malmgren, Fauvel, and others that there are two achaetous segments
preceding the branchiae in all genera. The first branchiferous segment, which
bears the paleae in some genera, thus becomes segment III.

The branchiferous region (segments III-VI). In the more primitive genera,
including Phyllocomus, Melinna, Isolda, Amphicteis and. Ampharete, there are four
pairs of gills. Dissection of the blood-vessels of Amphicteis gunnert, described by
Day (1961), showed that the four gills are supplied by four blood-vessels
corresponding to segments III, IV, V and VI. With the telescoping of the
anterior segments, however, there is a tendency for the four pairs of gills to
be grouped on the dorsal surface of segments III and IV. In Phyllocomus the
first two pairs are on segment III and the last two pairs are segmentally
arranged. The same arrangement has been reported in Amage auriculata and
Amphicteis posterobranchiata. In other genera the arrangement may be 2:: 2 or
3: 1 where three pairs of gills are arranged in a transverse row across segment
III with the fourth pair immediately behind.

In many genera the primitive number of four pairs of gills has been
reduced to three and in Auchenoplax Eblers and its synonym Melinnoides Benham
only two pairs have been reported.

The gills may be separate from one another or united by a basal web.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Commonly this web unites only the first three pairs of gills, leaving the fourth
posterior pair free. The gills may be smoothly cylindrical in shape or may
develop lateral papillae or flanges or even a series of lateral lamellae. Commonly
the branchial papillae are arranged in two lateral rows, and such branchiae
have been termed bipinnate, but this is not always the case. In Pterampharete
luderitzi, for example, numerous irregularly arranged papillae arise from the
anterior surface of the branchial axis. In generic descriptions such gills are
best described by the broader term papillose. Similarly the lateral lameliae of
such forms as Phyllamphicteis collaribranchis Augener and Phyllocomus (olim
Schistocomus) hiltont (Chamberlin) may be arranged in one or more rows.

The telescoping of the branchiferous region often results in the fusion of
segments and the loss of setae. In Jsolda and Melinna of the sub-family Melznninae
the neurosetae of the four segments III-VI are present though those of segment
VI may be lacking in some species. Segment III never has notosetae, and
segment IV has notosetae in the form of stout hooks behind the gills, but the
notosetae of segments V and VI are either small or absent. In Melinnopsis
McIntosh the hooks of segment IV are lacking, but notopodial capillaries are
present on segments V and VI.

In the sub-family Ampharetinae neurosetae are never developed on segments
III to VI. In a few cases all four segments may be distinct, but usually segments
III and IV are fused and occasionally segments III, IV and V. The notosetae
are variously developed. In primitive forms such as Amphicteis the notosetae of
segment III are enlarged to form stout paleae and those of segments IV—VI are
all present as normal capillaries. Often, however, the capillary notosetae of IV
and V are small or absent. Similarly the notosetae of segment III] may remain
small and not form paleae, or may be entirely lacking. In Sabellides, for example,
the notosetae of segment III are either absent or represented by small capillaries,
segment IV is fused to segment III and lacks setae, and the first normal capil-
laries are on segment V. In Neosabellides elongatus there are no setae on segments
III and IV, and in Paramage madurensis Caullery reports that the first bundle of
notosetae appears on segment VI though earlier achaetous notopodia remain.
In view of the many variations in the setation of the branchiferous segments
III to VI and the difficulty of deciding whether small setae present on the fused
third and fourth segments represent small paleae of segment III or small
capillaries of segment IV, it would seem undesirable to base generic divisions
on the presence or absence of paleae as has been done in the past.

Internal characters: diaphragm, nephridia, stomach. Apart from a few minor
details no new researches are reported here under this heading. The information
concerning the type-species of each genus has been extracted from Hessle
(1917), supplemented by additional information from Annenkova (1930), and
has been summarized in tables I and II. In several genera no information is
available concerning the internal anatomy.

The coelomic cavity of the first few segments is separated from the rest of
the thorax by a relatively stout septum or diaphragm. According to Hessle this

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) IOI

always lies between segments IV and V, but according to the enumeration of
segments adopted here it lies between segments V and VI. Following Hessle’s
terminology the nephridium in segment V (Hessle’s segment IV) is termed the
anterior nephridium, and all later ones are posterior nephridia.

Never more than five pairs of nephridia have been reported; in some cases
there are only three and in exceptional cases only two. At first sight the number
of nephridia would appear to provide a reasonable basis for classification, but
as Hessle himself has pointed out, their distribution is not constant. In some cases
four pairs may be present in segments V to VIII, in others the nephridium in VI
is missing and the four pairs are in V, VIT, VIII and [X; again V may be missing
and the four pairs are in segments VI-IX. The same variability occurs in those
species in which there are only three pairs of nephridia. They are located in
segments V, VI and VII in Sosanopsis wireni, Parhypania brevispina and Hypania
invalida, and in segments V, VII and VIII in Sabellides octocirrata. Even when the
number is reduced to two pairs there are differences in arrangement. In
Phyllocomus crocea and Schistocomus (=Phyllocomus) hiltont the nephridia are in
segments VII and VIII, and in Lysippides fragilis they are in segments V and
VII.

Hessle further reports that the number of pairs of nephridia is not constant
within the single genus Ampharete but may vary from two pairs situated in
segments V and VII for A. acutifrons and A. arctica to four pairs situated in
segments V, VII, VIII and IX for A. lindstromi and A. kerguelensis. The num-
bering of the segments given here is of course one higher than that given by
Hessle.

In view of this variability of the nephridia and the labour involved it would
seem undesirable to base generic divisions on these structures as has been done
by Hessle. None the less the nephridial papillae of the anterior nephridia in
segment V are occasionally visible. In Sabellides and Pterampharete the ducts are
elongated dorsally and two nephridial papillae may be found on the branchial
ridge between the two groups of gills. In Anobothrus, Sosane and Asabellides the
ducts open on a single nephridial papilla in the same position. These characters,
while not in themselves sufficient for generic distinction, provide useful
confirmatory evidence.

The alimentary canal is normally a fairly straight tube divided into
pharynx, oesophagus, stomach and intestine. At the anterior end of the stomach
where it joins the oesophagus a pair of lateral pouches project forward in
Ampharete, Microsamytha and Anobothrus, but not in Melinna nor fourteen other
genera which have been investigated by Wiren (1885), Hessle (1917) or
Annenkova (1930). There is also a curious invagination of the ventral wall of
the stomach of some genera, which is referred to by Hessle as an ‘innere Blind-
sack’ and is here termed an internal diverticulum. It has been found in Amage,
Amphicteis, Amphisamytha and Hypania, but not in fourteen other genera which
have been examined. Since such characters demand dissection they are of
doubtful value in practical classification.

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

The posterior thorax. As stated earlier, the first branchiferous (or paleal)
segment is reckoned here as segment III and the branchiferous region as
extending over segments III to VI. The beginning of the posterior thorax on
segment VII is marked by the first appearance of uncini in the neuropodia in
all genera. It is the most important and most easily recognizable landmark on
the body. Segments anterior to it may be distorted or fused and individual
segments may lack notosetae or nephridia, but segment VII and the posterior
thoracic segments are always well defined and fully developed. The maximum
number is 14 and the minimum number is 11 in all genera except Mugga,
where there are 9. It is suggested therefore that the numbers of such uncigerous
thoracic segments provide better generic characters than the total number of
thoracic setigers which have led to such confusion.

The notopodia of the posterior thorax are conical projections which in
Amphicteis and several other genera bear a terminal papilla or ‘notopodial
cirrus’. The neuropodia are short projecting pinnules and in a few genera such
as Phyllocomus they bear a superior papilla or cirrus above the row of uncini.

The notosetae are winged capillaries which are very uniform in shape
with few exceptions. In Anobothrus the 8th uncigerous segment has its notopo-
dium elevated and the tips of the notosetae are minutely spinulose. In Mugga
the same phenomenon occurs on uncigerous segment 9, and in Sosane and
Sosanopsis it occurs on uncigerous segment 12.

The uncini are flattened tooth plates of various shapes but are all roughly
triangular to quadrangular, with one or more series of teeth above the base.
In the Melinninae there is always a single series of teeth, but in the Ampharetinae
the teeth of the thoracic uncini may be arranged in 1, 2 or as many as 5 vertical
rows. Abdominal uncini often have more rows of teeth. The base of the uncinus
may be long and well separated from the tooth rows, or it may be short and
curve up towards the lowest tooth to form a bluntly rounded prow. Hessle and
others have quoted such differences as generic distinctions, but to me they
appear to be no more than specific characters, for the shapes vary very consider-
ably. The number of tooth rows seems to be more important when there are
only one or two vertical series, but when there are three or more they lose their
value.

The abdomen. There is seldom any abrupt narrowing between thorax and
abdomen, merely a change in the parapodia. This, however, is well marked.
The notosetae disappear and the notopodia are either reduced or absent. In
Melinna, Amphicteis, Amage, Phyllocomus and many other genera, rudimentary
notopodia persist on abdominal segments, and this is regarded as the more
primitive condition. In Ampharete, Sabellides and others, notopodia are entirely
lacking. Where notopodial! cirri are present on the thorax they tend to persist on
the rudimentary notopodia of the abdomen, as may be seen in Amphicteis and
Amage. Similarly, where neuropodial cirri are present on the thorax these also
are continued on the abdomen. Actually the uncigerous pinnules very often
have an obscure superior papilla and this tends to become better marked towards

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 103

the end of the abdomen. This is particularly well shown by Sabellides octocirrata
where the superior papillae become long cirriform projections.

The whole abdomen consists of a variable number of segments. In the
Melinninae there is always a large number of segments, the extremes being a
minimum of 20 in Melinnexis antarctica and a maximum of go in Irana hetero-
branchia. In a few primitive genera of the Ampharetinae there are also large
numbers of abdominal segments. Thus Phyllocomus crocea has 45 but in the great
majority of genera the number is much less, 12-18 being the usual range. It
is doubtful whether the exact number is constant for a species, and certainly
it is not of generic importance.

The abdominal uncini are more or less similar to those of the thorax but
usually there are more teeth arranged in more vertical rows or transverse arcs.

The pygidium is terminal and may be encircled by a number of low
indistinct papillae or may bear a reduced number of longer anal cirri.

DIVvISsION INTO SUB-FAMILIES AND GENERA

Hessle (1917) does not divide the Ampharetidae into sub-families, although
his remarks on p. go show that he is inclined to separate Melinna and Isolda
from other genera in this way. Chamberlin (1919) proposed three sub-families,
namely the Melinninae with dorsal hooks behind the gills but no paleae, the
Ampharetinae with paleae but no hooks, and the Samythinae with neither paleae
nor hooks.

Since the publication of Chamberlin’s monograph many new genera
have been described. The diagnostic characters of the various type-species are
set out in tables I and II, and it will be immediately obvious that there are
several genera related to Melinna. Not all of these possess dorsal hooks behind
the gills, but they all possess fine acicular neurosetae in segments III, IV, V
and often VI. I agree with Hessle in regarding this as an important and primi-
tive feature which links the Amphaetidae to Terebellides and other genera of the
Trichobranchidae and through them to the Terebellidae. I have therefore enlarged
Chamberlin’s diagnosis of the Melinninae to include all Ampharetidae with
acicular neurosetae in segments III-VI whether they possess dorsal hooks or
not. The amended characters of the sub-family are given later.

Chamberlin’s sub-families Ampharetinae and Samythinae are distinguished
by the presence or absence of paleae. The variability of these setae has been
described earlier; as the accompanying tables show, the setation of the whole
branchial region is not sufficiently reliable for the distinction between genera,
let alone sub-families. Several other possible groupings of genera have been
tried on the basis of the number of branchiae, the number of uncigerous thoracic
segments and the number of tooth rows on the thoracic uncini. None of them
were satisfactory in that they distinguished groups of genera with many charac-
ters in common. It was concluded that all the Ampharetidae apart from the
Melinninae must be included within the sub-family Ampharetinae and that
Chamberlin’s sub-family Samythinae must be sunk.

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ANNALS OF THE SOUTH AFRICAN MUSEUM 105

The reliability of the various characters which may be used to distinguish
genera has been discussed earlier. To be of practical value in a taxonomic key
such characters must not only be constant within a group of species which are
obviously similar in other ways, but they must also be easily seen when sorting a
sample. It is suggested that among such characters are the number of gills, and
the number of uncigerous thoracic segments, the presence of glandular ridges
on the prostomium and the possession of notopodial cirri. These have therefore
been used as the main basis for grouping genera with the addition of several
other features in specialized cases. It is tempting to use the obvious ornamenta-
tion of the gills as well as their number, but after careful consideration this
was omitted for it would have increased the number of monotypic genera
without breaking up the three large genera Amphicteis, Amage and Ampharete. In
any case the gills are often missing so that their ornamentation cannot be
decided, and only the scars remain to determine their number.

As stated earlier, 49 genera are currently recognized as valid. In the
accompanying key these have been reduced to 33 of which 6 are assigned to the
Melinninae and 27 to the Ampharetinae. This means that 16 genera have been
sunk, and reference to the tables will show that 11 of these were monotypic and
the other 5 contained 2 species each. One new genus has been erected in the
sub-family Melinninae.

It is hoped that this revision will make the recognition of genera much
simpler, but inevitably it will be found that many species have been assigned
to the wrong genus. This is certainly true of two species assigned to Lysippe by
myself and one referred with hesitation to Neosabellides.

Key to sub-families
1. Segments III-V (or III-VI) with fine acicular neurosetae. No paleae.
Post-branchial hooks sometimes present .. ai .. Melinninae

Segments III-VI without neurosetae. No a erates hooks. Paleae
sometimes present .. A a6 a’ 56 a6 .. Ampharetinae

106 ANNALS OF THE SOUTH AFRICAN MUSEUM

Sus-FAMILY Melinninae Chamberlin 1919 (characters amended)

Buccal tentacles never pennate or 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, IV and often V and VI. Uncini are from segment VII and always have a
single series of teeth. Numerous (20-90) abdominal segments.

Key to genera of Melinninae

1. Stout notopodial hooks behind the gills .. Sa ae ee sic 2
No notopodial hooks ae a, aie 5
2. Notopodial hooks on two eueacnk Prostoraittin ‘Alongared a .. Moyanus
(doubtful)
Notopodial hooks on segment IV only. Prostomium short... Sa 3
os wh pairs of gills 56 Sc ate 4
3 pairs of gills, including both sinoain intl mailioce fone D0 .. Irana
2 pairs of papillose gills .. s'6 se ae we Sa .. Ocorpata
(doubtful)
4. Allgillssmooth .. ih be As xe .. Melinna
Some gills smooth, some pemillocel a = tte eee .. Isolda
5. 4 pairs of smooth gills Bo ae ee ae ae 5 .. Melinnopsis
3 pairs of smooth gills We 53 Pe ae bc 4 .. Melinnopsides

MOYANUs Chamberlin 1919

Prostomium elongated and probosciform. Buccal tentacles mounted on
long tongue-shaped projection. Four pairs of smooth gills. A dorsal crest on
segment VI. Segments III—VI with notopodial hooks on segments ITV and V
and fine acicular neurosetae on segments III, IV and V. Twelve uncigerous
thoracic segments and about 65 abdominal ones. Thoracic uncini with a-single
series of teeth. Type-species Moyanus explorans Chamberlin 1919.

MELINNA Malmgren 1866

Buccal tentacles smooth with a groove on 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
neurosetae on segments ITI—V and sometimes on VI as well. Fourteen uncigerous
thoracic segments and 30-50 abdominal ones. Thoracic uncini with a single
peues oF teeth, Type-species Sabellides cristata Sars 1851.

IsoLDA Miiller 1858

Buccal tentacles smooth with a groove on one side. Four pairs of gills of
which 2 are smooth and 2 are papillose. A dorsal crest across segment VI.
Segments ITI-VI with notopodial hooks on segment IV and notopodial
capillaries on segments V and VI. Fine acicular neurosetae on segments III-V
and sometimes on VI as well. Twelve to 13 uncigerous thoracic segments and
25-36 abdominal ones. Thoracic uncini with a single series of teeth.

Type-species Isolda pulchella Miller 1858.

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 107

IRANA Wesenberg-Lund 1949

Buccal tentacles smooth. Three pairs of gills of which 1 is smooth and 2 are
papillose. A dorsal crest across segment VI. Segments III-IV with notopodial
hooks on segment IV and fine acicular neurosetae on segments III-VI. Twelve
uncigerous thoracic segments and about 90 abdominal ones. Thoracic uncini
with a single series of teeth.

Type-species Irana heterobranchia Wesenberg-Lund 1949.

OEORPATA Kinberg 1867

An incompletely described genus with two pairs of pennate gills; noto-
podial hooks on segment IV; many abdominal segments; uncini with a

pete cics.o: teeth: Type species Ocorpata armata Kinberg 1867.

MELINNOPSIS McIntosh 1885
Synonyms Melinnexis Annenkova 1930 and Melinnides Wesenberg-Lund. 1950.

Buccal tentacles smooth with a groove along one side. Four pairs of smooth
gills. Dorsal crest on segment VI present or absent. Segments III-VI without
notopodial hooks on segment IV but with notopodial capillaries on segments V
and VI. Fine acicular neurosetae on segments III-V and sometimes on VI as
well. Ten to 14 uncigerous thoracic segments and 25-50 abdominal ones.
Thoracic uncini with a single series of teeth.

Type-species Melinnopsis atlantica McIntosh 1885.

MELINNOPSIDES gen. nov.

Buccal tentacles smooth with a groove along one side. Three pairs of
smooth gills. No dorsal crest on segment VI. Segments III-VI without noto-
podial hooks on segment IV but with notopodial capillaries on segments V and
VI. Fine acicular neurosetae on segments III-V. Ten uncigerous thoracic
segments and about 30 abdominal ones. Thoracic uncini with a single series of
teeth.

Type-species Melinnopsis capensis Day 1955.

SUB-FAMILY Ampharetinae Chamberlin 1919 (characters amended)
(including Samythinae Chamberlin 1919)

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 to VI. Neuropodial uncini start on segment VII
and may have one or more series of teeth. Few (8) to many (60) abdominal
segments.

ANNALS OF THE SOUTH AFRICAN MUSEUM

108

(posuepy a

(3161 youqoayjo\A

IIA 8 A ‘yjoours &) sysolf sranyqupy ad44)
98 ou ou ou ur‘sida ‘sid? & ti OY 9-9 @¢G Yoows ou L161 a[ssopy saps]
(aye]JourRy o (g161 ‘sny
“yjoouus @) siygun.igi.wvjjo9 *g addy)
‘TID eue aed 1 GI d é sof é ‘idbo1r Vi Qj 9 9 gq Yoows ou gr6r s9UNSNY sraznydGunjjdyg
(6161
“siago1yg (qyQoouIS []z) ‘ueyy issdqo “gq 2dAq)
“uD dyg Jo wuikuouKks § 0G é 8 sah é ‘sad? 63 FI 9 9) 9 ¢ Woows ou 6161 urproquiey'y vag
(Sggr
(q}ooUuIS []z) “POW vIyILDJUD *) 9dAj)
‘1110 eue ared 1 Gz é A é é ‘udboG %t1 959) 29 -— yioous sok Sger ysowupopy vpjauviqnixy
(0981 “1D
IIA-A_ (qnoous T[e) ppyvaur srajnydupy ody)
of&-G1 soAM ou ou ur‘sid & ‘adbo o¢ 61 9 9 29 g wqoous sok L6g1 AnoowoIsCQ vunddzT
(Oggt “ay snus
IIA-A_ (yjoours T]2) -1naig = siajpnryquyy ad Ay)
: g[qooy Os-gI OU ou ou urT'sid & ‘sidvb 31 Vi OF DD ad Moows sok gzbr veaoyusuuy viundcy.wg
(9981
IIIA-A_ (y}00uISs [[R) wRYy YyV]NILUND “Py IdAq)
‘Tad Teue sired 3% 8 ss ou sah ut‘sid ¥ ‘sIdsy c= fl) 2 9 — UOOUIS = asaK QOgt usIsURYy app
(8481 “1D
(yjoours []2) QSO] pyofiysnanv saprpjaqns ad Ay)
‘Tda10 yeue aed 1 GI é & so é ‘sad bP «1 Fr yj 9 9 — -fided: sok Phb1 AtoT[NVeD srajoyqupiwg
(S&gI saeg
XI-A _(yjoours T]2) aauuns § apapydupy adj)
‘qa19 yeue sed 1 Gi é é sof url ‘sid G sid? I +I OD DD d wYyloows sox IGQI aqnay siajnyqupy
oud ct n [o}
Z > oe 8 f Z A Sel oS Us An ee
Ss Se Aw S sel ere 3 Q, O85
Be ee ed . Bee oe ASE 3
SYIVUIOY 5 5 (® am ae Qu = evipouvig © Gy GR s]USUIS9S sopor}uo} s sy satoads-adA}
ee ee E See y 5 5 & jouonejag yeong 8 3 pue snuay
Se Ae eG: Be iS ee
= + gS E58 > B 09
ge 4 a ae ae g B
ue) oO =} $s)
fo} 5 _ fe)
ee 5 o
5 5

‘yuasqe JO yUasard aq Aut avjas a}edIpUT sIsoyyUared UT s19}30[— (F) 10 (9)

‘TI Fav L

‘[]] JuowSas uo ovoyed [ews 10 o81e] = J Jo g ‘sorreyideo [erpodojou [[eus 10 as1e] = 9 10 H

AVNILAUVHINV ATINVAANS AO SAIOUdS-"AdAL CNV VYENHOD AO SYALOVUAVHO

109

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA)

“aUuDSOS
jo wituouds ¢
*poyipour g yUOUISIS
o19vI0Y} — sNOIASTO
-un JO 9¥}9S80}0N
‘ovTyouRIq Us9M0q
eyided jerprrydou 1
peo o pour
yUoTUSIS d1I9B.IOY
St] JO 9¥}9S0}0NJ
‘ovIyouRIq UsI9M19q
eyided jerpraydou 1

“snuloz0j] Cy
jo wtAuouAs ¢

‘Tmo «-Jeue sued &

‘oeqqided

qeue jo 9[1I0 VW

Bee
-jaqusy jo wikuouds ¢

‘T1119 Teue sted 1

‘Ida yeue sed 1

caddisd] yo urAuousds ¢

‘1a19 [Rue aed 1

‘sopud
-fisty jo wthuouks ¢
‘sapug
-gisty jo viAuouks

é

sok

[Tews

sok

sod

ou

sok

ou

sod

sok

Oe

sok

ou

ou

sak

ou

ou

ou

ou

ou

ou

sod

ou

ou

ou

ou

ou

ou

ou

XI ‘IIIA
‘IIA ‘A (qioouts T[e)
ut ‘sid 7 ‘sid F
XI
‘IIIA
‘IIA ‘A (q}OoUuES [[2)
ut ‘sid ‘sid

IIIA = (a7e]Jauuey &
BIIA ‘qoours 1)

ut ‘sid & ‘sid 7
IIIA

2 IIA (pesuey)

ut ‘sid & ‘sid

(yjoours [][e)
é ‘sid 7

(qjoows qe)
é ‘sid

IIA 22 A (qioours [[e)
ur ‘sid & ‘sid

(osoyided 1
‘yjoous &)
é ‘sad

IIIA-A_ (qi00urs ][)
ut ‘sid 7 ‘sid

IIA-A_ (qoours Tye)
ur ‘sad & ‘sad

TITA~A (tnoours qe)
ut ‘sad ‘sad ‘b

a

nN

a

Co.

oI

GI

oI

SI

€1

Vi

br

Ay

qjoours

yoours

qyjoours

qoours

asoypided

asoypided

asoyided

yloouts

qyioows

yjoours

qoows

ou

ou

ou

ou

ou

ou

ou

ou

ou

ou

ou

(9981 “WIR
syinas §=ayaapyduyy adj)
QQ UasUIADT snsyjoqoupy

(9981
“we Bypajns +6) adA})
QOQI UdISsWRP auvsog

(6161
ueYy 2uoqj17y = *6' 9dA})

6161
ulploquieyy ——- smauo20js2y96)

(gL4Q1 “15 v99019 “gq adAq)
gle aqnax snwos0p yg

(SP61

"A 2B “a Syyv40j74] “q 9049)

P61 AapayIog

2%» AapoyIog sapzppaqvsopnasg

(Sggr WoT

vowngis sapyyaqug dq)
6z61 vaoyuouUY sapijaquspy

(0981

“Igy suosfignon “Fr 3dAq)
QggI uUsIswpepyL aauwydup

(gr61

‘smy vipuuadig *q dq)
gi61 szouasny addistjo.g

(9981

Wye, Vipin) “7 adAq)
ggg uosswuypeyy adie T

(4Zgr wun 2zys

-maypmoy siajnyqupy addy)
gs61 vaoyusuuy vpjoupddpy

(4161

Q[ssopy varuodvl *zr addy)
LiGI o]ssopy vyjdiuvsrydiupy

ANNALS OF THE SOUTH AFRICAN MUSEUM

IIo

“UMOUW]

(oz61

jou propyl ‘vyzdiuvs (qj0ouIs |[z) ysr0Fy tuliuos “fy ody)
-o1npy yo widuouds ¢ é G1 é A A ‘sid § €1 4 OZOI ISIOP_Y DUDULY) PT
(ga61
(yjoouis |[z) ‘sny pupiyst4 “py addy)
a Gr d ‘sad & €1 - ou gz61 r9uasnYy vy;Ciuvsosnpy
(4ogt
“Sqyy sywpis “py addy)
‘i < a ‘sad & F1 é Logi Ssioquryy sapupdipy
(paaooais (1261
pur yj0ouIs) ‘uag piafiupiquau "yy ad Ay)
ou +41 ii ‘sad bi = ou 1c61 wieyuog spypdiup
(9Sgr savg
IIA-A (yjoours) pypiuoxas sapipyjaqns addy)
‘tao Jeue aed 1 sah 1 ur ‘sad & ‘sad & Pr - ou Q9gI UoAsWI[RP, DY;dUDS!
(yjoours) (Sggr “POY 22qnad +g 9dAq)
‘amo yeue ared 1 ~~ sok GS 5 ‘sad & Fi - soX Gggr ysoyupop sesdoysdiuns:
SH
-jaqvs yo utAuouds ¢ IIA (osoypided (g161
‘avIyouRiIq U99Mj0q GUN. CAN le) ‘sny 1zpuapny ‘gq odA})
oeyided jerprydou « ou II ur ‘sad ‘sid II ou 9161 souasny aasvydupsag
IIIA (SEgr savy
‘oeIyouRIq U99M}0q TILA ‘A (yjoouIs [[2) 40414190390 Djjaqvs ; oacdA}4)
ovyided jerprydou o ou Li ‘9 ur ‘sid & ‘sid II ) - (4) osopided ou gf sprempy—y sapzjaqns'
“A pure AT (bb61
sjuouIsas UO ovI}0S (yjoours {]2) "[ney sisuaipou *gq ody)
ou ynq_ veipodojoN ou 11 a ‘sid II ou Pr61 Aragynery adpwunwwg
*poyIpour ZI JUST IIA (4161
“Sas o10v10y} sno TA ‘A (yiooUurSs [T[e) QISsoFT tuaum 5 ad)
-1a810uN jJoaejasojON] sad II ou ur‘sad & WEGClty th fai ou L161 ossopzy sisdouvsos:
ee 2 O85 85 a“ eres fo Si WAN AS ANT 8 Q
ep a a ee ae 38.8 8. n=
ge ee ey SE Ea 5B
SYIVUWIOY 5" ap 48 a ah 8. a. ovipourig a3 2 a s = sataads-ad Ay
= BP pea & BS be § 3 % jo uoneyag 3 3 pue snuosy
3 4 q “es Spe) EG a:
See ee en 8 7H, BO cs ata
Piemmecat ie s 5
fs) 3 = °
=r B 5 oe
5 5

“panurjuoI—JJ{ AAV],

III

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA)

“xp qouaymnpy
jo wiAuouds ¢ ‘SuOT
trun JO MOI 4SILJ

*pooryd

AyjeqyusA pue SuoyT
Turoun JO MOI 4SsdTJ

*poyIpour ov}0s

puv |esiop 6 juour

-S9S o10e10Yy} sno
-Ig819uN JO 9v19SO}ON)

“wnuojsouny gay

jo utAuouds ¢ *sopoe}
-u9} yNo}s Moy AIDA YW

‘1qd19 Jeue ared 1

ZuL Ce
JO} sjfis jo sured P
soais Q661 joAneqy
‘oeypided
jeue jo opm vy
‘posuo]

-o1d oqo], aepnorjuay,

‘1aa10 yeue ared 1

‘TIA

pue JA s}uoursas
us9MyJoq ISpld [esIOp WV

é al é é

¢ Gi-z1 3 é
OA or Ge = ae aod

ou ZI é Al

é Saou, ou
ou aI é é

ou 1§ é iG

i GE-6e ¢ é

yews 61 ou sak

é

ou

ou

ou

ou

Maj

ou

ou

ou

ou

(yjoours)
a ‘sid 3

(qj00urS)
‘sad &

Cue

(yyoours ¢)
é “sad §&

(umouyun)
é “sid §

IIIA-A_ (q}00UuIS ][Te)
ut ‘sid ‘sad &

(yyoouus ][e)
é x sid

(yoours [[e)

A ‘sad €
(posury

pu yoours)

é ‘sad &

(qyjoours Ile)
é ‘sid §
Sal

‘IIIA

‘ILA ‘A (yoouts TT)

ur ‘sid ‘sad §
IIIA

‘ITA ‘A (yyoouts []@)

ur ‘sad & ‘sad §

od
te

GI

oI

II

GI

ol

oI

oI

yjoouws

yjoouls

yioours

yoows

yjoouls

qoous

qyoours

qoours

yjoous

asoyided

asoypided

ou

ou

ou

ou

ou

ou

ou

ou

ou

ou

ou

(1261
‘uog 2uosjau “Py adA})
1Z61 weyuog saplouurpapy

(4981 ‘Ty vyuis9 “yy ody)
Lggi siapyq xvjgouayany

(SS61

“WA wuaquyom “py eddy)
CG6I uoselly vasnpy

(SS61

‘ITW Snssopso.vut "Fy addy)
CG61 uoseITY sapisvyztiup

(8481 [°ULL

suaosayjog vyjtuvg adA,)

6991
wnuojsouvygqy

(g£61

peane.q 2jauva addis¢T adAq)
GC61 uoserpy adqistjaq

(Sggt

“PW vafisvgd -q adh)
Cggi ysoIUp vYy;duvsnzy

(€Ler

“IQA vDyvsuoja *5 3dAq)
ELQr [MIVA DppayzCiuns

(6161

“wey snuaposap *g addy)
6161 ulproquieyy syqvg

UOSUIAD'T

(S161 “Ty
snjosuoja sapryjaqvs addy)
LIGI ISsoFy sapzppaqvsoane
(4€61 suouuy voa “pyr adAy)
LE6r
vAOyUoUUY ajaivygiupuur/ayy

T12

ANNALS OF THE SOUTH AFRICAN MUSEUM

Key to genera of Ampharetinae

I.

10.

18} fe

12.

Te

14.

15.

16.

17:

4 pairs of gills
3 pairs of gills
2 pairs of gills; first row on uncini ran fone

Glandular ridges on prostomium. Buccal tentacles always smooth and
grooved along one side aa

No glandular ridges on prostomium. Raced enacts eer sino or
papillose

Notopodial cirri present
No notopodial cirri

14 uncigerous thoracic segments . .
II uncigerous thoracic segments ..

14 uncigerous thoracic segments
13 uncigerous thoracic segments
II uncigerous thoracic segments

14 uncigerous thoracic segments
13 uncigerous thoracic segments
12 uncigerous thoracic segments
II uncigerous thoracic segments

Notopodial cirri present
Notopodial cirri absent

Tentacles papillose. Abdominal notopodia seldom present
Tentacles smooth and grooved. Abdominal notopodia always ariesomn!

No nephridial papilla on the branchial ridge
One nephridial papilla on the branchial ridge

No specialized posterior notosetae or elevated notopodia. Over 30
abdominal segments ..

Specialized notosetae present in some ley ated posterior samen.
Less than 20 abdominal segments

One nephridial papilla on the branchial ridge ..
No nephridial papilla on the branchial ridge

Tentacles smooth with a longitudinal groove. No nephridial papillae on
the branchial ridge
Tentacles papillose. 2 nephridial aap on the peach tile:

14. uncigerous thoracic segments .

13 uncigerous thoracic segments ..

12 uncigerous thoracic segments ..

II uncigerous thoracic segments ..

g uncigerous thoracic segments. Last neraeadiaes plead andl fea
modified notosetae

Glandular ridges on prostomium
No glandular ridges on prostomium

Abdominal notopodia present. Tentacles numerous and smooth. Paleae
usually absent : Be at Me 25 oe dc

Abdominal notopodia absent. Tentacles represented by a folded
membrane. Paleae usually absent sf =

(With paleae, other characters unknown)

Tentacles papillose

Tentacles smooth 35 Se Be ae

A dorsal ridge between Bees Vi and VII. Paleae present on segment
Ill vs

No dorsal ridge. No : setae on sgEune | IIT

Tentacular lobe prolonged
No elongate tentacular lobe

2

13
Auchenoplax

art DD ow

Amphicteis
Amage
Parphypania
Aypania
Grubianella

7
Lysippe
8
12
Phyllamphicteis
Lysippides
9
10

Ampharete
Asabellides

Phyllocomus

II

Sosane
Sosanopsis

Paramage

Sabellides

14
Microsamytha

1600
Glyphanostomum

Mugga
Samythopsis
15

Samytha

Amythas
Aryandes

17
18

Melinnampharete
Neosabellides

Pabits
Samythella

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 113

AMPHICTEIS Grube 1851

Synonyms Crossostoma Gosse 1855, ?Rytocephalus Quatrefages 1865, and Paramphicteis Caullery

1944.

Prostomium with a pair of glandular ridges. Buccal tentacles smooth with
a groove along one side. Four pairs of gills. Segments III-VI without neuro-
setae. Segment ITI often with paleae; segments IV—VI with notopodial capil-
laries. Fourteen uncigerous thoracic segments. Notopodial cirri present.
Thoracic uncini with a single vertical series of teeth. Thirteen to 19 abdominal
segments sometimes with rudimentary notopodia as well as uncigerous pinnules.

Type-species Amphitrite gunnert Sars 1835.

AMAGE Malmgren 1866

Prostomium with glandular ridges. Buccal tentacles smooth with a groove
along one side. Four pairs of gills. Segments III-VI without neurosetae.
Segments IV—VI usually with notopodial capillaries. Eleven uncigerous
thoracic segments. Notopodial cirri present. ‘Thoracic uncini with 1 or 2 series
of teeth. Eight to 15 abdominal segments usually with rudimentary notopodia
as well as uncigerous pinnules.

Type-species Amage auriculata Malmgren 1866.

PARHYPANIA Annenkova 1928

Prostomium with glandular ridges. Buccal tentacles smooth. Four pairs
of gills. Segments III—VI without neurosetae. Segment ITI often with paleae,
segments IV—VI usually with notopodial capillaries. Fourteen uncigerous
thoracic segments. Notopodial cirri absent. About 16-20 abdominal segments
sometimes with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Amphicteis brevispinus Grube 1860.

HYPANIA Ostrooumovy 1897

Prostomium with glandular ridges. Buccal tentacles smooth. Four pairs
of gills. Segments III-VI without neurosetae. Segment III often with paleae,
segment IV may have notosetae, segments V and VI always with notopodial
capillaries. Thirteen uncigerous thoracic segments. Notopodial cirri absent.
About 15-30 abdominal segments sometimes with rudimentary notopodia as
well as uncigerous pinnules.

Type-species: Amphicteis invalida Grube 1860.

GRUBIANELLA McIntosh 1885

Prostomium with glandular ridges. Buccal tentacles smooth. Four pairs
of gills. Segments III—VI without neurosetae. Notopodial capillaries present
on segments V and VI and sometimes on segment IV. Eleven uncigerous
thoracic segments. Thoracic uncini usually with 2 series of teeth. Notopodial
cirri doubtful. About 25 abdominal segments; the last few are reported to be
swollen in the type but this is probably an abnormality.

Type-species: Grubianella antarctica McIntosh 1885.

IIi4 ANNALS OF THE SOUTH AFRICAN MUSEUM

LYSIPPIDES Hessle 1917
Synonyms Amphisamytha Hessle 1917 and Hypaniola Annenkova 1928.

Prostomium without glandular ridges. Buccal tentacles smooth with a
groove along one side. Four pairs of gills. Segments III-VI without neurosetae.
Notosetae present on segments V and VI and usually III and IV as well.
Fourteen uncigerous thoracic segments. Notopodial cirri absent. Thoracic
uncini with 1-3 series of teeth. Between 8 and 23 abdominal segments usually
with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Amphicteis fragilis Wollebaek 1912.

PHYLLAMPHICTEIS Augener 1918

Synonym Paiwa Chamberlin 1919.

Prostomium without glandular ridges. Buccal tentacles smooth. Four pairs
of gills. Segments III-VI without neurosetae. Notosetae present on segments V
and VI and usually on III and IV as well. Fourteen uncigerous thoracic
segments. Notopodial cirri present. Thoracic uncini with 1-3 series of teeth.
Between 15 and 20 abdominal segments sometimes with rudimentary notopodia
as well as uncigerous pinnules.

Type-species: Phyllamphicteis collaribranchis Augener 1918.

LysIPPE Malmgren 1866
Synonym Prterolysippe Augener 1918.

Prostomium without glandular ridges. Buccal tentacles smooth. Four
pairs of gills. Segments III-VI without neurosetae. Small notosetae often
present on segments III and IV and notosetae always present on V and VI.
Thirteen uncigerous thoracic segments. Thoracic uncini with 2-3 series of
teeth. Notopodial cirri absent. About 12-15 abdominal segments sometimes
with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Lysippe labiata Malmgren 1866.

AMPHARETE Malmgren 1866
Synonym Branchiosabella Claparéde 1863.

Prostomium without glandular ridges. Buccal tentacles papillose. Four
pairs of gills. No nephridial papilla on the branchial ridge. Segments III-VI
without neurosetae. 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 2 series of teeth. About 12-15 abdominal segments
which have uncigerous pinnules but usually lack rudimentary notopodia.

Type-species: Amphicteis acutifrons Grube 1860.

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 115

ASABELLIDES Annenkova 1929
Synonym Pseudosabellides Berkely and Berkely 1943.

Prostomium without glandular ridges. Buccal tentacles papillose. Four
pairs of gills. One nephridial papilla on the branchial ridge between the two
groups of gills. Segments III—VI without neurosetae. Notopodial capillaries
present on segments V and VI but usually absent from III and IV. Twelve
uncigerous thoracic segments. Notopodial cirri absent. Thoracic uncini usually
with 2 series of teeth. About 21 abdominal segments which have uncigerous
pinnules but lack rudimentary notopodia.

Type-species: Sabellides sibirica Wiren 1883.

PHYLLOCOMUs Grube 1878
Synonym Schistocomus Chamberlin 1919.

Prostomium without glandular ridges. Buccal tentacles smooth with a
groove along one side. Four pairs of gills. Segments III-VI without neurosetae.
Notopodial capillaries present on segments IV, V and VI. Twelve uncigerous
thoracic segments. No specialized notosetae. Notopodial cirri present or absent.
Thoracic uncini with a single series of teeth. Numerous (over 30) abdominal

segments. Type-species: Phyllocomus crocea Grube 1878.

SOSANE Malmgren 1866
Synonym Anobothrus Levinsen 1883.

Prostomium without glandular ridges. Buccal tentacles smooth. Four
pairs of gills. One nephridial papilla on the branchial ridge between the two
groups of gills. Segments III—VI without neurosetae. Notosetae usually present
on all four segments III to VI and often enlarged to form paleae on segment ITI.
Twelve uncigerous thoracic segments with one or more of the posterior noto-
podia elevated and bearing specialized notosetae. Notopodial cirri absent.
Thoracic uncini with 1—2 series of teeth. About 13 abdominal segments usually
with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Sosane sulcata Malmgren 1866.

sosANopsis Hessle 1917

Prostomium without glandular ridges. Buccal tentacles smooth. Four
pairs of gills. No nephridial papillae on the branchial ridge. Segments III-VI
without neurosetae. Segments V and VI with notopodial capillaries and usually
segment IV as well. Twelve uncigerous thoracic segments with the last noto-
podium elevated and bearing specialized notosetae. Notopodial cirri absent.
Thoracic uncini with 2-3 series of teeth. About 11 abdominal segments usually
with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Sosanopsis wirent Hessle 1917.

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

PARAMAGE Caullery 1944

Prostomium without glandular ridges. Buccal tentacles smooth. Four pairs
of gills. No nephridial papillae on the branchial ridge. Segments III-VI without
neurosetae. Segment VI with notopodial capillaries but notosetae often absent
from more anterior segments. Eleven uncigerous thoracic segments. No
specialized posterior notosetae. Notopodial cirri absent. Thoracic uncini usually
with a single series of teeth. About 11 abdominal segments with rudimentary
notopodia as well as uncigerous pinnules.

Type-species: Paramage madurensis Caullery 1944.

SABELLIDES Milne-Edwards 1838

Synonyms Heterobranchus Wagner 1885 and Pterampharete Augener 1918.

Prostomium without glandular ridges. Buccal tentacles papillose. Four
pairs of gills. A pair of nephridial papillae on the branchial ridge between the _
two groups of gills. Segments ITI-VI without neurosetae. Segments V and VI
with notopodial capillaries, segment IV usually fused to segment III and
without notosetae, but notosetae often present on segment III. Eleven uncig-
erous thoracic segments. No specialized posterior notosetae. Notopodial cirri
absent. Thoracic uncini with 1 or 2 series of teeth. Between 11 and 18 abdominal
segments with uncigerous pinnules but no rudimentary notopodia.

Type-species: ? Sabella octocirrata Sars 1835.

sAMYTHOPSIS McIntosh 1885

Prostomium with glandular ridges. Buccal tentacles smooth. Three pairs
of gills. Segments III-VI without neurosetae. Segments I[V—VI with notopodial
capillaries. Fourteen uncigerous thoracic segments. Notopodial cirri doubtful.
About 22 abdominal segments which may have rudimentary notopodia as well
as uncigerous pinnules.

Type-species: Samythopsis grubet McIntosh 1885.

SAMYTHA Malmgren 1866

Prostomium without glandular ridges. Buccal tentacles numerous and
smooth. Three pairs of gills. Segments II-VI without neurosetae. Segments
IV—VI with notopodial capillaries. Fourteen uncigerous thoracic segments. No
notopodial cirri. Thoracic uncini with 2—3 series of teeth. About 13 abdominal
segments with rudimentary notopodia as well as uncigerous pinnules.

Type-species: Sabellides sexcirrata Sars 1856.

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 117

AMYTHAS Benham 1921

Prostomium without glandular ridges. Buccal tentacles replaced by a
frilly membrane. Three pairs of gills. Segments III-VI without neurosetae.
Segments IV—VI with notopodial capillaries. Fourteen uncigerous thoracic
segments. No notopodial cirri. Thoracic uncini with 2 series of teeth. Fourteen
or more abdominal segments without rudimentary notopodia above the
uncigerous pinnules.

Type-species: Amythas membranifera Benham 1921.

ARYANDES Kinberg 1867

A questionable and incompletely described genus generally similar to
Samytha but with paleae.
Type-species: Aryandes gracilis Kinberg 1867.

MICROSAMYTHA Augener 1928
Synonym ? Alkmaria Horst 1920.

Note: Both genera are incompletely described and may belong to the family
Terebellidae. More is known of Microsamytha and this name is retained pro tem.

Prostomium without glandular ridges and ‘of the Ampharete type’. Buccal
tentacles smooth. Three pairs of gills. Segments III-VI without neurosetae.
Segments IV, V and VI with notopodial capillaries. Thirteen uncigerous
thoracic segments. Between 13 and 19 abdominal segments.

Type-species Microsamytha rychiana Augener 1928.

MELINNAMPHARETE Annenkova 1937

Prostomium without glandular ridges. Buccal tentacles papillose. Three
pairs of gills. Segments III-VI without neurosetae. Notosetae usually present
on all four segments (III-V1) and often enlarged to form paleae on segment III.
A dorsal ridge between segments VI and VII. Twelve uncigerous thoracic
segments. No notopodial cirri. ? number of abdominal segments.

Type-species: Melinnampharete eoa Annenkova 1937.

Neosabellides Hessle 1917

Prostomium without glandular ridges. Buccal tentacles papillose. Three
pairs of gills. Segments III-VI without neurosetae. Notosetae present on
segments V and VI but usually absent from III and IV. No dorsal ridge
between segments VI and VII. Twelve uncigerous thoracic segments. No
notopodial cirri. Thoracic uncini usually with 2 series of teeth. About 19
abdominal segments.

Type-species: Sabellides elongatus Ehlers 1913.

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

PABITS Chamberlin 1919

Prostomium without glandular ridges. Buccal tentacles smooth and borne
on a long probosciform organ. Three pairs of gills. Segments III-VI without
neurosetae. Notosetae present on segments V and VI and usually on IV as well.
Twelve uncigerous thoracic segments. No notopodial cirri. Thoracic uncini
with 2 series of teeth. Number of abdominal segments unknown.

Type-species: Pabits deroderus Chamberlin 1919.

SAMYTHELLA Verrill 1873
Synonyms Eusamytha McIntosh 1885 and ? Ecelysippe Eliason 1955.

Prostomium without glandular ridges. Buccal tentacles smooth and not
borne on an elongate tentacular lobe. Three pairs of gills. Segments III-VI
without neurosetae. Notopodial capillaries present on segments IV—VI and
sometimes on III as well. Twelve uncigerous thoracic segments. Notopodial
cirri may be present on the last few thoracic setigers. Thoracic uncini with 1 or
2 series of teeth. Up to 36 abdominal segments.

Type-species: Samythella elongata Verrill 1873.

GLYPHANOSTOMUM Levinsen. 1883
Synonym Amythasides Eliason 1955.

Prostomium without glandular ridges. Buccal tentacles smooth (? always).
Three pairs of gills. Segments III-VI without neurosetae. Segments IV—VI
with notopodial capillaries and segment III sometimes with paleae. Eleven
uncigerous thoracic segments. Notopodial cirri absent. Thoracic uncini with
2 or more series of teeth. Twelve to 25 abdominal segments without rudimentary
notopodia but with uncigerous pinnules.

Type-species Samytha pallescens Theel 1878.

MUGGA Eliason 1955

Prostomium without glandular ridges. Buccal tentacles smooth. Three pairs
of gills. Segments [II-VI without neurosetae but all of them may have notosetae.
Nine uncigerous thoracic segments. Notopodium of the last thoracic segment
dorsally situated and bearing modified notosetae. Thoracic uncini with more
than 2 series of teeth. Number of abdominal segments unknown.

Type-species: Mugga wahrbergi Eliason 1955.

AUCHENOPLAX Ehlers 1887

Synonym Melinnoides Benham 1921.

Prostomium without glandular ridges. Buccal tentacles smooth with a
groove along one side. Two pairs of gills. Segments III-VI without neurosetae.
Segments V and VI with notopodial capillaries. Twelve uncigerous thoracic
segments and about 12-15 abdominal ones. First row of thoracic uncini on
segment VII very long. Thoracic uncini with 1-2 series of teeth. Notopodial

cirri absent. Type-species: Auchenoplax crinita Ehlers 1887.

A REVIEW OF THE FAMILY AMPHARETIDAE (POLYCHAETA) 119

SUMMARY

The characters of existing genera are discussed and it is shown that
confusion has arisen because the anterior segments have been telescoped and
there are two different systems of numbering them. It is proposed that the
separation of genera be based primarily on the number of gills and the number
of uncigerous thoracic segments. The main diagnostic characters of 49 type-
species have been tabulated and it is suggested that 16 genera which are mostly
monotypic be sunk. Keys and definitions of the remaining 33 genera are
provided.

REFERENCES

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INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,
including TaBLE OF CONTENTS and SumMaARy. Position of text-figures and tables must be
indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 43 in. X 7 in. (7$ in.
including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in
alphabetical order of authors’ names (Harvard system). References at end of paper must be
given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,
not ‘and’. Year of publication; several papers by the same author in one year designated by
suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper
names (except in German). Title of journal, abbreviated according to World list of scientific
periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B).
Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (black type).
Part number, only if separate parts of one volume are independently numbered. Page numbers,
first and last, preceded by a colon (without prefix ‘p’). Thus:

Smitu, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;
date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,
with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of
author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of
author of book; his initials; title of book, underlined; edition, if any; volume number, if any,
in arabic numerals, with wavy underlining; pagination of paper; place of publication; name
of publisher. Thus:

SmirH, C. D. 1954. South African plonias. Jn Brown, X. Y. Marine faunas. 2nd ed. 3: 63-95.
London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species
has been previously designated (subsequent to 1758) are listed in chronological order, with
abbreviated bibliographic references to descriptions or citations following in chronological
order after each name. Full references must be given at the end of the paper. Articles and
recommendations of the International code of zoological nomenclature adopted by the XV International
congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:
Euplonia capensis (Smith) Brown, 1955: 259.
When misidentified as another species:
Plonia natalensis (non West), Jones, 1956: 18.
When another species has been called by the same name:
[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).

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